Ecosystems (2017) 20: 229–236

DOI: 10.1007/s10021-016-0071-2
Ó 2016 Springer Science+Business Media New York

20th Anniversary Paper

Next-Generation Individual-Based
Models Integrate Biodiversity and
Ecosystems: Yes We Can, and Yes We
Must
Volker Grimm,1,2* Daniel Ayllón,1 and Steven F. Railsback3

1
Department of Ecological Modelling, Helmholtz Centre for Environmental Research-UFZ, Permoserstr. 15, 04318 Leipzig, Germany;
2
German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Deutscher Platz 5e, 04103 Leipzig, Germany; 3Lang
Railsback & Associates, 250 California Avenue, Arcata, California 95521, USA

ABSTRACT
Ecosystem and community ecology have evolved
along different pathways, with little overlap.
However, to meet societal demands for predicting
changes in ecosystem services, the functional and
structural view dominating these two branches of
ecology, respectively, must be integrated. Biodi-
versity–ecosystem function research has addressed
this integration for two decades, but full integration
that makes predictions relevant to practical prob-
lems is still lacking. We argue that full integration
requires going, in both branches, deeper by taking
into account individual organisms and the evolu-
tionary and physico-chemical principles that drive
their behavior. Individual-based models are a ma-
jor tool for this integration. They have matured by
using individual-level mechanism to replace the
demographic thinking which dominates classical
theoretical ecology. Existing individual-based
ecosystem models already have proven useful both
for theory and application. Still, next-generation
individual-based models will increasingly use
standardized and re-usable submodels to represent
behaviors and mechanisms such as growth, uptake
of nutrients, foraging, and home range behavior.
The strategy of pattern-oriented modeling then
helps make such ecosystem models structurally
realistic by developing theory for individual
behaviors just detailed enough to reproduce and
explain patterns observed at the system level. Next-
generation ecosystem scientists should include the
individual-based approach in their toolkit and focus
on addressing real systems because theory devel-
opment and solving applied problems go hand-in-
hand in individual-based ecology.
Key words: biodiversity research; emergence;
first principles; individual-based ecology; individ-
ual-based modeling; pattern-oriented modeling;
predictions; structural realism; theory develop-
ment.

Received 7 May 2016; accepted 17 July 2016;
published online 10 November 2016
Author contributions VG, SR, and DA wrote the paper; DA did the
literature review of Table 1.
*Corresponding author; e-mail: volker.grimm@ufz.de
INTRODUCTION
In Richard Wagner’s opera ‘‘Siegfried,’’ the dwarf
Mime tries to weld together the pieces of the magic
but broken sword Nothung. After welding fails,
Siegfried, a headstrong and uncouth young man,

229
230 V. Grimm and others
tries an entirely new approach: he reduces No-
thung to filings, melts them down, and succeeds in
re-forging the sword entirely anew. The broken
sword reminds us of the separate disciplines of
ecosystem and community ecology, not yet inte-
grated via traditional approaches such as aggre-
gated system-level modeling. We certainly do not
suggest reducing ecosystem and community ecol-
ogy to filings, but the metaphor lies in our con-
viction that we need to revert to the building blocks
of communities and ecosystems, individual organ-
isms, before a true integration can be achieved.
The separation of ecology into largely disjunct
fields has been bemoaned for decades, for example
from the perspective of modeling: ‘‘Each level of
organization has traditionally been a separate disci-
pline (for example, physiological ecology, behav-
ioral ecology, population ecology, and ecosystem
ecology). Each field has its own set of phenomena to
explain, and most have their own distinctive types of
models.’’ (Huston and others 1988, p. 690).
Ecosystem ecology, for example, focuses on biogeo-
chemistry and the stocks and flows of energy and lim-
iting nutrients, whereas community ecology focuses on
organisms and structure in terms of species richness and
composition. Organisms and species were ignored in
ecosystem ecology, while energy and nutrients were
ignored in community ecology. Over the last two dec-
ades, though, a merger has started—referred to as bio-
diversity–ecosystem function (BEF) research (Loreau
2010; Cardinale and others 2012)—which aims at
integrating structural and functional perspectives. This
merger was driven by the societal demand to under-
stand and predict how ecosystem function and services
respond to changes in land use and climate. To meet this
demand, neither energy and nutrient constraints nor
organisms and how they interact with each other and
their environment can be ignored (Evans and others
2013a; Mokany and others 2016).
Still, there is no true integration of ecosystem
and community ecology yet. BEF research often
relates biodiversity patterns to ecosystem functions
such as productivity and stability properties, but
mostly via small-scale experiments or meta-analy-
ses, which have limited ability to elucidate the
mechanisms underlying observed relationships.
These experiments have also focused on grasslands,
so research over more systems and larger scales is
still needed. Moreover, BEF research often does not
explicitly refer to biogeochemistry and energy
considerations. Loreau (2010) and others before
him (Schulze and Mooney 1993; Tilman 1999)
have therefore called for a more mechanistic inte-
gration of biodiversity research, which is largely
community ecology, with ecosystem ecology.
How could this integration be achieved? Loreau
(2010) proposes ‘‘community evolution models,’’ in
which trait compositions evolve in response to
environment and species interactions. This approach
has led to interesting results, but the models are
usually aggregated at the population or species level
and again often only weakly linked to ecosystem
ecology and evolutionary principles. A fundamen-
tally different approach was suggested almost
30 years ago: individual-based modeling. In their
still intriguing vision, Huston and others (1988, p.
690) wrote: ‘‘Individual-based models link all of
these separate levels in the ecological hierarchy. The
responses of individuals to their local environment
are based on physiological and behavioral responses.
The aggregation of all individuals of a species pro-
duces the population dynamics of that species.
The aggregation of all individuals of many species
interacting with each other and with their environ-
ment produces community dynamics. Ecosystem
dynamics result from the aggregation of individual-
environment interactions into large-scale material
and energy fluxes’’.
Huston and others (1988) believed that individ-
ual-based models will unify ecological theory and
predicted their rapid development. Progress,
though, has been slow (Grimm 1999; Grimm and
Railsback 2005; DeAngelis and Grimm 2014). Re-
cent attempts to outline how structure and function
could be integrated in large-scale ecosystem models
do not even mention the individual-based approach
(Loreau 2010; Mokany and others 2016). IBMs are
an established tool for population ecology, but many
ecologists consider IBMs unsuitable for communities
and ecosystems. Although we acknowledge why this
notion exists, we consider it fundamentally wrong.
Here, we discuss why progress integrating com-
munity and ecosystem ecology has been slow and
how, nevertheless, major obstacles have been
overcome recently. Our central tenet is that, iron-
ically, ecosystem and community ecology can only
be fully and mechanistically integrated when we
revert to their building blocks and acknowledge
individual organisms and their traits and adaptive
behaviors. We believe, as Huston and others
(1988), that a unified ecology must start from
individual-based ecology (Grimm and Railsback
2005). This belief does not imply that all ecological
models should be individual-based but that indi-
vidual-based models must play a more central role
by complementing more aggregated approaches.
We discuss what this perspective means for com-
bining empirical study with modeling and theory in
ecosystem ecology and for the training of ecosys-
tem ecologists for the 21st century.
 Next-Generation Individual-Based Models Integrate Biodiversity and Ecosystems
FROM DEMOGRAPHIC THINKING TO FIRST
PRINCIPLES
Classical theoretical ecology focuses on demographic
rates of populations. Even if these rates depend on
size, stage, or age in structured population models,
they remain empirical (Figure 1). They reflect the
observed net outcome of survival and reproduction
in a certain group of organisms over a certain period
of time. These rates can be used to extrapolate pop-
ulation growth rates, but only for those environ-
mental conditions and densities at which the rates
were observed. This dependence on empirical rates
limits the usefulness of classical theory for predicting
responses to novel conditions and for linking eco-
logical dynamics to first principles like fitness seek-
ing and energy budgets.
Although most early IBMs also used demographic
rates as input, newer IBMs increasingly have demo-
graphics that emerge from mechanisms such as fit-
ness-seeking behavior (Railsback and Harvey 2002;
Stillman and Goss-Custard 2010), energy budgets
(Martin and others 2012, 2013; Sibly and others 2013;
van der Vaart and others 2016), stoichiometry (Kaiser
and others 2014; Smith and others 2014), or photo-
synthesis (Fischer and others 2016). These mecha-
nisms make IBMs more complex but also more
coherent because they are based on first princi-
ples—processes we can observe in the laboratory in
real organisms or believe from the fundamental
concepts of biochemistry, evolution, and so on. The
same mechanisms can be used for different species
and contexts. For example, the energy budget theory
of Kooijman (2010; see also Martin and others 2012,
Figure 1. The structure and dynamics of ecological sys-
tems emerge from how individual organisms perform
and how they interact with their biotic and abiotic
environment. Vital and growth are used in models as a
summary representation of processes at the individuals’
level, but they are linked to specific environmental
conditions.
231
2013; Sibly and others 2013) uses the same model for
all heterotrophic species, with species differing only in
parameterization. Likewise, ecophysiological forest
gap models like FORMIND are based on general
submodels of photosynthesis and competition for
light (Köhler and Huth 1998; Fischer and others
2016). Moreover, unlike system-level models, IBMs
are naturally suited for adaptive mechanisms from life
history selection (for example, Railsback and Harvey
2013) to microevolution (for example, Ayllón and
others 2016).
The research fields dealing with these first princi-
ples usually focus on the individual organism and
ignore the population context. Next-generation
individual-based models (Grimm and Berger 2016)
link these principles to higher levels: populations
and communities. Because model individuals can
respond to and process nutrients and energy
explicitly, IBMs of this type link community and
ecosystem ecology. Pioneering prototypes of such
models were already developed more than 20 years
ago, about moose foraging affecting plant commu-
nities and then nutrient cycling dynamics (Pastor
and Naiman 1992; Pastor and others 1998). IBMs
also link community and ecosystem ecology by
allowing adaptation and evolution as driving pro-
cesses (Loreau 2010).
The integration of community and ecosystem
ecology must start from the community side, as in
BEF research, as this is the less aggregated ap-
proach. Integration has thus to add function to
structure, or biogeochemistry to demographics.
However, classical models of community ecology,
still mostly Lotka–Volterra type, do not provide
mechanistic interfaces to nutrients and energy. This
missing link can be made by going down two levels
of organization: to individuals and the principles
that drive their behavior.
FROM POPULATIONS TO COMMUNITIES AND
ECOSYSTEMS: COPING WITH COMPLEXITY
IBMs of populations can be complex if they are
based on first principles and designed to predict the
response of populations to heterogeneous and
changing environments. Developing them can be
‘‘big science,’’ requiring years for development and
testing (Stillman and others 2015).
Extrapolating from this experience with popula-
tion IBMs makes IBMs of species-rich communities
and ecosystems seem impossible. However,
ecosystem IBMs have existed for decades, in par-
ticular for forests (Botkin and others 1972; Shugart
1984; Bugmann 2001) and, to a lesser extent,
marine fish communities (Shin and Cury 2001;
232 V. Grimm and others
Giacomini and others 2009, 2013; Rose and others
2010). The trick to keep complexity at bay for
multiple species is to use the same submodels for all
species within a functional group, for example,
trees, and the same submodel for interactions
among individuals of all species. Forest gap models
(Botkin and others 1972), for example, use the
same growth equation for all trees and represent
species differences via parameters. Competition
among trees for light depends only on the vertical
leaf area distribution. Other plant models represent
competition via overlapping zones of influence
(Weiner and others 2001; May and others 2009;
Lin and others 2012), which can also be used for
animals (Booth 1997; Piou and others 2007). In
marine fish community models, trophic interac-
tions are based on size differences among individ-
uals, independent of species (Shin and Cury 2001;
Giacomini and others 2009, 2013). Grimm and
Berger (2016) describe elements of ‘‘next-genera-
tion ecological modeling’’ that help cope with
complexity while aiming for structural realism and
predictions.
YES, WE CAN: MODELS LINKING
BIODIVERSITY AND ECOSYSTEMS EXIST
Quite a few models linking communities and
ecosystems already use the individual-based ap-
proach. However, these models are, perhaps be-
cause of their complexity, not yet generally
perceived as good starting points for integrating
biodiversity and ecosystems. In biodiversity re-
search, they seem to largely be ignored. In Table 1,
which was inspired by a similar table in Mokany
and others (2016), we compiled an overview of
spatially explicit, dynamic IBMs that link ecosys-
tem structure and function with varying levels of
integration. These models use a wide range of ap-
proaches and scopes and address the full spectrum
of scales of application and resolution. They de-
scribe composition and diversity either by treating
each of multiple species differently or by applying
trait-based functional groupings, and thus are
capable of simulating driving ecological processes at
both species and ecosystem level.
These IBMs incorporate, to different degrees,
many of the essential modeling strategies and fea-
tures of next-generation ecological models and
therefore have the potential to provide not only
predictions of ecosystem dynamics in a changing
world but also insights into fundamental theoreti-
cal and applied issues in ecosystem ecology (for
example, stability properties and resilience and the
mechanisms controlling them, biodiversity, and
ecosystem services). However, the design of these
models is still very heterogeneous, reflecting dif-
ferent backgrounds, modeling traditions, and re-
search questions of their developers. We expect
that in the future individual-based modeling of
ecosystem will be more coherent by using generic
representation of individual organisms and their
interactions.
FROM OBSERVED PATTERNS TO STRUCTURAL
REALISM, PREDICTIONS, AND THEORY
As key features of next-generation ecological
models, Grimm and Berger (2016) list emergence,
structural realism, and prediction. Emergence from
first principles is needed to predict responses to
novel conditions, but how can we achieve struc-
tural realism? Striving for structural realism means
trying to make models reproduce observed patterns
for the right reasons instead of forcing the right
output via calibration: being able to parameterize a
Lotka–Volterra community model to reproduce, for
example, realistic cycles does not mean the model
captures the mechanisms actually causing the cy-
cles.
Pattern-oriented modeling (POM; Grimm and
others 1996; Wiegand and others 2003; Grimm and
others 2005; Grimm and Railsback 2012) is a gen-
eral strategy to achieve structural realism. Multiple
patterns, observed at different scales and levels of
observations, are used in model design as filters for
(1) selecting variables and processes, (2) designing
submodels for mechanisms including individual
behavior, and (3) parameterizing full models. POM
is not unique to ecological IBMs: using observed
patterns (referred to as ‘‘stylized facts’’ in eco-
nomics) to select variables and processes, and to
parameterize models (known also as ‘‘inverse
modeling’’), is common in many kinds of model-
ing, often unconsciously. Moreover, the pattern-
oriented development of submodels corresponds to
the rationale of ‘‘strong inference’’ introduced by
Platt (1964). What is new is using patterns to find
IBM submodels that link individual-level mecha-
nisms to higher levels of organization. In particular,
realistic mechanisms can be designed via a
hypothesis-testing approach: posing alternative
submodels for individual behavior and falsifying
them if they do not cause the IBM to reproduce
patterns observed at higher levels.
There are several examples of this POM strategy
for developing models of individual traits that ex-
plain population or higher-level ecological
Table 1. Main Features Defining the Structural Realism of a Number of Existing Spatially Explicit, Dynamic Individual-Based Models Linking
Ecosystem Structure and Function
Element Features/Models1 aDGVM2 Madingley OSMOSE Giacomini iLand FORMIND WIST WEAVER DOVE

Scales2 Spatial scale of application Global Global Regional Regional Landscape Stand Local Micro-site Virtual
Fine spatial resolution - - - Pot. + + + + Na
Fine temporal resolution + + + + + + + + Na
Ecosystem Multitrophic - + + + – – + + +
structure Multispecies/functional groups + + + + + + + + +
and function Parameterized at species level + - + + + + + + +
Non-trophic interactions + - - - + + + + -
Mass/energy fluxes across trophic - + + + - - + + +
levels
Biogeochemistry + - + - + + + - -
Model struc- First principles + + + + + + + + +
tural realism Spatio-temporal heterogeneity + + + + + + - + -
Trait-based approach + + - + - + + + +
Microevolution + - – + - - - + +
Resilience3 + (a,b) + (a) + (a,c) + (d) + (e) + (e) + (e) + (e) Nd
Model allows for simulation of + Pot. + Pot. + + Pot. + Pot.
multiple-stressors
Link species distribution to mecha- + + - + + + Pot. + Pot.
nisms
Modeling Pattern-oriented theory develop- + Nd Nd Nd Nd Nd Nd Nd Nd
strategies ment
Use of standardized generic sub- + + + + + + + + +
models
Use of standard models of interac- + + + + + + + + +
tions
Pattern-oriented calibration/testing + + + - + + - - -
Robustness analysis4 Nd Nd Nd Nd + Nd Nd Nd Nd
Human Can explicitly incorporate human Pot. Pot. + Pot. + + Pot. Pot. Pot.
dimension management actions

‘‘+’’ means ‘‘yes,’’ ‘‘-‘‘ means ‘‘no’’.

1
Model scope and reference: aDGVM2 (dynamic global vegetation; Scheiter and others 2013), Madingley (global ecosystem; Harfoot and others 2014), OSMOSE coupled with ROMS-N2P2D2Z2 (end-to-end marine ecosystem; Shin and Cury
2001; Travers-Trolet and others 2014), Giacomini (marine fish community; Giacomini and others 2009, 2013), iLand (forest landscape; Seidl and others 2012), FORMIND (forest gap; Köhler and Huth 1998; Fischer and others 2016),
WIST (terrestrial food web; Parrott and Kok 2002), WEAVER (soil food web; Moya-Laraño and others 2014), DOVE (terrestrial food web; Peacor and others 2007).
2
The spatial scale shown refers to the typical scale of application the model was designed for, but most models can be applied at different scales. Fine spatio-temporal resolution refers, for example, to <1 km2 and <1 month, respectively.
3
Resilience theories addressed, examples: (a) system stability and persistence, (b) shifts in trait diversity after fire, (c) regime shifts, (d) stability and persistence under environmental gradients, (e) recovery and persistence after natural/
anthropogenic disturbance events.
4
Next-Generation Individual-Based Models Integrate Biodiversity and Ecosystems

Robustness analysis refers to the systematic deconstruction of a model by forcefully changing the model’s parameters, structure, and representation of processes to detect the model mechanisms that explain a certain phenomenon break
down (Grimm and Berger 2016).
5
Nd Not documented, Pot. Potentially, Na not apply.
233
234 V. Grimm and others
dynamics (for example, Railsback and Harvey
2002; Amano and others 2006; Cury and others
2008; Railsback and Johnson 2011). In a study
designed in part to illustrate POM, Railsback and
Johnson (2011) developed a model of bird foraging
behavior for an IBM of how pest-control services
by songbirds depend on land use on coffee farms.
They identified nine patterns observed at the indi-
vidual bird, bird population, and pest population
levels. Then they posed four alternative submodels
of foraging behavior as hypothesized traits. These
traits included a ‘‘null’’ model (random foraging), a
simple model that assumed birds can sense local
prey availability, a simple model that assumed birds
can sense prey over large distances, and a model
based on classical optimal foraging theory. They
implemented each of the four submodels in a sep-
arate version of their IBM and conducted simula-
tion experiments to see which versions reproduced
which observed patterns. They found that only the
simple model with local sensing of prey availability
could reproduce all the observed patterns. The high
level of structural realism achieved with this ap-
proach made it possible to predict bird populations,
their pest-control services, and coffee production
under alternative scenarios of land sharing versus
land sparing (Railsback and Johnson 2014).
POM is the link we need between empirical sci-
ence and theory: we use empirically observed pat-
terns to develop theory (submodels; Grimm and
Railsback 2005) for individual-level mechanisms
that explains, via the IBM, observed patterns at the
population, community, and ecosystem levels.
Theory development via POM is a cycle: after one
round of testing hypothesized submodels against
observed patterns, we can identify additional
observations that would help further refine the
theory. Therefore, POM is a productive way to
identify the right mix of empirical science and
theory in ecosystem ecology: we should collect
empirical observations that help us develop and
refine theory via the POM cycle.
NEXT-GENERATION ECOSYSTEM
ECOLOGISTS: SEEING THE TREES FOR THE
FOREST, AND THE FOREST FOR THE TREES
We claim here that ecosystem ecology can be
integrated with community ecology and biodiver-
sity research by going all the way down to the
individual level. This idea may seem irrational until
we remember that individuals are where theory is
most firmly tied to reality: we can easily measure
and model individual mechanisms (physiology, and
so on) in the laboratory, and the ecological theory
we have most confidence in is that individuals
behave to increase their future fitness. Therefore,
as Huston and others (1988) pointed out, letting
system dynamics emerge from what individuals do
is a productive way to integrate ecological levels. If
we focus on individual-based approaches to
ecosystems, how does our research and education
change?
The biggest overall change we need is learning to
think and model across ecological levels: instead of
building models and testing them with data just on
ecosystems, we need to think about and model
how ecosystem characteristics emerge from char-
acteristics and behaviors of individuals. But when
modeling and measuring individuals, we need to
search for simple representations of individuals that
are useful for explaining higher-level phenomena.
We do not want, for example, everything we know
about individual energetics in an IBM but only just
enough to model energy flow through an ecosys-
tem of unique and adaptive individuals. Seeing not
only the forest for the trees, but also the trees for
the forest is an apt metaphor for this kind of inte-
grative individual-based ecology of communities
and ecosystems.
A second change is focusing more on real sys-
tems and applied problems, and making predictions
useful to managers. Classical theoretical ecology
has sacrificed prediction for generality, which has
delayed the development of predictive theories for
decades (Evans and others 2013b). Ecosystem
ecology traditionally has a strong applied focus
(Mokany and others 2016), but biodiversity re-
search is still largely focused on theoretical issues
like stability–diversity relationships. A stronger fo-
cus on applied questions and realistic settings is
important not just to make ecology more relevant,
but also because it produces better models and
science (Stillman and others 2015). Addressing
applied problems not only forces us to tackle hard
and, therefore, unexplored questions, but also
makes productive strategies such as POM possible.
POM greatly facilitates model and theory develop-
ment but only works in real systems with real
observations.
Using POM also implies a closer integration of
theoretical and empirical research. Individual-
based ecology and POM provide a productive
framework for designing empirical research that
directly supports theory and model development.
This approach is a way to design laboratory and
field studies at the individual level, where research
is much less expensive and uncertain, that still
contribute directly to system-level understanding.
 Next-Generation Individual-Based Models Integrate Biodiversity and Ecosystems
We also continue to need system-level observa-
tions, but can focus on (often, qualitative and
short-term) patterns useful for developing and
testing IBMs instead of more extensive and quan-
titative data. In the current fixation on ‘‘big data’’
and huge and ever-growing databases of, for
example, plant traits it is important to remember
that data by themselves are useless; it is multiple
patterns that we need to identify and decode. As
Platt (1964) put it: ‘‘Many—perhaps most—of the
great issues of science are qualitative, not quanti-
tative, even in physics and chemistry. Equa-
tions and measurements are useful when and only
when they are related to proof: but proof or dis-
proof comes first and is in fact strongest when it is
absolutely convincing without any quantitative
measurement.’’ (p. 352).
A final change is a culture change in theoretical
ecology, so ‘‘theoretical’’ no longer means ‘‘hypo-
thetical’’ but instead refers to the kinds of theory
used in hard sciences: models shown via testing to
be useful and general for solving real problems. The
POM theory development cycle is a hypothesis-
testing approach for developing and documenting
the ability of IBMs to explain and predict ecosystem
phenomena.
Returning to the broken sword metaphor from
the Introduction, we are convinced that the bot-
tom-up approach we suggest is needed to make
progress in ecosystems ecology and is feasible.
Simple and elegant models that can be formulated
and analyzed within a few weeks are still important
as heuristic tools, but predictive modeling of
ecosystems, to tackle both applied and basic ques-
tions, will to a large degree be ‘‘big science,’’
requiring long-term collaboration of interdisci-
plinary teams.
After re-forging the magic sword Nothung,
Siegfried demonstrates its strength by cutting, with
one strike, the anvil in two pieces. This is where the
metaphor ends, but it confirms our vision of a
powerful predictive and integrated ecosystem and
biodiversity research, where structure and function
are linked via the individual-based perspective.
ACKNOWLEDGEMENTS
We thank Monica Turner and Steve Carpenter for
their invitation to contribute to this special feature
of Ecosystems and for their comments on an earlier
draft.
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