Seeing Forests through the Trees

Summary
Under a changing climate it remains unclear to what extent our forests will continue to provide their
current services to society. Key to assessing forest resilience is properly understanding how small-
scale heterogeneity affects overall forest function. Current dominant approaches neglect most or
all of this heterogeneity, calling their conclusions into question. With a focus on Sweden, with the
possibility to extend the project to Cambodia, this project will provide the evidence base to assess
the influence of topographic, compositional and structural heterogeneity on forest response to
environmental stress and develop a methodology to bridge across scales to deliver answers that
will inform both forest managers and policymakers.

Motivation
Forests are one of the primary natural resources in Sweden. Over the last 100 years, forestry has
transformed Sweden's landscapes in a remarkable fashion, shifting from structurally complex primary
forest, to even-aged plantations. But we are not only relying on forests for producing biomass for timber,
paper and energy. Forests are also providing other crucial ecosystem services (ES) to the society, such
as: recreation, water purification and pollination. Another ES for which forests are playing a vital role is
climate change mitigation1. Swedish forests are a strong carbon sink and the Swedish climate change
policy relies heavily on a continued carbon uptake1. Yet it is unclear whether forests will be able to
maintain the delivery of these services at the same level, or even at all, under a rapidly changing climate.
Recent events across the world show forests to be apparently increasingly susceptible to climate
change2. Being able to effectively assess the resilience to climate change of services delivered by a
particular forest, forest management or planting strategy is crucial to both forest owners and
policymakers alike.

A discourse has increasingly grown up around forest resilience to climate change2 and a range of
modelling studies have been conducted to try and assess this 3. However, these studies have neglected,
or only partially considered a key factor that could fundamentally change the conclusions that they draw:
The role of heterogeneity.

Forest response to stress varies according to topography, soil type4, species composition and stand
structure5. The difference in response can be dramatic and highly non-linear, even in a highly-managed
system such as Sweden, these factors typically vary substantially across a landscape6. That means that
taking into account heterogeneity from tree, to stand, to landscape scale is crucial in order to assess
overall system resilience in the face of a changing climate. I pose the hypothesis that within-
landscape heterogeneity substantially changes the magnitude, and even the sign, of forest
responses to climate stress, compared to a homogeneous landscape characterised by the most
abundant landscape characteristics. I further contend that topographic position, variation in functional
composition and in stand structural complexity are all major, and interacting, contributors to this
landscape-level response.

This project will test these hypotheses for boreal, temperate forests, using test cases in Sweden, whilst
developing technical infrastructure that will enable policy and management-relevant outputs. There are
already established plans to take it further to Cambodia as well as wider application globally.

Background
Recent evidence from Sweden6 shows that vegetation greenness response to the 2018 drought was
strongly dependent on both species composition and topographic position within the landscape, with the
sign of the response often differing (Fig. 1). Differing tree mortality responses based on topography have
also been reported in a pioneering study in the tropical forest 7 and productivity is well established to be
affected by slope and aspect. It has also been found that the mix of species can strongly influence
productivity, with mixed forest more productive compared to monocultures8 compared to monocultures.
Differences in the vulnerability of species to environmental stresses also implies that species diversity
strongly buffers forests against dramatic transitions 9. These diversity effects also appear to extend into
structural diversity. Ahlström et al.10 found that, after correcting for stand age, structurally-diverse pristine
forests had higher productivity compared to managed forests.

Figure 1: Results of the impact of the 2018 drought in Sweden on the greenness as an anomaly (z-
score) compared to the preceding 10-year period and how that differed between forest stands of different
composition and how the topography was a factor, represented here by the topographic position index
which is relating a grid cell to its surrounding cells. Results from master thesis6.

Despite this emerging evidence of the importance of heterogeneity, we lack (a) systematic
characterisation of how the different aspects of heterogeneity (topography, composition, structure)
interact with forest response to stress and (b) methodologies to scale up these effects across
landscapes and explore their response under climate change. Dynamic vegetation models (DVMs) have
been widely used to study long-term changes in biogeochemical exchanges in terrestrial ecosystems
as a response to climatic and environmental changes11. They use process-based mechanistic
representation of the main ecosystem processes to simulate biogeochemical cycling and vegetation
dynamics; this process-based approach makes them applicable to study the effects of changing
conditions, such as the forecasting of future climate change 12. DVMs have been shown to reproduce
broad-scale patterns and inter-annual trends in terrestrial ecosystem carbon exchanges with acceptable
accuracy13,14. They also provide an opportunity to follow vegetation structure through space and time.
However, the simplifications in the DVMs introduce errors and result in the loss of landscape variation
in the models. The current generation of DVMs neglect high-resolution topographic heterogeneity,
typically use a very limited number of plant functional types which are not capable of co-existing in the
same stand and greatly simplify or eliminate structural diversity.

Part of the challenge relates to a lack of suitable information to build model parameterisations. For
decades, understanding of forest structure and dynamics, along with wider forestry and forest ecology,
has been built on measuring diameter at breast height (DBH) and total height of trees 15. Generalised
allometric equations relate these measured quantities to biomass and other variables of interest.
Capturing the widespread spatial variability in allometry16 is necessary to allow us to understand how
stand structure varies as a function of environment17, species composition and competition18 (see also
Figure 2). To simulate the knock-on effects on forest function we need to combine modelling with direct
links between forest state and function. There have been studies looking at the connection between
carbon storage in an ecosystem and its biodiversity19, but paired analysis analyses across structurally,
topographically and compositionally differing forests (e.g. managed and unmanaged forests), that are
part of the same landscape or that shares certain environmental similarities, are missing. Advances in
these aspects can be combined with emerging potential of trait-based modelling approaches20 and the
attendant underlying data to capture how functional composition interacts over larger areas21.

How are key traits affected by: Competition, Landscape position, management? And how does this in
turn affect functions in the forest stands, and the associated ecosystem services?

We will answer these questions through conducting measurements and collecting data from literature
to synthesise and bringing that into the DVM, LPJ-GUESS13.

Measurements
Starting autumn 2021 colleagues Anders Ahlström (Lund University) and Daniel Metcalfe (Umeå
University) will establish long term forest monitoring plots where they will establish pairs of old growth
and managed forests, I will do measurements with a Terrestrial Laser Scanner (TLS) in these. The TLS
enable allometric variables to be obtained at a greatly reduced effort, with very high precision and without
destruction22. I recently led a successful application to the Lund University infrastructure fund to
purchase a TLS, which will revolutionise our ability to assess whole tree biomass, growth and structural
parameters within forest stands. With the TLS we are able to do more accurate measurements of trees
than with more traditional methods. This also makes it possible to do repeated measurements with a
higher frequency. Currently, estimating local allometry equations requires measurement of all
components of the trees in a plot, often through harvesting, and it is therefore unrealistic to capture
variations in allometry in space and time. Such limitations can be remedied with a large number of highly
precise assessments of tree allometry that the TLS would allow. Capturing this spatial variability would
allow us to study impact of stand heterogeneity, known to be widespread 16. In addition to the TLS
measurements, this project will also measure: wood density (WD), specific leaf area (SLA) and leaf
nitrogen concentration (NL), which are known to be important when assessing forest functioning23.

Vegetation models
LPJ-GUESS have a unique way of dealing with this in that it represents the variation over a landscape
by starting multiple duplicate stands with unique initial conditions, but what have been lacking so far is
to represent this variability over the landscape based on data, e.g. topography and soil structure.
Another level of heterogeneity that have not been incorporated into LPJ-GUESS, is varying traits, which
are known to vary for a species within one plot, and even more so between locations24. Following along
in what is known as trait-based modelling, where the traits of the simulated trees is allowed to vary
(constrained by data) and the trees with the best trait combinations are the ones that will thrive 20. No
one has until now, included all these levels of heterogeneity: i) landscape, soils, slopes and elevation;
ii) within forest stands, having real gaps in the simulated stands; iii) trait variation, both dynamic as a
response to the environment and genetic diversity; in a global ecosystem model. There is now a unique
opportunity to bring all of these together and put LPJ-GUESS in the front seat compared to other global
models. We will do this by first learning from Sweden to be able to extend further: Europe, tropical
regions where I and colleagues are working, Cambodia and Senegal. The current project is focused on
ii and iii, the implementation of i is in another PhD project.

Will adding these levels of heterogeneity change the projections of carbon balance in a
changing climate?

Project implementation
This project have 3 main objectives:

A. Understand the role of heterogeneity in the function and resilience of forests

B. Implement new levels of stand level heterogeneity in LPJ-GUESS
C. Communicate our findings with relevant stakeholders

An overview of the project implementation with approximate timing is available in Figure 3.

The role of heterogeneity

To reach objective A, we need to do more forest inventories, and in a structured manner. There is a
need to do paired measurements in e.g. managed and unmanaged forests on similar soils, slopes and
latitudes. With the TLS we will quantify the structural differences between forest stands located in
different parts of a landscape or with different management, with the TLS we will also be able to
quantify the edges in patches in the forest and to do more frequent measurements allowing for better
estimates of annual increments. Linking ongoing detailed plot scale ecosystem monitoring (structural
and functional), regular forest inventories (structural), remote sensing (structural and functional) will
enable us to extrapolate over larger areas. It is also of importance that we already in this step are
bringing in fractal analysis to be able to bring in the heterogeneity in the generalisation step, following
e.g. Jupp & Twiss25. Today there is a capacity to do this for Sweden and Cambodia by extending
already existing projects. We will start by focusing on Sweden and move on to Cambodia when we
have established the methods. If pending grants are successful, there is the possibility to extend to
other tropical regions such as: Senegal (dry tropical) and Rwanda (tropical gradient, wet- dry).

As entire forest stand level experiments are very sparse or non-existing26 what we can do instead is to
observe changes in ecosystems using remote sensing substituting time with space 26,27 to find
ecosystems facing the challenges of today in past records. When these have been found we can use
fractal analysis to attribute the variability in landscape responses to the various landscape elements 25,28.
Milestones:

A1. Perform measurements of traits in new and already ecosystem monitoring plots
A2. Establish the role of heterogeneity for tree functioning at stand level
A3. Synthesise the results and extrapolate the relationships from A2 to landscapes and beyond
using remote sensing

Model implementation
Objective B builds directly on A. Some of the features are already implemented in versions of LPJ-
GUESS, but all is not included in one version yet. I have implemented, in LPJ-GUESS, a dynamic
allometry that let the trees react to
changes in experienced competition
(not published, initial results in Figure
2). To further refine and publish this
will be among the tasks of PhD
student #2 in the project. The new
allometry allows for dynamic
responses from management
measures such as thinning on the
growth strategy of the trees, it also
allows for a better representation of
growth strategies, yielding a more
realistic representation of succession
compared to what is currently in the
model. Flexible traits, will be
implemented by me and tested and
evaluated by the student. Previously
we have had good experience of Figure 2: Allometric relationship between height and
employing students from the technical diameter at breast height for a spruce-beech forest in
faculty to do implementations in the southern Germany. Observations from Hans Pretsczh, the
LPJ-GUESS framework that are of a red line shows results using LPJ-GUESS with the
more technical nature, I will do that standard constant allometric relationship and in black, the
new updated allometry.
within this project as well for the
graphical user interface.

Milestones:

B1. New functions in LPJ-GUESS representing heterogeneity mechanistically

B2. Performing simulations of historic and future carbon balance using the updated LPJ-GUESS
B3. New or updated graphical user interface for LPJ-GUESS

Outreach and communication

Objective C will be done throughout the project, it is of importance to include the stakeholders early on
in the process, so that we are making adjustments to the graphical user-interface that will be of use for
the foresters.

The collaboration with e.g. Katam (see below) will facilitate bringing the results from this project to an
audience outside of academia. The app from the company Katam is also useful to include in the forest
inventories, I see that interaction as being continuing throughout the project and hopefully beyond.

I have not specified in Figure 3 when papers coming out of the work are anticipated, the main results
and thus papers, are expected in connection to A3 and B2.

Milestones:

C1. Stakeholder interactions, such as workshops

C2. Applications of the graphical user interface for LPJ-GUESS
 * 2023 2024 2025 2026 2027
C3. Aims Milestones Responsible
4 1 2 3 4 1 2 3 4 1 2 3 4 1 2 3 4 1 2 3
A. the role of A1 fieldwork a a a a PhD1,Tech, students
heterogeneity
A2 analysis b b b PhD1, PhD2,Tech,SO
A3 synthesis b b b PhD1, PhD2,SO
B. Model B1 science SO
implementation
B2 simulate b b b b PhD2,SO
B3. GUI c c c c c c c c c c c c Students, SO
C. Outreach and C1. stake b b SO with PhD students
communication
C2. GUI c c c c c c c c c c c c c c c c SO with PhD students

Figure 3 Gantt-chart showing the Aims and milestones with a rough estimate on when in the project they will
be addressed and implemented. a and b, if the students are hired in autumn 2022 and stays on track, this is
after their defences. c, the timing of this is not crucial, sometime after the first stakeholder interaction.

Group and networks

I am currently supervising 4 PhD students at INES, 3 that do work directly connected to this project. C.
Sovann (main supervisor) is working on quantifying ecosystem services across landscapes in
Cambodia, and how they have been affected by climate and land-use change. H. Zhou (co-supervisor),
is working on bringing landscape heterogeneity into LPJ-GUESS. A. Gärtner (co-supervisor) is building
a model on the fate of dead wood in forests with the aim of bringing it into LPJ-GUESS.

I am also co-supervising 3 PhD students at other universities, in common for them is that they are
applying model features that I have implemented but not published: H. Camargo (Birmingham, UK),
ozone stress in plants; M. Jianyong (KIT, Germany), biological nitrogen fixation in crops (manuscript in
review); Q, Guan (University of Copenhagen, Denmark), paddy rice (draft ready so submit).

I am part of the core development group of LPJ-GUESS in Lund. In the wider LPJ-GUESS community I
have collaborations with research groups in Germany: KIT, Garmisch-Partenkirchen; TUM, München;
SBiK-F, Frankfurt; PIK, Potsdam; and HIE in Sydney, Australia and IMBE in Aix, France and
UCPH, Copenhagen, Denmark.
Planned group
Within the project a PhD student and a lab-technician will be employed. The task of the technician will
be to do measurements primarily with the TLS and to apply and develop algorithms of using the point
clouds that are the result of the TLS. The tasks of the technician is not limited to the described project,
but might assist in other TLS related tasks. The PhD student will together with the technician do the
experiments and perform the analysis of the results. It is also a task for the PhD student to perform
simulations with LPJ-GUESS when the model implementation is done. I will do the implementation in
the model. Three senior researchers from INES will be loosely connected to the group: Torbern
Tagesson, remote sensing; Thomas Pugh, ecosystem modeller; Anders Ahlström, forest inventories,
they will be asked to co-supervise the students, depending on the needs of the hired student. Also the
students already working on projects connected to this (see above) will be welcome to participate.

Project partners
Within this project I will deepen the collaboration with Pretsczh, a renowned forestry professor of the
Technical University in München (TUM), he have deep knowledge in the field and the research questions
posed in this project, he have also agreed to host me and PhD students.

I will continue and broaden the already established collaborations with Kim Soben (Royal University of
Agriculture (RUA), Cambodia) on performing forest inventories in the already established plots in
Cambodia. There already plans to perform ecosystem modelling teaching at RUA, specifically targeting
the application of LPJ-GUESS for landscape studies within Cambodia.

Dan Metcalfe (Umeå University) and Anders Ahlström (INES) will establish long term ecosystem
monitoring plots in Sweden with the aim of addressing questions at the core of this project, namely the
difference between managed and natural forests and landscape position.
Katam, a Swedish company focused on bringing digitalisation to the forestry sector. They make use of
mobile applications and drones to deliver management plans to foresters. I have had initial talks with
them, and there is scope to take the lessons learned from this project into their allocation model

Within the three Lund University strong research areas BECC (Biodiversity and Ecosystem services in
a Changing Climate), MERGE (Modelling the regional and global earth system) and eSSENCE (The e-
Science Collaboration), I have established several collaborations on various aspects of modelling.

Fractal group, I have established a group (with initial funding from MERGE 2020) of experts from
different fields (soil scientist, animal ecologist, remote sensing, mathematician, etc.) to work on how to
make use of fractals when going from one scale to another within landscape analysis and in modelling
of landscapes. Some initial findings from that work (the allocation in trees show a fractal relationship29)
have lead up to the ideas put forward here, a path this project will continue.

References
1. Lundmark, T. et al. Potential Roles of Swedish Forestry in the Context of Climate Change Mitigation. For. 2014, Vol. 5,
Pages 557-578 5, 557–578 (2014).
2. Reyer, C. P. O. et al. Forest resilience and tipping points at different spatio-temporal scales: approaches and
challenges. J. Ecol. 103, 5–15 (2015).
3. Dolan, K. A. et al. Disturbance Distance: quantifying forests’ vulnerability to disturbance under current and future
conditions. Environ. Res. Lett. 12, 114015 (2017).
4. Fredeen, A. L., Bois, C. H., Janzen, D. T. & Sanborn, P. T. Comparison of coniferous forest carbon stocks between
old-growth and young second-growth forests on two soil types in central British Columbia, Canada.
https://doi.org/10.1139/x05-074 35, 1411–1421 (2011).
5. Calama, R. et al. Mixture mitigates the effect of climate change on the provision of relevant ecosystem services in
managed Pinus pinea L. forests. For. Ecol. Manage. 481, 118782 (2021).
6. Asch, J. Topographic controls of drought impact on Swedish primary forests. (Lund University, 2021).
7. Zuleta, D., Duque, A., Cardenas, D., Muller-Landau, H. C. & Davies, S. J. Drought-induced mortality patterns and rapid
biomass recovery in a terra firme forest in the Colombian Amazon. Ecology 98, 2538–2546 (2017).
8. Pretzsch, H. & Schütze, G. Tree species mixing can increase stand productivity, density, and growth efficiency and
attenuate the tradeoff between density and growth throughout the whole rotation. Ann. Bot. (2021)
doi:10.1093/AOB/MCAB077.
9. Sakschewski, B. et al. Resilience of Amazon forests emerges from plant trait diversity. Nat. Clim. Chang. 6, 1032–1036
(2016).
10. Ahlström, A., De Jong, G. E., Nijland, W. & Tagesson, T. Primary productivity of managed and pristine forests in
Sweden. Environ. Res. Lett. 15, 94067 (2020).
11. Prentice, I. C. et al. Dynamic Global Vegetation Modeling: Quantifying Terrestrial Ecosystem Responses to Large-
Scale Environmental Change. in Terrestrial Ecosystems in a Changing World (eds. Canadell, J., Pataki, D. & Pitelka,
L.) 175–192 (Springer Berlin Heidelberg, 2007). doi:10.1007/978-3-540-32730-1_15.
12. Friedlingstein, P. et al. Uncertainties in {CMIP5} Climate Projections due to Carbon Cycle Feedbacks. J. Clim. 27, 511–
526 (2013).
13. Smith, B. et al. Implications of incorporating {N} cycling and {N} limitations on primary production in an individual-based
dynamic vegetation model. Biogeosciences 11, 2027–2054 (2014).
14. Le Quéré, C. et al. Global carbon budget 2014. Earth Syst. Sci. Data Discuss. 7, 521–610 (2014).
15. Maas, H. G., Bienert, A., Scheller, S. & Keane, E. Automatic forest inventory parameter determination from terrestrial
laser scanner data. in International Journal of Remote Sensing vol. 29 1579–1593 (Taylor and Francis Ltd., 2008).
16. Hulshof, C. M., Swenson, N. G. & Weiser, M. D. Tree height-diameter allometry across the United States. Ecol. Evol. 5,
1193–1204 (2015).
17. Hulshof, C. M., Swenson, N. G. & Weiser, M. D. Tree height-diameter allometry across the United States. Ecol. Evol. 5,
1193–1204 (2015).
18. Pretzsch, H. Canopy space filling and tree crown morphology in mixed-species stands compared with monocultures.
Forest Ecology and Management vol. 327 251–264 (2014).
19. Strassburg, B. B. N. et al. Global congruence of carbon storage and biodiversity in terrestrial ecosystems. Conserv.
Lett. 3, 98–105 (2010).
20. Berzaghi, F. et al. Towards a New Generation of Trait-Flexible Vegetation Models. Trends in Ecology and Evolution
vol. 35 191–205 (2020).
21. Chapin, F. S. Effects of plant traits on ecosystem and regional processes: A conceptual framework for predicting the
consequences of global change. Annals of Botany vol. 91 455–463 (2003).
22. Calders, K. et al. Terrestrial laser scanning in forest ecology: Expanding the horizon. Remote Sensing of Environment
vol. 251 112102 (2020).
23. Reich, P. B. Key canopy traits drive forest productivity. Proc. R. Soc. B Biol. Sci. 279, 2128–2134 (2012).
24. Falster, D. S., Duursma, R. A. & FitzJohn, R. G. How functional traits influence plant growth and shade tolerance
across the life cycle. Proc. Natl. Acad. Sci. U. S. A. 115, E6789–E6798 (2018).
25. Jupp, T. E. & Twiss, S. D. A physically motivated index of subgrid-scale pattern. J. Geophys. Res. 111, D19112 (2006).
26. Ewers, R. M. et al. A large-scale forest fragmentation experiment: the Stability of Altered Forest Ecosystems Project.
Philos. Trans. R. Soc. B Biol. Sci. 366, 3292–3302 (2011).
27. Gardner, R. H., Milne, B. T., Turnei, M. G. & O’Neill, R. V. Neutral models for the analysis of broad-scale landscape
pattern. Landsc. Ecol. 1987 11 1, 19–28 (1987).
28. Milne, B. T. Measuring the fractal geometry of landscapes. Appl. Math. Comput. 27, 67–79 (1988).
29. Duursma, R. A. et al. Self-shading affects allometric scaling in trees. Funct. Ecol. 24, 723–730 (2010).