Journal of Biogeography (J. Biogeogr.

) (2012) 39, 22402252

SPECIAL
ISSUE

Connecting dynamic vegetation models

to data an inverse perspective
Florian Hartig1*, James Dyke2, Thomas Hickler3, Steven I. Higgins4,
Robert B. OHara3, Simon Scheiter3 and Andreas Huth1

UFZ Helmholtz Centre for Environmental

Research, 04318 Leipzig, Germany, 2Institute
for Complex Systems Simulation, University of
Southampton, Southampton SO17 1BJ, UK,
3
Biodiversity and Climate Research Centre
(LOEWE BiK-F), 60325 Frankfurt am Main,
Germany, 4Institut fur Physische Geographie,
Goethe Universitat Frankfurt am Main, 60438
Frankfurt am Main, Germany

*Correspondence: Florian Hartig, UFZ

Helmholtz Centre for Environmental Research,
Permoserstr. 15, 04318 Leipzig, Germany.
E-mail: florian.hartig@ufz.de

ABSTRACT

Dynamic vegetation models provide process-based explanations of the dynamics

and the distribution of plant ecosystems. They offer significant advantages over
static, correlative modelling approaches, particularly for ecosystems that are
outside their equilibrium due to global change or climate change. A persistent
problem, however, is their parameterization. Parameters and processes of
dynamic vegetation models (DVMs) are traditionally determined independently
of the model, while model outputs are compared to empirical data for validation
and informal model comparison only. But field data for such independent estimates of parameters and processes are often difficult to obtain, and the desire to
include better descriptions of processes such as biotic interactions, dispersal,
phenotypic plasticity and evolution in future vegetation models aggravates limitations related to the current parameterization paradigm. In this paper, we
discuss the use of Bayesian methods to bridge this gap. We explain how Bayesian
methods allow direct estimates of parameters and processes, encoded in prior
distributions, to be combined with inverse estimates, encoded in likelihood
functions. The combination of direct and inverse estimation of parameters and
processes allows a much wider range of vegetation data to be used simultaneously,
including vegetation inventories, species traits, species distributions, remote
sensing, eddy flux measurements and palaeorecords. The possible reduction of
uncertainty regarding structure, parameters and predictions of DVMs may not
only foster scientific progress, but will also increase the relevance of these models
for policy advice.
Keywords
Bayesian statistics, calibration, data assimilation, forest models, inverse modelling, model selection, parameterization, plant functional types, predictive
uncertainty, process-based models.

INTRODUCTION

Correlative versus dynamic vegetation models

Plants are essential for life on Earth. Not only is their

photosynthetic production the primary energy source for the
biosphere, plants also influence the main global biogeochemical cycles, structure terrestrial habitats, and interact with other
important compartments of the Earth such as soil and
atmosphere. Moreover, the global climate is substantially
influenced by vegetation through the exchange and accumulation of carbon, and through effects on the Earths albedo,
water balance and evapotranspiration (Bonan, 2008). Understanding and predicting how climate and other factors such as
soil, water and nutrients influence vegetation dynamics and
species distributions is therefore a key question in ecology
(Grace, 2004).

Current approaches to modelling plant species, communities

and biomes, particularly in response to climate change, broadly
fall into two categories: correlative models on the one hand
and dynamic vegetation models (DVMs) on the other (Pereira
et al., 2010; Dormann et al., 2012).
Correlative modelling approaches, also referred to as
climatic envelope models, species distribution models (SDMs)
or niche models, use statistical methods to relate environmental variables such as climate or soil conditions to the presence
or absence of a species (Elith & Leathwick, 2009). An
important factor for the success of these models is the way
they make use of empirical data: in correlative species
distribution models, parameters and model structure are

2240

http://wileyonlinelibrary.com/journal/jbi
doi:10.1111/j.1365-2699.2012.02745.x

2012 Blackwell Publishing Ltd

Dynamic vegetation models an inverse perspective

inferred inversely by comparing model predictions with
observed species distribution patterns. It is widely acknowledged that the resulting phenomenological descriptions of the
species niche must be interpreted with care (Araujo & Guisan,
2006; Elith & Leathwick, 2009), but the ability of these models
to rapidly convert distribution data into predictive models for
a large range of taxa has made them a widely used tool in
ecology and conservation biology (Zimmermann et al., 2010).
Dynamic vegetation models, on the other hand, provide a
bottom-up description of plant communities by explicitly
modelling physiological and population level processes such as
growth, photosynthesis, carbon allocation, regeneration and
mortality. For many questions that arise in the context of
environmental change, they have considerable advantages
compared to static correlative models. For example, dynamic
models do not need to assume that the vegetation is in
equilibrium with the environment. Thus, they may be used to
model succession, impacts of disturbances, and transient
vegetation dynamics that result from changes in environmental
and climatic conditions (Shugart, 1998; Prentice et al., 2007;
Hickler et al., 2012), as well as reactions to biotic changes or
invasions (Gritti et al., 2006).
Dynamic vegetation models, however, are also associated
with a major practical disadvantage: the model structure and
model parameters are traditionally derived from independent
experiments or empirical observations. We call this a direct
parameterization, as opposed to the inverse parameterization
approach used with correlative models. A direct parameterization is certainly appealing because it derives model outputs
from basic processes, and allows the validity of these processes
to be tested with independent data. However, the necessity for
direct observations of all parameters also severely limits
complexity in terms of processes, diversity and locations that
can be modelled. Process-based models are therefore often
characterized as being data-hungry (Guisan & Thuiller, 2005).
The best of both worlds combining direct and
inverse data sources with process-based models
The term data-hungry suggests that parameterizing DVMs
requires large amounts of data. This may be the case, but the
more pressing problem is the type of data required. Direct
parameterizations require all model parameters (e.g. on growth
or physiology) to be determined by direct measurements or
observations, which are often difficult or laborious to obtain.
On the other hand, many existing data, such as species
distribution databases, eddy flux data or palaeorecords, cannot
be used because they describe variables that are modelled on
the basis of multiple, interacting processes and therefore
depend on many parameters at the same time.
Inverse modelling is the key to using these data. As with
correlative models, inverse modelling methods adjust model
parameters and processes according to their ability to reproduce field observations. Moreover, Bayesian methods offer the
possibility of systematically combining information gained by
this inverse modelling process with other independent inforJournal of Biogeography 39, 22402252
2012 Blackwell Publishing Ltd

mation. There are already some successful applications of these

techniques (e.g. Van Oijen et al., 2005), and we anticipate that
their wider application, particularly on more complex models,
is only a matter of time and computing power. In particular,
we believe that inverse modelling will improve parameterizations of processes such as local interactions, dispersal, migration and plant physiology, enable better representations of
functional and species diversity, and allow accommodation of
additional processes such as phenotypic plasticity and evolution. Inverse modelling may therefore be a crucial ingredient in
making DVMs a viable alternative to correlative species
distribution models for many questions of global change and
conservation.
The aim of this paper is to discuss the use of inverse
modelling with Bayesian statistics as a means to combine direct
and inverse parameterization paradigms for model selection
and parameter estimation. In the following, we first discuss the
properties of DVMs and the types of predictions they make.
Then, we explain the general principles of inferring parameters
and underlying processes by inverse modelling, and discuss
computational and methodological challenges. Finally, we
discuss data types and sources that we consider promising for
constraining parameters and underlying processes of DVMs,
and give some specific hints for including these data in an
inverse modelling process.
DYNAMIC VEGETATION MODELS
We understand a DVM to be any process-based model that
predicts the population dynamics of plant species or plant
functional types over time. A large number of different model
types fall into this category, from dynamic global vegetation
models (DGVMs; see Prentice et al., 2007) to more local
approaches such as forest gap models (see Bugmann, 2001).
The conceptual differences between these model approaches,
however, will not be the focus of this paper (see e.g. Bugmann,
2001; Prentice et al., 2007; Jeltsch et al., 2008).
A central component of most DVMs is the description of a
plants response to its environmental conditions (which may
be influenced by competition with other plants, see below) in
terms of processes such as resource uptake, photosynthesis,
respiration, resource allocation, mortality and reproduction
(Fig. 1). This description may be based on differential or
difference equations, and can be provided at different levels of
aggregation, ranging from fully individual-based representations over age classes up to population-level descriptions
(Smith et al., 2001; Prentice et al., 2007). When sufficient data
are available, all species of interest can be modelled explicitly.
Forest models, for example, often include parameterizations
for 530 species (e.g. Bugmann, 1996; ForClim model; Pacala
et al., 1996; SORTIE model; Xiaodong & Shugart, 2005;
FAREAST model; Hickler et al., 2012; LPJ-GUESS model).
For global models and for local models in highly diverse
ecosystems such as grasslands, savannas and tropical forests,
species are typically grouped into plant functional types (PFTs)
that represent a number of species with similar ecological
2241

F. Hartig et al.

Individual scale

Plot scale

Geographical scale

Projection into
geographic space

Carbon
allocation

Seeds

Upscaling

species1

Factor 2

Factor 2

Latitude

species1

species 2

species1

species 2

Factor 1

Factor 1

Fundamental niche,
growth rate > 0
in isolation

Realized niche,
growth rate > 0
in community

Longitude
Realized distribution,
map of realized niche to
environmental conditions

properties (Kohler et al., 2000; see also Huth & Ditzer, 2000;
FORMIX model; Sitch et al., 2003; LPJ model; Scheiter &
Higgins, 2009; aDGVM model). It should be noted, however,
that the number of PFTs that are locally co-occurring in global
vegetation models is typically much lower than in their more
local counterparts (e.g. Huth & Ditzer, 2000).
Central to most DVMs are also interactions between plants,
for example through competition for resources such as light,
water and nutrients that are available in a common resource
pool (Fig. 1). These competition processes are major determinants for predicted community composition, non-equilibrium
dynamics and succession. Examples are the dry limit of trees,
which is determined in many DGVMs by competition with
grasses and fire (Sitch et al., 2003; Scheiter & Higgins, 2009),
and successional dynamics in forest gap models, which are
based on vertical light competition between trees of different
sizes (Bugmann, 1996; Huth & Ditzer, 2000). Finally, geographical distribution and productivity maps can be created by
upscaling model predictions to the scale at which environmental information is available (Fig. 1).
Thus, DVMs take a bottom-up approach to predict from
basic processes the responses of plants to environmental
conditions and competition. There are, however, limitations
to constraining all necessary parameters and processes bottomup. For example, despite the fact that many processes in DGVMs
are relatively well understood, small differences in parameterization and model design can propagate to create large
uncertainties (Cramer et al., 2001; Sitch et al., 2008; Galbraith
et al., 2010). These uncertainties are unlikely to have been
reduced by the newer generation of DVMs (Moorcroft et al.,
2242

species 2

Figure 1 Structure of dynamic vegetation

models and the connection to the niche. At
the core of the model are the properties of the
individual species (physiological,
morphological, phenological, life history and
bioclimatic parameters), which determine the
range of environmental factors at which an
individual plant is able to maintain positive
growth at low population density
(corresponding to its fundamental niche).
Species interactions determine the actual
community composition under given
environmental conditions (corresponding to
its realized niche). Finally, the realized niche
may be mapped into geographical space by
connecting a spatial map of environmental
conditions with the realized niches of the
species.

2001; Smith et al., 2001; Sato et al., 2007). Moreover, the desire
to include additional processes, such as dispersal and migration
(Lischke et al., 2006), more detailed plant physiology (Higgins
et al., 2012), seed competition (Bohn et al., 2011), environmental modulation (Linder et al., 2012) or phenotypic plasticity
and evolution (e.g. Parmesan, 2006; Kramer et al., 2010), is
bound to add to the problem of constraining uncertainty about
parameters and processes. A further issue in global vegetation
models is the practice of summarizing a large number of species
or individuals by an average functional type. The Amazonian
Basin, for example, is home to c. 11,000 tree species (Hubbell
et al., 2008), but many global vegetation models summarize this
diversity by only two functional types. While we concede that it
may not always be necessary to model each species explicitly,
there is substantial evidence that interspecific and intraspecific
diversity in species traits may be important for understanding
and correctly predicting community dynamics (Daz & Cabido,
2001; Clark, 2010; McMahon et al., 2011).
To address these challenges, we have to find additional
information to constrain parameters and processes. To that
end, it seems natural to explore the potential of inverse
modelling approaches, which allow for the use of a large range
of data types from different sources and scales that cannot be
used for direct parameter estimation. Modern Bayesian
methods provide the means to combine such inverse parameter estimates with any direct parameter estimates that are
available, and thus provide a method to synthesize all available
data sources. In the following section we review the general
technical prerequisites of Bayesian methods for parameter
estimation and model selection. This will be useful for
Journal of Biogeography 39, 22402252
2012 Blackwell Publishing Ltd

Dynamic vegetation models an inverse perspective

understanding our subsequent discussion of promising data
types and their properties.
INFERRING PARAMETERS AND PROCESSES
BY INVERSE MODELLING
Let us assume that we have some empirical observations D that
can be reproduced with a DVM, and that we want to compare
several versions of this model that may differ in the choice of
model parameters (parameter estimation) or in the choice of
processes (model selection) in their ability to reproduce these
data.

(a) Normal error model

Model prediction M() Observation Dobs

Likelihood: p(D obs|M())~
exp[-1/(22)(M()-Dobs)2]
e.g. Biomass
(b) Bernoulli error model

Dobs = n observations, k presence

The likelihood as the goodness of fit
Clearly, a core question for any inverse methodology is to
define a measure that quantifies how well the predictions of
any particular model or parameterization fit to our observations. Different approaches have been advocated. A common
method is to define ad hoc some objective function that
measures goodness of fit, that is, the difference between
predicted and observed data (see e.g. Schroder & Seppelt, 2006).
Usually, this function is subsequently used to find the best fitting
parameter set by optimization. Although useful in some
situations, a disadvantage of the ad hoc choice of objective
functions is their lack of statistical interpretability, which
prevents a clear definition of confidence boundaries and creates
problems when multiple data sources have to be merged into one
single objective function (see Hartig et al., 2011).
An alternative is provided by the likelihood, a statistical
measure of the quality of the fit (e.g. Gelman et al., 2003).
Given that we want to compare different parameterizations Q
of a model in terms of their ability to reproduce some
empirical data D, the likelihood p(D|Q) is defined as the
probability of obtaining the observations D given the model
parameterized with Q. One may ask why there should be a
probability of obtaining the data, given that even stochastic
vegetation models often produce very stable outputs and
therefore either reproduce the data or not. An analogy that
some readers may find useful is that of a nonlinear regression: in
such a regression, the aim is to fit a nonlinear function (which
could also be a DVM) that relates a dependent variable (in our
case the structure or dynamics of the vegetation) to an
independent variable (the environment). To this end, a regression uses a probabilistic model (e.g. that observations are
distributed around the true value according to a normal
distribution) to represent additional processes that lead to
unexplained variation in the data (observation error, unobserved covariates). This error model allows a probability p(D|Q)
(called the likelihood) to be assigned to the observed deviations
between observation and model prediction (residuals).
Thus, the likelihood is given by the probability of obtaining
the observed deviations between model predictions and data,
based on an error model that needs to be defined. This error
model can have a large effect on the results of the inference. It
must therefore be chosen with care to reflect our knowledge
Journal of Biogeography 39, 22402252
2012 Blackwell Publishing Ltd

M() = pred. occurrence probability

Likelihood:
p(Dobs |M()) ~ k (1-)(n-k)

Figure 2 Examples of calculating the likelihood of observing Dobs

given the model predictions M(F): (a) a likelihood based on a
normal observation error. (b) A Bernoulli process is one of the
simplest error models for presence/absence observations, given
that the model predicts occurrence probabilities p.

about the processes that lead to variation in system states

(process error) and observations (observation error), as well as
with the actual deviations between model predictions and
observed data after the fit (below).
If we assume that the error is given by an independent,
normally distributed random variable (typically an observation
error), the likelihood is proportional to:
pDjH / e

MHD2
2r2

where M(Q) are the model predictions resulting from the

parameterization Q, D the observed data, and r the standard
deviation of the error (see Fig. 2a). Because of its simplicity and
general acceptance, the normal error model is widely used (e.g.
Lasslop et al., 2008 or Higgins et al., 2010), but many other
options are possible (e.g. Fig. 2b, see also Gelman et al., 2003).
When using multiple datasets, likelihoods can often be
combined with relative ease: when deviations between predicted and observed data are uncorrelated, their corresponding
likelihoods represent independent probabilities, and so their
joint likelihood is simply the product of the individual
likelihoods. However, difficulties arise when deviations
between model predictions and data are correlated within or
across data sets. Luckily, we can build on a large stock of
existing knowledge about how to treat these problems from
statistical models of the vegetation. Dormann et al. (2007), for
example, discuss the problem of specifying error models for
spatially autocorrelated data. Previous work on more complex
(hierarchical) error models that include several layers of
dependent stochastic processes, such as environmental
stochasticity, intraspecific variability, process errors and
observation errors is also of interest in this context (Clark,
2243

F. Hartig et al.

Weak / noninformative prior

Parameter value
Bayes theorem
By specifying the likelihood function, we have established a
statistically interpretable measure of fit, which could, for
example, be used to search for the model parameters Q with
the highest likelihood (maximum likelihood estimation).
Often, however, there is additional, independent information
about parameters or processes available, for example from field
data that provide direct parameter estimates. Bayes theorem
offers the possibility of merging such independent information
with the inversely generated information that is contained in
the likelihood. The theorem states that
pHjD

pDjH  pH
pD

p(Q|D) is called the posterior density, and is interpreted as a

probability density that summarizes our information about the
probable values of Q. The posterior p(Q|D) depends on the
likelihood p(D|Q) (see above), and a new term p(Q), which
will be discussed below (see Gelman et al., 2003 and Stephens
et al., 2007 for more details on the Bayesian approach and its
relation to other measures of statistical inference). The
denominator, p(D), is the integral of the numerator over Q,
and acts only as a normalization constant.
Specifying prior information
The new term, p(Q), is called the prior. It quantifies the
uncertainty about the model parameters before comparing
model outputs to empirical data. As an example, if measurements
about a parameter such as the specific leaf area (SLA) of a plant
are available, we may summarize this information by a normal
distribution for this parameter (Fig. 3). Also, if a model was
already parameterized with Bayesian methods, and new information becomes available, this new information can be merged
with the existing information by using the posterior distribution
from the old data as the prior for the new data (Fig. 4).
The possibility of including prior information about a
parameter in the inference is one of the great advantages of
Bayesian statistics. Often, however, there is a desire to avoid
doing so, either because there is no prior knowledge, or to
avoid the criticism of biasing the posterior in a particular
direction (let the data speak). For this purpose, so called
vague, weak, non-informative or reference priors are
used. Intuitively, one might think that the natural choice for
such a prior is the uniform (flat) prior, which associates the
2244

Strong / informative
prior
Probability density

Probability density

2005; Schurr et al., 2012). Likelihood functions may be

difficult to formulate for such complicated error structures
within DVMs, but new methods allow approximate likelihood
functions to be constructed directly from the stochastic model
outputs (Hartig et al., 2011). This allows us to go beyond
fitting process-based models with phenomenological error
models towards an approach where dynamic models explain
both the mean effects and the stochasticity in the data through
mechanistic descriptions of the ecological system and the data
acquisition process (Zurell et al., 2009).

Parameter value

Figure 3 A conceptual illustration of the difference between a

weak (non-informative) prior on the left, where all parameter
values are equally likely, and a strong (informative) prior on the
right.

same probability with all parameter values (Fig. 3). However,

it is well known that this is not generally true because it leads
to logical inconsistencies, for example when considering
nonlinear parameter transformations. As a rule of thumb, flat
priors are a reasonable way of expressing the absence of prior
information for parameters that act linearly and uncorrelated
on the model predictions. In other cases, corrections may be
needed (see Kass & Wasserman, 1996).
Checking model assumptions and posterior
diagnostics
A core assumption of all likelihood-based inference is that
deviations between model predictions and observations originate from a random source of variation that is captured well
by the error model underlying the likelihood function. If there
are structural errors in the model, however, or systematic
measurement errors, these assumptions may not hold. Posterior distributions can then be biased because parameters are
adjusted to compensate for structural or systematic errors.
Detecting such systematic errors in a Bayesian fit, however, is
still helpful because it aids in the identification of problems in
data acquisition and model assumptions.
One diagnostic to check for is the consistency of prior and
posterior values. Typically, one would expect likelihood and
prior distributions to point towards the same range of
parameter values, which will result in a reduction of parameter
uncertainty. However, when there are substantial differences,
posterior uncertainty will remain large, or can even increase
compared to the prior. Although it is possible that such effects
result from pure chance, they may also suggest a structural
problem in the vegetation model, a systematic observation
error in the data, or an error in the specification of the priors.
A second aspect to examine is systematic patterns in the
deviations between model predictions and observed data
(systematic mismatch, correlation, heteroscedasticity) that
are not in accordance with the error model on which the
likelihood was based. Such discrepancies can often be
addressed by correcting the error model. Remaining systematic
Journal of Biogeography 39, 22402252
2012 Blackwell Publishing Ltd

Dynamic vegetation models an inverse perspective

Information independent of the model

(prior information)

Comparison of model outputs and observed data

Measurements

Forest size structure

predicted

Abundance

NPP predicted

e.g. specific
leaf area

Stem diameter

Prior
parameter
estimate
Repeat to
add new
information

Likelihood
Dynamic of model
vegetation predictions
given the
model
observed
Posterior
data
parameter
estimate

Error model

NPP observed

Abundance

Forecasting,
quantification of
predictive uncertainty

Forest size structure

observed

Model selection
and improvement

image source: NASA

Stem diameter

Analysis and applications of the fit

Figure 4 The inverse modelling cycle starts with the specification of prior information (top left). In the next stage, prior probabilities are
multiplied by the likelihood. As a result, we obtain a posterior estimate for parameters or model structure. This posterior estimate may be
used to calculate the predictive uncertainty of the model, to select and improve models, or as an input for a further inverse modelling cycle.
The comparison between model and data shows net primary productivity (NPP) predictions (T. Hickler, unpublished data) from LPJDGVM (Sitch et al., 2003) with hydrology updates by Gerten et al. (2004) as implemented within the generalized ecosystem modelling
framework LPJ-GUESS (Smith et al., 2001) compared with normalized difference vegetation index (NDVI) as proxy for observed NPP from
Terra/MODIS (February 2011), and abundance of trees in different size classes for a tropical mountain forest simulated by the forest model
FORMIND and compared with field data (Dislich et al., 2009).

deviations, however, that cannot be sensibly related to a

stochastic process, hint towards structural problems in the
vegetation model or systematic observation errors in the data.
Predictions from posterior parameter estimates
Apart from analysing posterior parameter estimates or using
them as new priors if new data become available (Fig. 4), the
posterior can also be used to estimate predictive uncertainties.
This is usually done by drawing a number of parameter
combinations from the posterior and calculating the
distribution of the model predictions that arises from these
draws.
Comparisons between structurally different models
The cycle of Bayesian parameter estimation, as summarized in
Fig. 4, can also be used to test or improve our knowledge
about underlying processes (e.g. ecophysiology or demograJournal of Biogeography 39, 22402252
2012 Blackwell Publishing Ltd

phy) by comparing several structurally different models and

seeing which receives more support from the data. Various
approaches have been advocated, and the choice of method
depends on whether the aim is to find one best model (model
selection), or an ensemble of models, weighted by their model
fit (model averaging, e.g. Butler et al., 2009). For an overview
of these approaches, see the reviews by Johnson & Omland
(2004) and Kadane & Lazar (2004).
Remarks on algorithms and computing time
The estimation of posterior densities can usually not be done
analytically. Therefore, Monte Carlo algorithms such as
importance sampling (IS), Markov chain Monte Carlo
(MCMC) and sequential Monte Carlo (SMC) algorithms are
commonly used. These algorithms generate approximate
posterior densities by evaluating a large number of model
runs with different parameterizations (Andrieu et al., 2003;
Hartig et al., 2011). For most computing languages such as R,
2245

F. Hartig et al.
MATLAB or PYTHON, there are packages that provide
implementations for the most common algorithms. Also,
there are stand-alone implementations such as OpenBUGS or
JAGS.
A point of practical importance is the number of model
calculations needed for these algorithms: an inverse Bayesian
parameter estimation typically requires about 5 104 to
5 105 model runs for a model of the order of 20 parameters.
Given that there are limits to the parallelizability particularly of
MCMC algorithms, it follows that a single model run should
not take longer than a minute (a runtime of 1 min for a single
simulation and 105 steps of the algorithm result in a total
runtime of c. 70 days). For models with longer runtimes,
model simplification, model parallelization or model emulation (Conti & OHagan, 2010) can be considered.
SELECTING FIELD DATA FOR INVERSE
MODELLING OF VEGETATION MODELS
The final part of this paper is devoted to data and applications.
We discuss what we perceive as the most promising data types
for constraining parameters and processes of DVMs, and argue
that combining multiple data types at different scales provides
a promising general strategy for obtaining strong, independent
data sets.
Data from local vegetation inventories
One traditional source of vegetation data is local vegetation
inventories. Such observations usually describe the vegetation
on sampling plots of up to a few hectares, listing species,
abundance, sizes or biomass, growth and sometimes also the
spatial location of plant individuals. Direct vegetation observations are comparably abundant for forests [Condit et al.,
2000; Jenkins et al., 2001; ter Steege et al., 2006; see also the
Forest Inventory Analysis program (FIA) and the Center for
Tropical Forest Science (CTFS) forest plot network], but there
are also good datasets available for grasslands (Roscher et al.,
2004), savannas (Sankaran et al., 2005) and from general
databases such as the Global Index of Vegetation-Plot

Databases (see http://www.givd.info/). Additionally, local

observations can be used to generate chronosequences from
stands of different ages (space for time substitution, Pickett,
1989; for an application see Purves et al., 2008). Data are also
accumulating on plant functional traits, for example in the
TRY database (Kattge et al., 2011).
Local vegetation data are particularly useful because they
contain species-specific information about the abundance and
size-structure of the vegetation. Interesting examples of
Bayesian calibrations with local inventory data are provided
by Van Oijen et al. (2005), Higgins et al. (2010), Martnez
et al. (2011) and Weng et al. (2011).
Extending the spatial scale remote sensing and
distribution data
Local vegetation inventories give a detailed picture of the local
vegetation structure, but they are expensive and so cannot be
extended to continental or global scales. Here, remote sensing
of the vegetation and species distribution databases (Fig. 4)
offer complementary information.
Distribution databases are usually generated from presence
or presence/absence observations as well as from expert
knowledge. Examples are GBIF (http://www.gbif.org/) or the
Flora Europaea database (http://www.faunaeur.org/). Distribution data are the main data source for the inverse calibration
of correlative species distribution models, but they are
currently rarely used for the calibration of DVMs (but see
Purves et al., 2007). One reason for this may be that most
large-scale vegetation models are calibrated for functional
types and not for species. In principle, however, these data
might provide valuable information and could be used with
error functions similar to those used for correlative species
distribution models (see Fig. 2b).
Remote sensing data can be subdivided into two main types:
passive sensors that detect reflected light from the vegetation
(optical, infrared and microwave spectrum), and active sensors
that record the backscattering from either a laser (lidar = light
detection and ranging) or a radar signal (see Table 1;
Chambers et al., 2007; Frolking et al., 2009). Remote sensing

Table 1 Overview of remote sensing technologies and their application.

Type

Notes, products, references

Passive

Sensors such as Landsat or MODIS (Moderate Resolution Imaging Spectroradiometer) observe the visible, infrared and
microwave spectrum; allows estimation of leaf area via the normalized difference vegetation index, NDVI
(see, e.g. Zhang et al., 2003), as well as water content and temperature profiles of canopies (infrared spectrum,
Fensholt et al., 2010) and soil moisture and temperature (microwave spectrum, Kerr, 2007) and even chemical and
taxonomic diversity (Asner & Martin, 2009).
A lidar device creates a laser beam and measures the reflection created by all optically reflecting material, i.e. ground,
leaves etc. May be used to estimate vegetation height and leaf area index, LAI (Asner et al., 2010; Lefsky, 2010).
A radar device creates an electromagnetic signal, typically radio waves or microwaves, and measures reflections of
this signal. Depending on the wavelength, reflections are triggered particularly by the massive parts of the vegetation,
i.e. branches and trunks. Used to estimate aboveground biomass, height/vertical structure of the vegetation (Krieger
et al., 2010; Kohler & Huth, 2010), also soil moisture (Kerr, 2007) and temperature.

Laser (Lidar)
Radar

2246

Journal of Biogeography 39, 22402252

2012 Blackwell Publishing Ltd

Dynamic vegetation models an inverse perspective

techniques have undergone substantial progress over the past
few decades, so that global maps and time series of vegetation
types, leaf area and productivity are becoming increasingly
available. For all these reasons, a number of authors have
recently stressed the potential of these data for vegetation
modelling, in particular for forest models (Frolking et al.,
2009; Hurtt et al., 2010; Kohler & Huth, 2010).
Error models for remote sensing data will depend on the
variables and the resolution of the data. One obvious issue is that
remote sensing data can be obtained at very high resolutions
(below a metre), which can result in strong spatial autocorrelation. To avoid problems associated with this, one can consider
upscaling the data to match the spatial scale of the model, and
one should account for remaining strong residual autocorrelation
(Dormann et al., 2007). In some cases, however, information
may be contained in the small-scale variation itself: Kohler &
Huth (2010), for example, suggest that the local variability of
biomass values contains additional information that may be
used to infer the degree of human forest use (logging). Another
interesting example comes from Patenaude et al. (2008) who
fitted a model with a combination of hyperspectral, lidar and
radar measurements, as well as with additional local stand data.
Extending the temporal scale: eddy flux data and
palaeorecords
In addition to the spatial dimension, a second dimension along
which we can extend our empirical baseline is provided by
observations over time-scales that are not covered by the
aforementioned data types. Here, we discuss two particularly
interesting examples, namely eddy flux measurements and
palaeorecords (Fig. 4).
An eddy flux experiment typically consists of recording
wind, temperature and gas concentrations at different heights
above a vegetated surface. From these data, one can estimate
the heat, water, and gas exchange of the vegetation at scales up
to 30 m. The number of eddy flux experiments has increased
significantly in recent years. Most of these are organized in
FLUXNET (Baldocchi et al., 2001), a network of over 500 eddy
flux sited all over the world.
Eddy flux data are particularly useful for estimating
parameters related to processes that act on short time-scales,
such as photosynthesis or evapotranspiration. Like remote
sensing data, eddy flux data can be measured at a very high
resolution (hours or minutes), but it is important to check
whether this resolution fits to the scale of the model that is
used. When errors originate predominantly from processes
that change slowly compared to the measurement frequency,
observations may show strong temporal autocorrelation.
Hence, either the data have to be summarized at a coarser
temporal scale, or the temporal autocorrelation must be
accounted for in the error model. Friend et al. (2007) and
Williams et al. (2009) give an overview of inverse modelling
with FLUXNET data, partly also coupled with remotely sensed
information (see also: Hanson et al., 2004; Braswell et al.,
2005; Knorr & Kattge, 2005; Medvigy et al., 2009).
Journal of Biogeography 39, 22402252
2012 Blackwell Publishing Ltd

Finally, an exciting possibility is to compare vegetation

models to palaeodata. Pollen data, for example, can be used to
reconstruct vegetation composition over long time-scales. At
the same time, global climate reconstructions are available that
may be used as inputs for vegetation models (Allen et al.,
2010). Together, these data might prove invaluable for
inferring processes that are rare, slow or difficult to test, such
as long-distance dispersal abilities of plants, phenotypic
plasticity and rapid evolution or large-scale vegetation
responses to elevated CO2 levels (Miller et al., 2008; Prentice
& Harrison, 2009; McMahon et al., 2011). We stress, however,
that large uncertainties exist due to limited dating accuracy
and spatial resolution, taxonomic uncertainty, unknown
disturbance events, and large differences between species in
pollen productivity (e.g. Sugita et al., 1999). Nevertheless,
palaeodata provide important information on a time-scale
unmatched by other data.
The need to use multiple data types at a time
Which of these many different data types should we use? When
deciding on field data for inverse modelling, we have two aims.
The first aim is to find data that are informative about the
parameters and processes of interest. When a model output is
independent of a subset of the parameters, we cannot expect
field data for this output to be informative about these
parameters. A global sensitivity analysis can be useful for
identifying the relevant parameters for the available data, or
conversely, to identify the data that are needed to fit a
particular parameter subset or submodel.
The second aim relates to the problem of correlation. In
principle, it is always better to have more data, but there is one
caveat: as mentioned before, likelihoods can most easily be
combined when the underlying data are uncorrelated. Combining strongly correlated data requires more complex error

Figure 5 Complementarity of spatial and temporal scales within

the data sources discussed.

2247

F. Hartig et al.
models, which may increase the computational burden and lead
to instabilities in inference if the correlation structure is not
captured adequately. Thus, information gains have to be
weighed up against potential problems that come with new data.
Correlations, however, may not only be influenced by the
types of data that are chosen, but also by the way the data are
aggregated and represented. Data can be transformed, changed
in scale (up/downscaling) or summarized. A sensible choice
here may considerably reduce correlations. Thus, we believe
that independence of data is a technical problem, but we still
recommend considering the full spectrum of vegetation data
for inference. In particular, we recommend including data that
are measured at different scales (Fig. 5) because errors will
tend to be less correlated and information more complementary than for data obtained at the same scale (Grimm &
Railsback, 2012). Using all available data sources will not only
facilitate a synthesis of our empirical information as a product
that is more useful for ecological theory and prediction, it will
also facilitate the further development of DVMs by systematically assessing their ability to match multiple empirical
observations at the same time.
CONCLUSIONS
We have discussed the principles of the Bayesian modelling
approach for inferring parameters and structure of DVMs.
Inverse parameterization of DVMs resembles the correlative
species distribution modelling approach. The importance of
prior knowledge about parameters, and also about model
structure, however, will remain an area where DVMs differ
significantly from correlative modelling approaches. We
therefore think that inverse modelling methods will not, as
one might fear, reduce DVMs to merely a very complicated
version of a correlative model that is blindly adjusted to data.
Rather, Bayesian methods offer a state-of-the-art technique
that allows us to use all available data (on model inputs and
model outputs) to improve our knowledge about the
processes that govern vegetation dynamics, and hence our
ability to predict the future development of the Earths
vegetation.
The additional information that can be gained by applying
inverse modelling methods to heterogeneous data types such as
species distributions, local vegetation inventories, eddy flux
measurements, and remote sensing data offers tremendous
scientific potential (see also Luo et al., 2011). Using many or
all of these data in parallel will allow us to build models that
provide a better representation of the dynamics and functional
diversity of the terrestrial vegetation. Bayesian methods will
also improve our knowledge about the uncertainty of model
predictions, which is an important factor for informing policy.
The benefits of such a route lie not only in the potential to
generate substantially better predictions, making DVMs more
relevant for applied questions, but also in the ability to test
ecological theory with DVMs at different temporal and spatial
scales, which allows fundamental questions of evolution,
biogeography and community ecology to be addressed. For
2248

all these reasons, we anticipate a further growth in the

popularity of Bayesian methods for inferring parameters and
processes of DVMs.
ACKNOWLEDGEMENTS
We would like to thank Christine Romermann, Steve Higgins
and Bob OHara for organizing the workshop from which
this paper originated, as well as the LOEWE initiative (see
below) for funding this event. We are grateful for comments
by Olga Bykova, Anne Carney, Carsten Dormann, Martin
Kazmierczak, Peter Linder, Glenn Marion, Peter Pearman,
Christopher Reyer, Stan Schymanski, Robert Whittaker and
three anonymous referees, which greatly improved this
manuscript. F.H. acknowledges support from ERC advanced
grant 233066. Funding for R.B.OH. and S.S was provided by
the Biodiversity and Climate Research Centre (BiK-F), which
is part of the LOEWE programme Landes-Offensive zur
Entwicklung Wissenschaftlich-okonomischer Exzellenz of
Hesses Ministry of Higher Education, Research and the
Arts. J.D. acknowledges support from the Helmholtz
Association through the research alliance Planetary Evolution
and Life.
REFERENCES
Allen, J.R., Hickler, T., Singarayer, J.S., Sykes, M.T., Valdes,
P.J. & Huntley, B. (2010) Last glacial vegetation of
northern Eurasia. Quaternary Science Reviews, 29, 2604
2618.
Andrieu, C., de Freitas, N., Doucet, A. & Jordan, M.I. (2003)
An introduction to MCMC for machine learning. Machine
Learning, 50, 543.
Araujo, M.B. & Guisan, A. (2006) Five (or so) challenges for
species distribution modelling. Journal of Biogeography, 33,
16771688.
Asner, G.P. & Martin, R.E. (2009) Airborne spectranomics:
mapping canopy chemical and taxonomic diversity in
tropical forests. Frontiers in Ecology and the Environment, 7,
269276.
Asner, G.P., Powell, G.V.N., Mascaro, J., Knapp, D.E., Clark,
J.K., Jacobson, J., Kennedy-Bowdoin, T., Balaji, A.,
Paez-Acosta, G., Victoria, E., Secada, L., Valqui, M. &
Hughes, R.F. (2010) High-resolution forest carbon
stocks and emissions in the Amazon. Proceedings of the
National Academy of Sciences of Sciences USA, 107, 16738
16742.
Baldocchi, D., Falge, E., Gu, L. et al. (2001) FLUXNET: a new
tool to study the temporal and spatial variability of ecosystem-scale carbon dioxide, water vapor, and energy flux
densities. Bulletin of the American Meteorological Society, 82,
24152434.
Bohn, K., Dyke, J.G., Pavlick, R., Reineking, B. & Reu, B.
(2011) The relative importance of seed competition,
resource competition and perturbations on community
structure. Biogeosciences, 8, 11071120.
Journal of Biogeography 39, 22402252
2012 Blackwell Publishing Ltd

Dynamic vegetation models an inverse perspective

Bonan, G.B. (2008) Forests and climate change: forcings,
feedbacks, and the climate benefits of forests. Science, 320,
14441449.
Braswell, B.H., Sacks, W.J., Linder, E. & Schimel, D.S. (2005)
Estimating diurnal to annual ecosystem parameters by
synthesis of a carbon flux model with eddy covariance net
ecosystem exchange observations. Global Change Biology, 11,
335355.
Bugmann, H. (2001) A review of forest gap models. Climatic
Change, 51, 259305.
Bugmann, H.K.M. (1996) A simplified forest model to study
species composition along climate gradients. Ecology, 77,
20552074.
Butler, A., Doherty, R.M. & Marion, G. (2009) Model averaging to combine simulations of future global vegetation
carbon stocks. Environmetrics, 20, 791811.
Chambers, J.Q., Asner, G.P., Morton, D.C., Anderson, L.O.,
Saatchi, S.S., Espirito-Santo, F.D.B., Palace, M. & Souza, C.,
Jr (2007) Regional ecosystem structure and function: ecological insights from remote sensing of tropical forests.
Trends in Ecology and Evolution, 22, 414423.
Clark, J.S. (2005) Why environmental scientists are becoming
Bayesians. Ecology Letters, 8, 214.
Clark, J.S. (2010) Individuals and the variation needed for high
species diversity in forest trees. Science, 327, 11291132.
Condit, R., Ashton, P.S., Baker, P., Bunyavejchewin, S., Gunatilleke, S., Gunatilleke, N., Hubbell, S.P., Foster, R.B., Itoh, A.,
LaFrankie, J.V., Lee, H.S., Losos, E., Manokaran, N., Sukumar,
R. & Yamakura, T. (2000) Spatial patterns in the distribution
of tropical tree species. Science, 288, 14141418.
Conti, S. & OHagan, A. (2010) Bayesian emulation of complex multi-output and dynamic computer models. Journal of
Statistical Planning and Inference, 140, 640651.
Cramer, W., Bondeau, A., Woodward, F.I., Prentice, I.C.,
Betts, R.A., Brovkin, V., Cox, P.M., Fisher, V., Foley, J.A.,
Friend, A.D., Kucharik, C., Lomas, M.R., Ramankutty, N.,
Sitch, S., Smith, B., White, A. & Young-Molling, C. (2001)
Global response of terrestrial ecosystem structure and
function to CO2 and climate change: results from six
dynamic global vegetation models. Global Change Biology, 7,
357373.
Daz, S. & Cabido, M. (2001) Vive la difference: plant functional diversity matters to ecosystem processes. Trends in
Ecology and Evolution, 16, 646655.
Dislich, C., Gunter, S., Homeier, J., Schroder, B. & Huth, A.
(2009) Simulating forest dynamics of a tropical montane
forest in South Ecuador. Erdkunde, 63, 347364.
Dormann, C.F., McPherson, J.M., Araujo, M.B., Bivand, R.,
Bolliger, J., Carl, G., Davies, R.G., Hirzel, A., Jetz, W., Kissling, D., Kuhn, I., Ohlemuller, R., Peres-Neto, P.R.,
Reineking, B., Schroder, B., Schurr, F.M. & Wilson, R.
(2007) Methods to account for spatial autocorrelation in the
analysis of species distributional data: a review. Ecography,
30, 609628.
Dormann, C.F., Schymanski, S.J., Cabral, J., Chuine, I.,
Graham, C., Hartig, F., Kearney, M., Morin, X., RomerJournal of Biogeography 39, 22402252
2012 Blackwell Publishing Ltd

mann, C., Schroder, B. & Singer, A. (2012) Correlation and

process in species distribution models: bridging a dichotomy. Journal of Biogeography, 39, 21192131.
Elith, J. & Leathwick, J.R. (2009) Species distribution models:
ecological explanation and prediction across space and time.
Annual Review of Ecology, Evolution, and Systematics, 40,
677697.
Fensholt, R., Huber, S., Proud, S.R. & Mbow, C. (2010)
Detecting canopy water status using shortwave infrared
reflectance data from polar orbiting and geostationary
platforms. IEEE Journal of Selected Topics in Applied Earth
Observations and Remote Sensing, 3, 271285.
Friend, A.D., Arneth, A., Kiang, N.Y., Lomas, M., Ogee, J.,
Rodenbeck, C., Running, S.W., Santaren, J.-D., Sitch, S.,
Viovy, N., Woodward, F.I. & Zaehle, S. (2007) FLUXNET
and modelling the global carbon cycle. Global Change Biology, 13, 610633.
Frolking, S., Palace, M.W., Clark, D.B., Chambers, J.Q., Shugart, H.H. & Hurtt, G.C. (2009) Forest disturbance and
recovery: a general review in the context of spaceborne remote sensing of impacts on aboveground biomass and
canopy structure. Journal of Geophysical Research, 114,
G00E02.
Galbraith, D., Levy, P.E., Sitch, S., Huntingford, C., Cox, P.,
Williams, M. & Meir, P. (2010) Multiple mechanisms of
Amazonian forest biomass losses in three dynamic global
vegetation models under climate change. New Phytologist,
187, 647665.
Gelman, A., Carlin, J.B., Stern, H.S. & Rubin, D.B. (2003)
Bayesian data analysis, 2nd edn. Chapman & Hall/CRC,
Boca Raton, FL.
Gerten, D., Schaphoff, S., Haberlandt, U., Lucht, W. & Sitch, S.
(2004) Terrestrial vegetation and water balance hydrological evaluation of a dynamic global vegetation model.
Journal of Hydrology, 286, 249270.
Grace, J. (2004) Understanding and managing the global carbon cycle. Journal of Ecology, 92, 189202.
Grimm, V. & Railsback, S.F. (2012) Pattern-oriented modelling: a multi-scope for predictive systems ecology. Philosophical Transactions of the Royal Society B: Biological
Sciences, 367, 298310.
Gritti, E.S., Smith, B. & Sykes, M.T. (2006) Vulnerability of
Mediterranean Basin ecosystems to climate change and
invasion by exotic plant species. Journal of Biogeography, 33,
145157.
Guisan, A. & Thuiller, W. (2005) Predicting species distribution: offering more than simple habitat models. Ecology
Letters, 8, 9931009.
Hanson, P.J., Amthor, J.S., Wullschleger, S.D., Wilson, K.B.,
Grant, R.F., Hartley, A., Hui, D., Hunt, E.R., Jr, Johnson,
D.W., Kimball, J.S., King, A.W., Luo, Y., McNulty, G., Sun,
G., Thornton, P.E., Wang, S., Williams, M., Baldocchi, D.D.
& Cushman, R.M. (2004) Oak forest carbon and water
simulations: model intercomparisons and evaluations
against independent data. Ecological Monographs, 74, 443
489.
2249

F. Hartig et al.
Hartig, F., Calabrese, J.M., Reineking, B., Wiegand, T. & Huth,
A. (2011) Statistical inference for stochastic simulation
models theory and application. Ecology Letters, 14, 816
827.
Hickler, T., Vohland, K., Feehan, J., Miller, P.A., Smith, B.,
Costa, L., Giesecke, T., Fronzek, S., Carter, T., Cramer, W.,
Kuhn, I. & Sykes, M.T. (2012) Projecting the future distribution of European potential natural vegetation zones with
a generalized, tree species-based dynamic vegetation model.
Global Ecology and Biogeography, 21, 5063.
Higgins, S.I., Scheiter, S. & Sankaran, M. (2010) The stability
of African savannas: insights from the indirect estimation
of the parameters of a dynamic model. Ecology, 91, 1682
1692.
Higgins, S.I., OHara, R.B., Bykova, O., Cramer, M.D., Chuine,
I., Gerstner, E.-M., Hickler, T., Morin, X., Kearney, M.R.,
Midgley, G.F. & Scheiter, S. (2012) A physiological analogy
of the niche for projecting the potential distribution of
plants. Journal of Biogeography, 39, 21322145.
Hubbell, S.P., He, F., Condit, R., Borda-de-Agua, L., Kellner, J.
& ter Steege, H. (2008) How many tree species are there in
the Amazon and how many of them will go extinct? Proceedings of the National Academy of Sciences USA, 105,
1149811504.
Hurtt, G.C., Fisk, J., Thomas, R.Q., Dubayah, R., Moorcroft,
P.R. & Shugart, H.H. (2010) Linking models and data on
vegetation structure. Journal of Geophysical Research, 115,
G00E10.
Huth, A. & Ditzer, T. (2000) Simulation of the growth of a
lowland Dipterocarp rain forest with FORMIX3. Ecological
Modelling, 134, 125.
Jeltsch, F., Moloney, K.A., Schurr, F.M., Kochy, M. & Schwager, M. (2008) The state of plant population modelling in
light of environmental change. Perspectives in Plant Ecology,
Evolution and Systematics, 9, 171189.
Jenkins, J.C., Birdsey, R.A. & Pan, Y. (2001) Biomass and NPP
estimation for the mid-Atlantic region (USA) using plotlevel forest inventory data. Ecological Applications, 11, 1174
1193.
Johnson, J.B. & Omland, K.S. (2004) Model selection in
ecology and evolution. Trends in Ecology and Evolution, 19,
101108.
Kadane, J.B. & Lazar, N.A. (2004) Methods and criteria for
model selection. Journal of the American Statistical Association, 99, 279290.
Kass, R.E. & Wasserman, L. (1996) The selection of prior
distributions by formal rules. Journal of the American Statistical Association, 91, 13431370.
Kattge, J., Daz, S., Lavorel, S. et al. (2011) TRY a global
database of plant traits. Global Change Biology, 17, 29052935.
Kerr, Y.H. (2007) Soil moisture from space: where are we?
Hydrogeology Journal, 15, 117120.
Knorr, W. & Kattge, J. (2005) Inversion of terrestrial ecosystem
model parameter values against eddy covariance measurements by Monte Carlo sampling. Global Change Biology, 11,
13331351.
2250

Kohler, P. & Huth, A. (2010) Towards ground-truthing

of spaceborne estimates of above-ground life biomass and
leaf area index in tropical rain forests. Biogeosciences, 7,
25312543.
Kohler, P., Ditzer, T. & Huth, A. (2000) Concepts for the
aggregation of tropical tree species into functional types and
the application to Sabahs lowland rain forests. Journal of
Tropical Ecology, 16, 591602.
Kramer, K., Degen, B., Buschbom, J., Hickler, T., Thuiller, W.,
Sykes, M.T. & de Winter, W. (2010) Modelling exploration
of the future of European beech (Fagus sylvatica L.) under
climate change range, abundance, genetic diversity and
adaptive response. Forest Ecology and Management, 259,
22132222.
Krieger, G., Hajnsek, I., Papathanassiou, K.P., Younis, M. &
Moreira, A. (2010) Interferometric synthetic aperture radar
(SAR) missions employing formation flying. Proceedings of
the IEEE, 98, 816843.
Lasslop, G., Reichstein, M., Kattge, J. & Papale, D. (2008)
Influences of observation errors in eddy flux data on inverse
model parameter estimation. Biogeosciences, 5, 13111324.
Lefsky, M.A. (2010) A global forest canopy height map from
the Moderate Resolution Imaging Spectroradiometer and
the Geoscience Laser Altimeter System. Geophysical Research
Letters, 37, L15401.
Linder, H.P., Bykova, O., Dyke, J., Etienne, R.S., Hickler, T.,
Kuhn, I., Marion, G., Ohlemuller, R., Schymanski, S.J. &
Singer, A. (2012) Biotic modifiers, environmental modulation and species distribution models. Journal of Biogeography, 39, 21792190.
Lischke, H., Zimmermann, N.E., Bolliger, J., Rickebusch, S. &
Loffler, T.J. (2006) TreeMig: a forest-landscape model for
simulating spatio-temporal patterns from stand to landscape
scale. Ecological Modelling, 199, 409420.
Luo, Y., Ogle, K., Tucker, C., Fei, S., Gao, C., LaDeau, S., Clark,
J.S. & Schimel, D.S. (2011) Ecological forecasting and data
assimilation in a data-rich era. Ecological Applications, 21,
14291442.
Martnez, I., Wiegand, T., Camarero, J.J., Batllori, E. & Gutierrez, E. (2011) Disentangling the formation of contrasting
tree line physiognomies combining model selection and
Bayesian parameterization for simulation models. The
American Naturalist, 5, E136E152.
McMahon, S.M., Harrison, S.P., Armbruster, W.S., Bartlein,
P.J., Beale, C.M., Edwards, M.E., Kattge, J., Midgley, G.,
Morin, X. & Prentice, I.C. (2011) Improving assessment
and modelling of climate change impacts on global terrestrial biodiversity. Trends in Ecology and Evolution, 26, 249
259.
Medvigy, D., Wofsy, S.C., Munger, J.W., Hollinger, D.Y. &
Moorcroft, P.R. (2009) Mechanistic scaling of ecosystem
function and dynamics in space and time: Ecosystem
Demography model version 2. Journal of Geophysical Research, 114, G01002.
Miller, P.A., Giesecke, T., Hickler, T., Bradshaw, R.H.W., Smith,
B., Seppa, H., Valdes, P.J. & Sykes, M.T. (2008) Exploring
Journal of Biogeography 39, 22402252
2012 Blackwell Publishing Ltd

Dynamic vegetation models an inverse perspective

climatic and biotic controls on Holocene vegetation change in
Fennoscandia. Journal of Ecology, 96, 247259.
Moorcroft, P.R., Hurtt, G.C. & Pacala, S.W. (2001) A method
for scaling vegetation dynamics: the ecosystem demography
model (ED). Ecological Monographs, 71, 557586.
Pacala, S.W., Canham, C.D., Saponara, J., Silander, J.A., Kobe,
R.K. & Ribbens, E. (1996) Forest models defined by field
measurements: estimation, error analysis and dynamics.
Ecological Monographs, 66, 143.
Parmesan, C. (2006) Ecological and evolutionary responses to
recent climate change. Annual Review of Ecology, Evolution,
and Systematics, 37, 637669.
Patenaude, G., Milne, R., Van Oijen, M., Rowland, C. & Hill,
R. (2008) Integrating remote sensing datasets into ecological
modelling: a Bayesian approach. International Journal of
Remote Sensing, 29, 12951315.
Pereira, H.M., Leadley, P.W., Proenca, V. et al. (2010) Scenarios for global biodiversity in the 21st century. Science,
330, 14961501.
Pickett, S.T.A. (1989) Space-for-time substitution as an alternative to long-term studies. Long-term studies in ecology:
approaches and alternatives (ed. by G.E. Likens), pp. 110
135. Springer-Verlag, New York.
Prentice, I.C. & Harrison, S.P. (2009) Ecosystem effects of CO2
concentration: evidence from past climates. Climate of the
Past, 5, 297307.
Prentice, I.C., Bondeau, A., Cramer, W., Harrison, S.P., Hickler, T., Lucht, W., Sitch, S., Smith, B. & Sykes, M.T. (2007)
Dynamic global vegetation modeling: quantifying terrestrial
ecosystem responses to large-scale environmental change.
Terrestrial ecosystems in a changing world (ed. by J.G.
Canadell, D.E. Pataki and L.F. Pitelka), pp. 175192.
Springer-Verlag, New York.
Purves, D.W., Zavala, M.A., Ogle, K., Prieto, F. & Benayas,
J.M.R. (2007) Environmental heterogeneity, bird-mediated
directed dispersal, and oak woodland dynamics in Mediterranean Spain. Ecological Monographs, 77, 7797.
Purves, D.W., Lichstein, J.W., Strigul, N. & Pacala, S.W. (2008)
Predicting and understanding forest dynamics using a simple tractable model. Proceedings of the National Academy of
Sciences USA, 105, 1701817022.
Roscher, C., Schumacher, J., Baade, J., Wilcke, W., Gleixner,
G., Weisser, W.W., Schmid, B. & Schulze, E.-D. (2004) The
role of biodiversity for element cycling and trophic interactions: an experimental approach in a grassland community. Basic and Applied Ecology, 5, 107121.
Sankaran, M., Hanan, N.P., Scholes, R.J. et al. (2005) Determinants of woody cover in African savannas. Nature, 438,
846849.
Sato, H., Itoh, A. & Kohyama, T. (2007) SEIB-DGVM: a
new Dynamic Global Vegetation Model using a spatially
explicit individual-based approach. Ecological Modelling,
200, 279307.
Scheiter, S. & Higgins, S.I. (2009) Impacts of climate change on
the vegetation of Africa: an adaptive dynamic vegetation
modelling approach. Global Change Biology, 15, 22242246.
Journal of Biogeography 39, 22402252
2012 Blackwell Publishing Ltd

Schroder, B. & Seppelt, R. (2006) Analysis of patternprocess

interactions based on landscape models overview, general
concepts, and methodological issues. Ecological Modelling,
199, 505516.
Schurr, F.M., Pagel, J., Cabral, J.S., Groeneveld, J., Bykova, O.,
OHara, R.B., Hartig, F., Kissling, W.D., Linder, H.P.,
Midgley, G.F., Schroder, B., Singer, A. & Zimmermann, N.E.
(2012) How to understand species niches and range
dynamics: a demographic research agenda for biogeography.
Journal of Biogeography, 39, 21462162.
Shugart, H.H. (1998) Terrestrial ecosystems in changing environments. Cambridge University Press, Cambridge.
Sitch, S., Smith, B., Prentice, I.C., Arneth, A., Bondeau, A.,
Cramer, W., Kaplan, J.O., Levis, S., Lucht, W., Sykes, M.T.,
Thonicke, K. & Venevsky, S. (2003) Evaluation of ecosystem
dynamics, plant geography and terrestrial carbon cycling in
the LPJ dynamic global vegetation model. Global Change
Biology, 9, 161185.
Sitch, S., Huntingford, C., Gedney, N., Levy, P.E., Lomas, M.,
Piao, S.L., Betts, R., Ciais, P., Cox, P., Friedlingstein, P.,
Jones, C.D., Prentice, I.C. & Woodward, F.I. (2008) Evaluation of the terrestrial carbon cycle, future plant geography
and climate-carbon cycle feedbacks using five Dynamic
Global Vegetation Models (DGVMs). Global Change Biology,
14, 20152039.
Smith, B., Prentice, I.C. & Sykes, M.T. (2001) Representation
of vegetation dynamics in the modelling of terrestrial ecosystems: comparing two contrasting approaches within
European climate space. Global Ecology and Biogeography,
10, 621637.
ter Steege, H., Pitman, N.C.A., Phillips, O.L., Chave, J.,
Sabatier, D., Duque, A., Molino, J.-F., Prevost, M.-F.,
Spichiger, R., Castellanos, H., von Hildebrand, P. & Vasquez, R. (2006) Continental-scale patterns of canopy tree
composition and function across Amazonia. Nature, 443,
444447.
Stephens, P.A., Buskirk, S.W. & del Rio, C.M. (2007) Inference
in ecology and evolution. Trends in Ecology and Evolution,
22, 192197.
Sugita, S., Gaillard, M.-J. & Brostrom, A. (1999) Landscape
openness and pollen records: a simulation approach. The
Holocene, 9, 409421.
Van Oijen, M., Rougier, J. & Smith, R. (2005) Bayesian
calibration of process-based forest models: bridging the
gap between models and data. Tree Physiology, 25,
915927.
Weng, E., Luo, Y., Gao, C. & Oren, R. (2011) Uncertainty
analysis of forest carbon sink forecast with varying measurement errors: a data assimilation approach. Journal of
Plant Ecology, 4, 178191.
Williams, M., Richardson, A.D., Reichstein, M., Stoy, P.C.,
Peylin, P., Verbeeck, H., Carvalhais, N., Jung, M., Hollinger,
D.Y., Kattge, J., Leuning, R., Luo, Y., Tomelleri, E.,
Trudinger, C.M. & Wang, Y.P. (2009) Improving land
surface models with FLUXNET data. Biogeosciences, 6,
13411359.
2251

F. Hartig et al.
Xiaodong, Y. & Shugart, H.H. (2005) FAREAST: a forest gap
model to simulate dynamics and patterns of eastern Eurasian forests. Journal of Biogeography, 32, 16411658.
Zhang, X., Friedl, M.A., Schaaf, C.B., Strahler, A.H., Hodges,
J.C.F., Gao, F., Reed, B.C. & Huete, A. (2003) Monitoring
vegetation phenology using MODIS. Remote Sensing of
Environment, 84, 471475.
Zimmermann, N.E., Edwards, T.C., Graham, C.H., Pearman,
P.B. & Svenning, J.-C. (2010) New trends in species distribution modelling. Ecography, 33, 985989.
Zurell, D., Berger, U., Cabral, J.S., Jeltsch, F., Meynard, C.N.,
Munkemuller, T., Nehrbass, N., Pagel, J., Reineking, B.,
Schroder, B. & Grimm, V. (2009) The virtual ecologist
approach: simulating data and observers. Oikos, 119,
622635.

BIOSKETCH
Florian Hartig is interested in the processes that govern the
assembly, dynamics, distribution and evolution of ecological
communities. His current research focuses on diversity
patterns of tropical rain forests. All authors of this paper
share a common interest in vegetation modelling and/or
statistical methodology.
Author contributions: All authors worked together in designing and writing this review. F.H. coordinated the discussion
and led the writing.

Editor: Peter Pearman

The papers in this Special Issue arose from two workshops
entitled The ecological niche as a window to biodiversity held
on 2630 July 2010 and 2427 January 2011 in Arnoldshain
near Frankfurt, Germany. The workshops combined recent
advances in our empirical and theoretical understanding of the
niche with advances in statistical modelling, with the aim of
developing a more mechanistic theory of the niche. Funding
for the workshops was provided by the Biodiversity and
Climate Research Centre (BiK-F), which is part of the LOEWE
programme Landes-Offensive zur Entwicklung Wissenschaftlich-okonomischer Exzellenz of Hesses Ministry of Higher
Education, Research and the Arts.

2252

Journal of Biogeography 39, 22402252

2012 Blackwell Publishing Ltd