Current Forestry Reports

https://doi.org/10.1007/s40725-023-00189-y

MODELLING PRODUCTIVITY AND FUNCTION (A ALMEIDA, SECTION EDITOR)

Latest Trends in Modelling Forest Ecosystems: New Approaches or Just

New Methods?
Juan A. Blanco1   · Yueh‑Hsin Lo1 

Accepted: 26 May 2023

© The Author(s) 2023

Abstract
Purpose of Review  Forest models are becoming essential tools in forest research, management, and policymaking but cur-
rently are under deep transformation. In this review of the most recent literature (2018–2022), we aim to provide an updated
general view of the main topics currently attracting the efforts of forest modelers, the trends already in place, and some of
the current and future challenges that the field will face.
Recent Findings  Four major topics attracting most of on current modelling efforts: data acquisition, productivity estimation,
ecological pattern predictions, and forest management related to ecosystem services. Although the topics may seem different,
they all are converging towards integrated modelling approaches by the pressure of climate change as the major coalescent
force, pushing current research efforts into integrated mechanistic, cross-scale simulations of forest functioning and structure.
Summary  We conclude that forest modelling is experiencing an exciting but challenging time, due to the combination of new
methods to easily acquire massive amounts of data, new techniques to statistically process such data, and refinements in mechanistic
modelling that are incorporating higher levels of ecological complexity and breaking traditional barriers in spatial and temporal
scales. However, new available data and techniques are also creating new challenges. In any case, forest modelling is increasingly
acknowledged as a community and interdisciplinary effort. As such, ways to deliver simplified versions or easy entry points to
models should be encouraged to integrate non-modelers stakeholders into the modelling process since its inception. This should be
considered particularly as academic forest modelers may be increasing the ecological and mathematical complexity of forest models.

Keywords  Forest modelling · Forests · Methods

Introduction how the ecosystem functions (i.e., nutrient, water and energy
cycles, gene flows, population, and successional changes).
Forests are one of the most complex ecosystems on Earth’s All this natural complexity poses a true challenge for
biosphere, as they host a large proportion of terrestrial bio- representing forest structure and functioning in scientific and
diversity and exist at the interface between the atmosphere technical studies, as well as for science-based management
and the pedosphere. In addition, forests are defined as such [2]. Traditionally, forest models have focused on the
because the dominant organisms are trees, which are long- dominant organisms (trees) and how they grow, survive, and
lived immobile individuals that are usually large [1]. These are distributed [3••]. This approach has been dominant since
features provide opportunity for forests to develop specific the beginning of early quantitative forestry in the eighteenth
spatial and temporal structures that have direct influence on century. However, for the last few decades, it has been well
known that understanding how trees function is not enough to
understand how forests function, as other forest components
(understory, wildlife, soil, and microbial communities) are
* Juan A. Blanco
juan.blanco@unavarra.es also influencing trees. Hence, forest models have constantly
evolved to incorporate some of forests’ complexity into their
Yueh‑Hsin Lo
yuehhsin.lo@gmail.com algorithms in order to produce the estimations that model
developers consider necessary to meet their objectives.
1
Departamento de Ciencias, Institute for Multidisciplinary The development of the first forest growth simula-
Applied Biology (IMAB), Universidad Pública de Navarra tor marked the beginning of a new approach to estimate
(UPNA), 31006 Pamplona, Spain

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tree growth. Since then, modelling has evolved from the “forest function,” “forest distribution,” “forest adaptation,”
data-based approach of using statistical tools to transform and “modelling forest function” (with their alternative spell-
observed data (“empirical models”) into an approach in ings) in the title and keywords of documents. We identified
which an understanding of causal relationships between var- a total of 4933 documents. Among those, we selected 154
iables was added to statistical relationships in order to pre- papers that were reviews of different modelling topics. After
dict variables of interest (“process-based models”) [4]. Soon screening for relevance, the selected review papers used for
after, the advantages and disadvantages of both approaches our narrative review were reduced to 79.
were identified [5, 6], and with the aim of solving them, an On a second phase, to objectively identify the most popu-
intermediary approach was proposed [7]. Since then, for- lar topics in the most recent literature, we used the visu-
est models have evolved considerably, and in the last few alization tool VOSviewer [13] with the database of 4933
years, important technical developments have revolution- documents to map the relationships between their keywords.
ized the forest modelling field [8], such as the following: the However, as the statistical term “random forests” was dis-
continuous increment of computing power [9]; the develop- torting the database (data not shown), we removed the
ment of new statistical methods [10]; the great expansion documents with this term. As a result, we retained 2040
in techniques for data acquisition such as LiDAR, spectral, documents for keyword mapping with VOSviewer v1.6.18
hyperspectral, thermal, or radar sensors that can be applied (Centre for Science and Technology Studies, Leiden Univer-
at broad scales [11]; or the development of autonomous sity, the Netherlands, http://​www.​vosvi​ewer.​com). We lim-
continuous measurement devices for soil, vegetation, and ited the minimum number of occurrences for each keyword
atmospheric variables [12]. Therefore, the aim of this review displayed in the map to 30 (Fig. 1). As a result, 20 different
is to identify the current focuses in forest modelling that are keywords were selected. This search was not intended to be
capturing most of the research effort. a formal or in-depth quantitative review but merely a way to
gain an unbiased and up-to-date insight on current popular
modelling trends.
Current Main Topics in Forest Modelling As main result of the keyword mapping, we found the
term “climate change” as the most cited. Climate change
To identify the current trends in forest modelling, we first also stood out in a central position among all other terms.
carried out a search in the Web of Science database (https://​ In addition, four different clusters of terms were identified,
www.​webof​scien​ce.​com/​wos/​woscc/​advan​ced-​search) for with climate change being the main connector among them.
the years 2018 to 2022 using the terms “forest modelling,” The first cluster (in red in Fig. 1) could be considered as

Fig. 1  Keyword map show-

ing relationships between the
20 most common keywords
in documents related to forest
modelling published in the Web
of Science in the 2018–2022
period. Different line and dot
colors indicate different clusters
of terms. Dot size is propor-
tional to the frequency of each
keyword, and line thickness is
proportional to the frequency
of co-occurrence of connected
keywords

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Current Forestry Reports

built around quantitative assessments of vegetation biomass precipitation. For example, a scarce availability of models
(or carbon) using remote-sensing techniques (either aerial able to link climate change with ecological disturbances has
or terrestrial). The second cluster (in yellow in Fig. 1) was been identified [26]. Similarly, most regeneration algorithms
composed of the relationships between growth, productiv- used in forest models do not capture the effect of climate
ity, and climate. The third cluster (in blue in Fig. 1) was change [27]. In any case, climate change needs to be directly
limited to more technical terms related to model building. linked to modelling physiological responses (e.g., phenol-
Finally, the fourth cluster (in green in Fig. 1) was related to ogy, photosynthesis, respiration) and to frequency and sever-
ecosystem services and management, in combination with ity of disturbances (fire, drought, insects’ outbreaks, etc.).
climate change. Below, we discuss the main trends in each In turn, changes in these processes will also affect other
of these clusters in the following sections, based on the 79 ecosystem processes (allocation, allometry, growth at tree
review papers identified as relevant. level, biodiversity, and competition at ecosystem level), and
therefore, simulating climate change effects will indirectly
be needed to improve how such processes are modelled.
Climate Change: the Main Driver for Forest
Modeling
Remote Sensing and Biomass Accounting
It is not surprising that climate change is at the center of
current forest modelling efforts, a pattern already noticed Biomass (in the form of timber, firewood, cork, fruit, resin,
in other recent reviews [14]. This result could just reflect charcoal, etc.) has traditionally been the most important
the generalized wish by forest researchers to link their work commodity obtained from forests. Therefore, it is not sur-
to the current widespread scientific polices focused on prising that the different ways to estimate forest biomass and
addressing climate change, but it could also genuinely indi- other closely related variables (i.e., timber volume, carbon)
cate the need for understanding how complex systems such are still among the most important topics in current forest
as forests will behave under unknown climate conditions. modelling efforts (Fig. 1). Among them, modelling strate-
Climate change is being observed as a major force behind gies to sequester C stands out as one of the most important
many changes in current and future forest environmental topics [28]. The large size and immobile nature of trees
changes [15–18]. Such changes will affect in different ways allow individual features such as diameter and height to be
the key factors driving tree physiology, and therefore, new measured at different times over extended periods. Such
modelling approaches need to disaggregate climate influ- an inventory-based approach can provide a wealth of data,
ences on those drivers. Hence, understanding detailed effects but it quickly becomes a cumbersome task when large and
of climate change, alone or in combination with other major diverse forest areas need to be assessed. However, the explo-
drivers for change such as land-use change or biodiversity sive development of remote-sensing techniques, the lower-
loss, is obviously the ultimate goal of much of the current ing prices of unmanned aerial vehicles, and the continuous
modelling effort. growth in computing capabilities are generating the ability
The realization of the first signs of climate change and the to finally obtain detailed assessment of not only the basic
need for early action in forest management well in advance population features but also the structure and spatial distri-
of other economic sectors (due to the long-lived nature of bution of individual trees over large areas [29••].
trees) has meant that for at least two decades, the need to A model convergence towards the tree scale for mean-
provide forest models with capabilities to simulate climate ingful C-cycle modelling, both from upscaling more physi-
change has been recognized [19, 20]. Such need has meant ologically oriented models and downscaling stand-level C
that the use of simple correlational models using traditional accounting models, has been noted [30•]. However, not until
data from permanent plots or inventories has long been seen very recently have researchers looked for ways to incorporate
as inadequate among the scientific community for climate structural diversity into process-based models. A detailed
change-related studies, although such an approach can be review on the potential and limitations of using terrestrial
very suitable for other research and management applica- laser scanning to calibrate functional-structural plant models
tions [21]. In addition, other models that had implicit rep- is available [31•]. One of the main advantages of linking
resentation of climate influences have moved into explicit both modelling approaches is the potential to include physi-
representations to keep up with the knowledge demands on ological models into a realistic structure of plant communi-
climate change effects on forest systems from different stake- ties. This could move structural modelling from individual
holders [22–25]. to community level. In fact, there are suggestions that the
Nevertheless, for the successful implementation of cli- merging of allometry, empirical observation, remote sens-
mate change simulation capabilities into forest models, mod- ing, and individual-based modelling will contribute to a
elers need to move beyond direct effects on temperature and more unified vision of forest ecology [32]. However, to reach

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such a level of integration, proper processing of terrestrial the usual time steps in forest models (seasonal or annual)
laser scanning data is needed. In addition, researchers should are too large to capture C allocation dynamics and resource
avoid the temptation of upscaling functional-structural plant acquisition. In summary, the generalized use of allometry
models to the landscape level, as it will be challenging due and inventory-based approaches is just not adequate to cap-
to the potential to misrepresent other ecological processes ture short-term C dynamics [40•].
more relevant at such a spatial scale [33, 34].
Another important challenge to incorporate more remote
sensing into forest models is the need for increased meas-
ures of standardization and uncertainty in observations [35]. Patterns vs. Processes
However, these authors also highlight the high potential of
remote-sensing data to automatize carbon models, which A second main topic in current forest modelling research
currently need manual and time-consuming calibration. is the development of new and refined methodological
In this respect, several issues have been identified when approaches, mostly through the use of advanced mathe-
increasing the importance of remote data acquisition of matical or statistical tools or by borrowing them from other
canopy structure, such as the need for standardization of fields. New progress is made almost daily in deep learn-
modeling approaches, the need for open datasets, the need to ing methods than are revolutionizing modern ecology [41].
improve allometric models, and the need for stronger valida- These methods have great potential to improve computa-
tion protocols [29••]. tionally costly tasks such as classification of information
Allometric models are as important as remote sensing to from remote sensing or simulation of interactions between
estimate timber volume, biomass, or carbon stocks. Such individuals in large forest areas. The use of these advanced
models have been extensively used in the past but usually statistical techniques is greatly expanding modelling capa-
using data from pure and coetaneous stands [14, 35]. This bilities to link research done at multiple scales, to simulate
situation introduces an important bias when estimating car- larger regions, and to incorporate dynamic changes at shorter
bon or biomass stocks in natural forests, which are usually temporal scales (crucial for accurate C flux modelling).
multispecies and multiaged, as species allometry changes in The need for such increasingly powerful approaches is
the presence of competitors [36]. Hence, using allometric clear by the two keywords highlighted in our review for this
equations from pure stands could be a source of uncertainty cluster (“prediction” and “pattern,” Fig. 1). There is a dire
when modelling mixed stands, as there are significant dif- need for tools that can provide usable predictions for manag-
ferences in allometry for a given species when growing in ers, as the forestry sector needs to adapt to climate change
a single- vs. multiple-species stand [37]. In addition, many even earlier than other sectors, given the long-term conse-
of these allometric models do not include climatic or stand- quences of current management decisions [42]. Hence, using
level features [14], although recent research has been under- techniques to simplify model use will undoubtedly facilitate
taken to address these shortcomings [25, 36]. the generation of tools easy to interpret and to share with
The combination of different remote-sensing techniques non-modelers, and that can be easily compared with expert
such as LiDAR and radar can help to accurately model for- knowledge [43]. This idea of simplification while retaining
est structure [29••]. An additional feature of models based the behavior of complex process-based models is behind the
on remote sensing is the potential to simulate and estimate developments of “model emulators” [44].
radiation levels through the canopy and on the understory Model emulators are built to mimic the same outputs
based on 3D data from LiDAR measurements. For exam- from complex (usually process-based) models, with the
ple, the division of the canopy into volumetric pixels (or main objectives of reducing computing requirements. This
“voxels”) allows for simulating the interaction between trees, simplification allows for integration of the emulator in other
understory, and radiation at individual-tree levels or even modelling platforms (and therefore connectivity with other
smaller scales. In fact, 3D canopy simulation can be a more models or submodules different from the original process-
reliable way to estimate energy and C fluxes than traditional based model), to expand temporal and spatial scales not
inventory-based approaches [38]. In addition, such models reachable with the original process-based models or to sim-
could help in improving connections between forest and plify interaction with model users. Such expansion of the
atmosphere models [39]. basic model could be crucial to understand ecological pat-
Additional issues when simulating C fluxes, particularly terns that emerge at higher scales and that otherwise would
C allocation, have been identified [40•]. These authors have not have been directly inferred by the underlying process-
highlighted that the common use of fixed ratios for allocat- based model [45]. Hence, emulators could be valuable tools
ing C to plant organs is a severe oversimplification under in the future to understand ecological patterns at large scales,
climate change, as it removes from the model the sensitivity particularly under novel ecological conditions created by the
to environmental conditions and disturbances. In addition, combination of climate, biodiversity, and land-use changes.

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Current Forestry Reports

However, the development of emulators also brings an developers (https://​mwp.​org/​link-​to-​mwp-​digit​al-​cerem​ony-​

important challenge to the field of ecological modelling. and-​sympo​sium/).
The substitution of process-based algorithms by machine- Productivity estimations will remain crucial in the near
learning based decision rules offers clear advantages. None- and medium future, as commercial forestry will likely
theless, it could also be considered as a process to create become more focused on high-yield intensively managed
“black box” models in which scientific understanding of plantations to sequester and substitute carbon-intensive
ecological process is impeded, as the mechanisms behind materials. Conservation forestry will increasingly expand
such processes are simplified to just algorithms that have the into forest reserves around the world to increase stored
same outcomes. A detailed review on model simplification carbon and protect biodiversity. In this context, the devel-
is available [46]. opment of basic (but management-friendly) correlational
Obviously, this situation highlights the need for a dual models such as allometric and inventory-based models is
direction in scientific advancement: while emulators are needed [35, 52, 53]. However, the need to include climate in
clearly useful tools to study ecological patterns, ecologi- all these new models is certainly a challenge for new regions
cal processes can be better studied with mechanistic pro- and species, as they would need either long-term data series
cess-based models (although such a division is not so clear or an extensive network of inventory plots to account for
[44]). Advancement along both lines will also support the climatic influences on tree growth rates or allometry. Hence,
development of “digital twins”: computer-based copies of new developments in automatic and climate-sensitive tree
real forests constructed to mimic the most intricate patterns monitoring devices may be helpful [12].
and processes, with visualization of virtual stands as one of
their main strengths. These digital tools are already being
proposed to train managers and researchers in understand- Modelling Forests Beyond Trees
ing how climate change and new management techniques
could facilitate the transition of the forest sector towards While tree growth and productivity are still an important
novel conditions [47]. Obviously, digital twins depend not topic, the largest cluster of research topics identified was
only on the simulation and visualization techniques used related to modelling forest components other than trees.
but also on particularly the availability of quality data to Most of this research is based on the clear understanding
calibrate them. Here again, remote sensing, forest inven- that for models to be able to handle climate change effects,
tories, and traditional fieldwork data will be crucial, as the it is essential to include more ecosystem components that
old-fashioned rule in ecological modelling is still valid: in historically have received less attention [21].
the absence of adequate data, all different modelling options Some key issues are the improved assessment of carbon
are equally valid [43]. and water cycles. For example, it has been stated that those
models using drought indexes that include an evaporative
component work better, but also that there is just a small
number of studies actually evaluating drought indexes
Productivity Still a Concern against physiological indicators of water stress [54]. In this
respect, a recent review of the way in which the representa-
A third popular topic in current forest modelling is related to tion of evapotranspiration processes has evolved in forest
forest productivity and growth (Fig. 1). This indicates that, models has noticed a trend towards the simplification from
even if for more than two decades efforts have been made to the initial attempts, achieved by the availability of more
add non-timber forest products to forest models (i.e., [48]), empirical data and model evaluation tests that have allowed
estimating forest productivity is still a major issue in the the refinement of simulation algorithms [55]. These authors
field. This research cluster is clearly focused on how tree have pointed that such simplification allows for further con-
growth is influenced by climate. One of the key features of nections to water flow models and the scalability of such
the research on this topic are the continuous calls for devel- research. This is an important advance, as the common
opment of new growth models for species and regions out- oversimplification of eco-hydrological processes in models
side North America, Europe, and to a lesser extent Asia [49, makes linkages with socio-economic values more difficult
50••]. An example of successful model application around to evaluate [56•]. These authors also pointed to a lack of
the world is the spread in the use of 3-PG, which was origi- empirical work on effects of water availability in forest pro-
nally developed for Australian eucalyptus plantations but has ductivity. Further developments that mechanistically link
been embraced and modified for its application in multiple hydraulic conductance with physiology, growth and mor-
regions and stand types [51]. The widespread application of tality are taking place [57–59].
3-PG by scientists and managers was recognized in 2020 by How biodiversity is integrated into forest models is
the Marcus Wallenberg Prize which was awarded to their another of the issues in this keyword cluster. Traditionally,

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there is a biodiversity bias towards trees in forest models concept of plant functional traits) and different ways to
[60]. This is not surprising, as biodiversity interactions (both incorporate linkages between plant and herbivores into
animal and vegetal) in forests are a complex and broad field process-based models have been suggested [3 ••]. This
that have not been incorporated into models until relatively issue is not limited to tropical or natural forests, as the
recently, and that still remains largely ignored in operational influence of large herbivores on tree and shrub density in
models used in forest management. In this regard, a lack of boreal [66] and temperate forests [67] has been reported,
integration of modelling approaches at different spatiotem- with or without management.
poral scales has been identified as a barrier to implement Other approaches to account for biodiversity include the
biodiversity into forest modelling [61]. Similarly, calls for use of habitat and species distribution models. They link the
more attention to the role of understory in key ecological smallest (habitat) to the largest (distribution) spatial scales
processes have been raised [62], even if early examples of and provide a better understanding of the potential impacts
the importance of tree-understory interactions when simu- of novel ecological conditions over the mid to long term.
lating commercial forestry are available [e.g., 63]. It is cur- The dramatic increase of available data on climate, soils,
rently advocated that the most efficient approach is to use and species distributions allows for finely gridded model-
plant functional traits that can accommodate the inherent ling at both temporal and spatial scales. This advance allows
complexity of understory communities. To do so, models statistically based species distribution models to be linked to
must have detailed time and spatial scale to allow for the process-based models [16, 68], although better understand-
different ecophysiological behaviors (many times resource ing of absence data and improved inclusion of abiotic inter-
opportunistic) that understory species usually display, par- actions will become crucial to estimate effects of climate
ticularly following disturbances [64]. change [69, 70].
An important effort currently taking place in vegeta- Finally, an always-important topic in forest models is the
tion science is determining how functional traits can be integration of management into modelling. Such integration
applied to models to understand how species with different has two clear foci: simulation of management practices and
traits interact. An important and ongoing development is involvement of forest managers into the modelling process
to expand the functional trait approach being developed for [71]. As forest management is inherently an ecological dis-
vegetation studies [65]. This is particularly important in turbance, including management simulation in forest mod-
highly diverse ecosystems such as tropical forests in which elling should not be limited to anthropogenic actions but
it is unrealistic to simulate forest dynamics with only a should include natural disturbances as well. However, the
few dominant species.The functional trait approach is now main limitation that needs to be solved is the lack of infor-
being expanded to model species interactions including mation on the specific mechanisms that link climate change
animals, particularly herbivores. However, mechanistic with disturbances [26]. This is especially important when
models of forest pests are usually based on correlations several disturbances can be connected through cascading
between environmental variables (e.g., degree days) and effects on the ecosystem [42, 72].
growth rates (usually at individual or population scales), Important conceptual advances in disaggregating distur-
and limited to some of the pest’s life cycle stages. Hence, bances into their constituent components and embedding
there is a need for models able to integrate current algo- disturbances into system dynamics have been recently com-
rithms that simulate specific pest and pathogens at differ- pleted [50••]. These authors have identified as important
ent development stages to obtain meaningful estimates of challenges the need for simulating nondeterministic compet-
their interactions with the rest of forest components [43]. itive interactions between tree species and their responses to
More intriguingly, concerns have been raised around the disturbances and suggest using life history traits to overcome
usually forgotten role of megafauna in forest models [3••]. this issue. However, although these linkages among distur-
Although it has been traditionally assumed that the effects bances have been long recognized in forestry, little research
of megafauna are realized at the forest level through seed has actually incorporated them into forest models, particu-
dispersal, arguments exist to also consider their impacts larly as multi-disturbance models [50••]. In addition, most
on nutrient cycling and plant demography, such as the role models that incorporate disturbances predict probabilities
of megafauna on predation of plant reproductive organs, for such disturbances to happen depending on different stand
mortality caused by herbivory or trampling, and nutrient features, but not the disturbances effects [26]. Among dis-
redistribution related to animal residues [3••]. A serious turbances, wildfire modelling is an important field by itself.
effort to better understand the role of megafauna in forests As in the case of other disturbances, abiotic factors such
is needed, given the current situation of defaunation in as slope, elevation, distance to roads, or weather patterns
many areas of the world, which in some areas is trying are important for incorporating complexity at small spatial
to be reversed by rewilding actions. The use of “herbi- and temporal scales [73]. However, getting good quality for
vore functional traits” (equivalent to the already accepted such small-scale variables could be a challenge in areas with

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dense forest cover and sparse road networks, as is the case such forests usually have limited resources to access and
in most tropical or boreal forests [74]. use models, and models usually lack representations of
Another important step in making forest models more external factors (such as the vicinity of agriculture lands)
meaningful for stakeholders include modifying the way that can be relevant for the functioning and structure of
models are created. The focus on participatory processes in small forests [76].
which model users and forest stakeholders interact with for- • Urban forests: As urban landscapes expand, urban forests
est modelers during the inception of the modelling studies is are becoming very important in delivering a multitude of
being increasingly recognized as fundamental for the model ecosystem services. However, urban forest models have
to make actual impact in the forest sector [44]. This approach been developed only for few regions around the world (i.e.,
aims to bring nonacademic forest stakeholders into the pro- USA, Europe, and China) and are mostly correlational in
cess at the beginning, so they develop a sense of ownership nature. To better assess the effects of climate change on
of the research outcome and therefore are much more likely ecosystem services, better linkages with ecophysiological
to implement the model outcomes. Three models for science- mechanisms must be incorporated into urban forest models
policy interaction have identified [74]: the “linear phase” [49]. Among the potential ecosystem services that could be
when science informed policy-making in a unidirectional modelled in urban forests are not only carbon sequestration
manner, the “interactive phase” when both sides found them- [78] but also aesthetic values [79].
selves in a continuous interaction, and the “embedded phase.” • The Global South: A recurrent finding in all recent for-
Our own experience is that the linear phase is still dominant est modelling reviews is the strong bias towards North
in many regions, with scientists developing models and sce- America and Europe [38, 50••, 52], followed by East
narios of their interest and then approaching nonacademic Asia to a lesser extent (mostly China and Japan). Some
stakeholders with their results. Only in some scarce cases isolated modelling hotspots in the southern hemisphere
the interaction has progressed and moved into the second are Australia (which has generated one of the most suc-
stage of science-policy interaction (i.e., [44]). It is then time cessful forest models, [51]) and Brazil (mostly focused
to push towards a multi-actor approach (the second “interac- on modelling plantation forests but also generated some
tive” phase of bringing science into practice). However, to work on Amazonian forests). More effort must be made
achieve this goal, models need to be accessible, relevant, and to better understand the applicability of models from
user-friendly for non-modelers and address current forest other regions to these areas that are underrepresented in
management concerns to actually bring change into forestry the scientific modelling literature. This is an important
practices [76]. A comparison on how different European deci- research area given regional variations in terms of tree,
sion support systems are facing these challenges has identified understory and wildlife species composition, and other
the need to incorporate forest owner behavior and accurate environmental constraints such as climate, edaphic fac-
spatial analysis to better estimate landscape-level provisioning tors, or human management models.
of ecosystem services [77]. • Overlooked physio-ecological processes: Two impor-
tant mechanisms have attracted little attention in forest
models until now. One is regeneration (including mast-
Next Challenges for Forest Model ing), which is now recognized as a process that can sig-
Convergence nificantly affect biomass allocation and hence carbon
and energy flows. Even if detailed conceptual models
Understanding how complex ecosystems such as forests are on forest regeneration have been available for some
structured and function as a system has been, still is, and time (i.e., [80]), regeneration has usually been oversim-
will be challenging. The challenge lies in understanding how plified in forest models [81]. However, recent important
climate change affects forests, while our understanding on advances in understanding the masting process allow
how to model forests under “normal” conditions is still far for the implementation of mechanistic models [82].
from complete. In addition to the most popular topics cur- Giving the inherent complexity and current incomplete
rently being explored in forest modelling discussed earlier, understanding of the process, modelling regeneration
we have identified through our review several topics that patterns could be a more practical approach than mod-
deserve mention due to their relevance, even if they did not elling processes in order to avoid error propagation,
explicitly appear in the keyword map in Fig. 1. Such topics especially if models are to be scaled up to regional or
include the following: larger areas [83 ••]. Another overlooked topic is root
growth and function. Traditionally, the simulation of
• Small forests: Landscapes around the globe are becoming fine roots has been underdeveloped compared to leaves,
increasingly fractioned, making small forests of a few and hence, a common approach has used allometric
hectares or smaller increasingly common. Managers of relationships of fine roots to other biomass fractions

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[84]. However, the latest research indicates that this is Funding  Open Access funding provided by Universidad Pública de
not always appropriate, but also that enough data for Navarra.
mechanistic root models are starting to be available Data Availability  The data presented in this study are available on
[85]. Given the important role of roots in carbon, nutri- request from the corresponding author.
ent and water cycles, and the influence of such cycles
on tree mortality [86], a more mechanistic modelling Declarations 
approach would be desirable.
Human and Animal Rights and Informed Consent  This article does not
• Uncertainty assessment: Traditionally, the study of cli-
contain any studies with human or animal subjects performed by any
mate change effects on forest has relied on modelling of the authors.
different climate scenarios, management options, and
their interactions. However, such an approach does not Conflict of Interest Juan A. Blanco and Yueh-Hsin Lo declare that
provide a clear picture of the uncertainty around model they have no conflict of interest.
predictions. Hence, moving from scenario assessment
Open Access  This article is licensed under a Creative Commons Attri-
towards uncertainty analysis has been proposed [56•, bution 4.0 International License, which permits use, sharing, adapta-
63]. To do so, using predictions from different models tion, distribution and reproduction in any medium or format, as long
would be useful, particularly if the models use different as you give appropriate credit to the original author(s) and the source,
approaches [16]. The viability of assessing uncertainty provide a link to the Creative Commons licence, and indicate if changes
were made. The images or other third party material in this article are
through using envelopes of models has been demon- included in the article's Creative Commons licence, unless indicated
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Conclusions

Our review of current trends in forest modelling has

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