Chapter 6

Changes of Tree and Stand Growth:

Review and Implications

H. Pretzsch, M. del Río, F. Giammarchi, E. Uhl, and R. Tognetti

Abstract In this chapter, we review the current long-term growth trends and short-­
term growth reaction to single or repeated stress events on tree and stand level in
Europe. Based on growth trend analyses, the chapter reveals the strong human foot-
print on forest ecosystems.
First, we use long-term experiments and increment cores to show change in
growth trends within the last centuries. Growth reactions are caused by deposition
and climate change rather than by silvicultural measures. Second, we look closer on
regional-specific deviations from the general trend. Climate change, drought events,
acid rain and O3 are causing regional-specific growth reaction patterns. Third, we
assess stress events and the resilience and resistance of monospecific and mixed
stands against biotic and abiotic stress in view of the ongoing growth trends.

H. Pretzsch (*)
Forest Growth and Yield Science, TUM School of Life Sciences Weihenstephan, Technical
University of Munich, Freising, Germany
e-mail: [email protected]
M. del Río
INIA, Forest Research Centre, Madrid, Spain
iuFOR, Sustainable Forest Management Research Institute, University of Valladolid & INIA,
Valladolid, Spain
e-mail: [email protected]
F. Giammarchi
Faculty of Science and Technology, Free University of Bolzano-Bozen, Bolzano, Italy
e-mail: [email protected]
E. Uhl
Forest Growth and Yield Science, TUM School of Life Sciences Weihenstephan, Technical
University of Munich, Freising, Germany
Bavarian State Institute of Forestry (LWF), Freising, Germany
e-mail: [email protected]
R. Tognetti
Dipartimento di Agricoltura, Ambiente e Alimenti, Università degli Studi del Molise,
Campobasso, Italy
e-mail: [email protected]

© The Author(s) 2022 189

R. Tognetti et al. (eds.), Climate-Smart Forestry in Mountain Regions, Managing
Forest Ecosystems 40, https://doi.org/10.1007/978-3-030-80767-2_6
190 H. Pretzsch et al.

The revealed tree and stand growth behaviours are highly relevant, as any changes
of forest growth and structure have strong impacts on the provision of goods and
ecosystem services. The results underline the importance of biomonitoring and sug-
gest counteracting measures by forest planning, adaptation of silvicultural guide-
lines for existing forest and innovative design of future forests stands.

6.1 I ntroduction: The Information Potential of Tree

and Stand Growth Trajectories

At first sight, the reports about the status quo of forest health, vitality and productiv-
ity in Europe seem contradictory; on the one hand messages about growth accelera-
tion (Kauppi et al. 2014), on the other hand forest dieback due to stress factors (Allen
et al. 2015). The situation seems similarly contrary at the global level, with accelerat-
ing growth in the northern latitudes and growth decline in the Mediterranean and dry
continental zones. At higher altitudes, the findings are similar to northern latitudes;
some tree species in some regions seem to benefit from the changing environmental
conditions, and other species in various regions suffer. In terms of canopy cover,
European and North American forests have experienced a history of recovery
(Nabuurs et al. 2013; Zhu et al. 2018). Climate change and land use changes interfere
with forest regrowth, changing the temporal trajectories of biomass recovery and the
age structure of forest stands. These forests, although continuing to sequester a sig-
nificant amount of carbon, already show signs of growth saturation, which may limit
their sink potential in the decades to come, as it will gradually saturate as trees age.
Information on carbon uptake in regrowth forests derives from modelling exercises
(Pugh et al. 2019). Sink saturation occurs due to a decrease in the assimilation com-
ponent or an increase in the emission component for a given period of time. Global
terrestrial carbon sink in forests is largely driven by tree physiological responses to
increasing CO2, N deposition, air pollution, evaporative demand, land use changes
(vegetation shift, forest regrowth) and changes in forest cover (forest fire, forest
dieback).
Certainly, the growth reaction pattern of a species depends on the respective site
conditions and the specific environmental changes. When growing at its ecological
optimum, i.e. in the centre of its realised ecological niche, a species will be less
affected by environmental changes than when growing at the edge or even beyond
its natural range. In Central Europe, a dieback of Norway spruce under drought in
the lowlands beyond its natural range is not surprising. As it is not adapted to the
increasing warmth and drought, the environmental conditions simply develop away
from its ecological niche. Similarly, a growth acceleration of European beech in the
mountain zone of the Alps and in its northern range area is not surprising as here the
environmental conditions may improve the growing conditions of beech as they
change towards its optimal growing conditions. A general increasing trend of basal
area increment was observed for European beech throughout the twentieth century,
across the Italian Peninsula, with the exception of southernmost populations
(Tognetti et al. 2014), which was accompanied by a continuous enhancement in
isotope-derived intrinsic water-use efficiency.
6 Changes of Tree and Stand Growth: Review and Implications 191

In addition to the interplay between initial site conditions and large-scale envi-
ronmental change, regional conditions may influence the growth behaviour.
Examples for this are recovery from litter raking (Gimmi et al. 2013), soil acidifica-
tion by deposition and over-exploitation (Lundström et al. 2003; Hoegberg et al.
2006), lowering of groundwater level (Pretzsch and Kölbel 1988), ozone exposure
(Matyssek et al. 2010) or eutrophication by N deposition (Hofmann et al. 1990;
Pretzsch et al. 2014a).
Yet, shifts of species’ relative abundance in mixed-species forests might strongly
affect regional carbon balances, which warrants the establishment/maintenance of
long-term experiments to measure and monitor growth dynamics and trends in rep-
resentative forests of major biogeographic areas (Pretzsch et al. 2019a). Water
retention experiments, fertilizing experiments and other long-term experiments
with different silvicultural treatments can reveal, for given site conditions and spe-
cies setups, how the effects of environmental changes can be modulated or miti-
gated by management activities. They may contribute to answering whether tree
species mixing, stand density reduction or choice of provenances can mitigate the
effects of climate change, extending the present limit of forest carbon sink. Trees
and forest stands may have the ability to acclimate to stress and to recover.
Morphological acclimation and epigenetic adaptation to stress are far from being
sufficiently understood. Due to the site dependency of growth behaviour, silvicul-
tural recipes that are useful on one site may be inappropriate for counteracting envi-
ronmental changes on other sites.
An important basis for fact-finding on growth reactions to external factors are
long-term observations by, e.g. national inventory plots, environmental monitoring
networks and networks of long-term experiments, covering extended environmental
gradients. For the study at hand, we tapped these databases:
(i) To provide the theoretical basis for better understanding the spatial-temporal
pattern of forest growth behaviour
(ii) To review and give overview of the growth trends and events of tree and stands
in order to derive measures for silvicultural adaptation
(iii) To outline the process of acclimation to stress, adaptation and recovery
(iv) To discuss the implications of the revealed growth behaviour for environmen-
tal monitoring, forest ecology and management

6.2  heoretical Considerations on Growth Changes: Effects

T
of Site Conditions and Species Identity

6.2.1 Standard of Comparison

The unimodal relationship between environmental conditions and growth (or any
other indicator of fitness) is helpful for understanding both the site-specific and non-­
linear growth reactions of monocultures in case of disturbances (Tognetti et al.
2019) and the growth-stabilizing effect of tree species mixtures (Bauhus et al.
192 H. Pretzsch et al.

Fig. 6.1 Growth of various tree species depending on their realised ecological niche and prevail-
ing site conditions in schematic representation. The vertical lines represent specific site conditions:
(a) Unimodal relationship between site conditions and growth of species 1. Any changes of site
conditions (grey bands left and right to site conditions 1, 2 and 3) have stronger non-linear
effects on sites 1 and 3 than on site 2
(b) Species 2 with a niche different from species 1 may compensate for growth losses when site
conditions change from 2 to 3 or from 3 to 4
(c) Tree species interactions in mixed stands can extend (facilitation) or reduce (competition) the
ecological niches of tree species (grey lines)
(d) By combining various tree species with different niches in mixtures (above numbers), stand
growth under climate change (below numbers) may be stabilised

2017). In Figure 6.1a, the environmental conditions change along the x-axis (for
simplicity one dimensional); growth is high in the centre of the real niche and
decreases if temperature, water and nutrient supply become scarce or excessive.
Growth is rather stable under site condition 2 (see middle vertical line), as it is close
to the saddle of the unimodal curve, where temporal variation of the resources sup-
ply (grey bands left and right of site 2) hardly changes the growth. However, on sites
represented by the steep left or right branch of the niche, changes may have strong
non-linear positive or negative effects (grey bands left and right of the sites 1 and 3,
left and right vertical lines). This is why climate warming by 2 °C may have hardly
any effect on temperate sites but strong positive and negative effects on cold and
warm sites, respectively.
6 Changes of Tree and Stand Growth: Review and Implications 193

A combination of two species with differing niches, as shown in Figure 6.1b,

may stabilise stand growth if site conditions change. In this case, species 2 may
compensate for growth losses of species 1, if the site conditions shift from 2 to 3 or
to 4. Combining two species in a stand can also trigger competition reduction and
facilitation (e.g. by hydraulic redistribution through hydraulic lift, atmospheric
nitrogen fixation by symbionts) that may extend their real niche (grey unimodal
curves in the background, Fig. 6.1c) and improve their growth stability (Fig. 6.1c).
Multispecies forest stands, as represented by their niche composition in
Figure 6.1d, may be inferior in growth under stable environmental conditions as
some species are growing suboptimally. However, if the environmental conditions
change, e.g. gradually to a warmer and drier climate, or episodically caused by
drought years, this may change. In this case an assemblage of various tree species
with different niches can mean a risk distribution and stabilisation of stand produc-
tivity, although one or the other tree species may decline or even drop out.

6.2.2 Long- and Short-Term Deviations from Normality

Useful references for detection of any trend or event of the tree or stand develop-
ment are basic growth and yield curves as shown in Figure 6.2a. Under constant
environmental conditions, the growth has a unimodal development over time, and
the yield, as the integral of the growth, has a sigmoid shape. With progressing size,
tree and stand growth declines, and the yield curve has an asymptotic course.
Marziliano et al. (2019) observed that the increase in tree size, more than senes-
cence, explained the reduction in height increment in older silver fir trees. Certainly,
the growth and yield curves of diameter, basal area and volume differ in rhythm
(slope, point of inflexion, turning point), and they are also species specific and
depend, among others, on site conditions, but the unimodal pattern and sigmoid
shape remain as their general characteristics and can be used as reference for detect-
ing abnormal developments in case of environmental changes.
Based on the unimodal growth curve (black) as a standard of comparison, any
acceleration or deceleration of growth by environmental changes (Fig. 6.2b) with
improved growing conditions may be interrupted by episodic stress events
(Fig. 6.2c), or normal growing conditions terminated by stress events with short- or
long-lasting growth reduction (Fig. 6.2d) can be revealed and quantified.

6.3 Empirical Evidence of Growth Trends and Events

6.3.1 Overarching Growth Trends in the Lowlands of Europe

Shortly after the acid rain phenomena from 1970 to 1980, Kenk et al. (1991) detected
a site index improvement by up to 7 metres of dominant height at age 100 according
to the yield table by Assmann and Franz (1963) for Norway spruce stands on poor
194 H. Pretzsch et al.

Fig. 6.2 Modification of the normal unimodal growth curve (black) of forest trees or even-aged
forest stands by changing environmental conditions. (a) Unimodal curve of tree and stand growth
(integrated yield in grey) under normal, constant and undisturbed growing conditions. (b)
Acceleration (green) or deceleration (red) of growth caused by environmental changes. (c)
Improved growing conditions, interrupted by episodic stress events (green). (d) Normal growing
conditions, halted by stress events with subsequent low (red) and high (green) resilient reaction

and medium sites. Among others, Spiecker et al. (1996), Bontemps et al. (2012),
Kauppi et al. (2014) and Pretzsch et al. (2014a) revealed that forest stand growth
dynamics in most parts of Europe have strongly accelerated in recent decades.
Pretzsch et al. (2019a) compiled a data set of several hundred long-term experi-
ments from Austria, Denmark, France, Germany, Great Britain, Poland, Spain,
Sweden and Switzerland for closer analyses of the current growth trends in European
lowlands. Figure 6.3 shows the total stand volume production of 577 fully stocked
and unthinned or just moderately thinned long-term experiments of Norway spruce,
Scots pine, European beech and common and sessile oak in comparison to the yield
tables for Norway spruce by Wiedemann (1936/1942), Scots pine by Wiedemann
(1943), European beech by Schober (1967) and sessile oak by Jüttner (1955). The
yield classes I and IV of these tables represent the development of the respective
species growing in the early twentieth century under optimal and poor conditions,
respectively.
6 Changes of Tree and Stand Growth: Review and Implications 195

Fig. 6.3 Development of the total volume production on long-term experiments of (a) Norway
spruce (n = 202), (b) Scots pine (n = 189), (c) European beech (n = 97) and (d) sessile oak (n = 89)
in Europe established between 1848 and 2010, according to Pretzsch et al. (2019a). The observed
trajectories strongly exceed the spectrum of common yield tables (yield classes I and IV) for
Norway spruce by Wiedemann (1936/1942), Scots pine by Wiedemann (1943), European beech by
Schober (1967) and sessile oak by Jüttner (1955). The German experimental plots were provided
by the Forest Research Station of Baden-Württemberg (BW), the Research Station of Lower
Saxony in Goettingen (GOE) and the Chair for Forest Growth and Yield Science at TUM in
München (MUE)

The same data set also revealed significant relocations of the age trajectories of
total stand volume production and related variables when considering the year of
stand establishment. Figure 6.4 shows, representatively for Scots pine, how the total
stand volume production, standing volume and absolute and relative cumulative
volume, achieved at a given age, changed during the last 150 years. Similar trends
were found for other major species. The total stand production and standing stock
in a mature Scots pine stand is reached 50 years earlier today than for stands that
196 H. Pretzsch et al.

Fig. 6.4 Change of (a) total stand volume production, (b) standing volume, (c) cumulative
removed volume and (d) percentage of removed (mortality, thinning) volume of fully stocked,
unthinned or just moderately thinned long-term experiments of Scots pine established in 1860,
1900 and 1940, according to Pretzsch et al. (2019a)

were established 100 years earlier. As a consequence, the intermediate yield (i.e. the
cumulative removed volume) is 200 m3 ha−1 at the age of 75 years at present, while
it was just 75 m3 ha−1 for stands that were established 100 years earlier. This reflects
an increase by 150%. The cumulative removed volume is the total production minus
standing volume; the percentage of removed volume results from (total produc-
tion − standing volume)/total production × 100.
For a subsample of the plots in Germany, Pretzsch et al. (2018) showed that
wood density decreased by 8–12% since 1900. Even if stand and trees grew much
faster in terms of wood volume, stand biomass increment increased by 9–24 per-
centage points less compared to volume increment. This does not at all cancel the
remarkable volume growth acceleration in the past 100 years, but it slightly reduces
the findings based on stand volume records. The decreasing wood density goes
6 Changes of Tree and Stand Growth: Review and Implications 197

along with an increased early wood fraction and points to the observed extension of
the growing season and fertilisation effect of dry deposition as the main causes of
the detected changes in growth trends.
The growth trends shown in Figures 6.3 and 6.4 indicate changes in growth con-
ditions in terms of rising temperature, extension of the growing season, rising atmo-
spheric CO2 level, nitrogen deposition and abandonment of nutrient-exporting
treatments such as litter raking (Pretzsch et al. 2014a). In other regions, environ-
mental changes can be, of course, also detrimental for growth rates and slow down
stand dynamics. A recent study about tree growth in forests and urban areas revealed
that the beneficial effects of climate change can turn into growth decrease and losses
in regions with limited water and nutrient availability (Pretzsch et al. 2017). Again,
without unmanaged long-term experiments, the knowledge of growth trends and
their causes would be strongly biased or blurred. Nevertheless, Keenan et al. (2016),
using global carbon budget estimates, ground, atmospheric and satellite observa-
tions, and multiple global vegetation models, reported increases in the terrestrial
sink during the past decade. This was associated with the effects of rising atmo-
spheric CO2, though many other factors may influence the carbon cycle at the local
scale. Indeed, partitioning of photosynthates can be largely influenced by environ-
mental factors, stand age and forest management.

6.3.2 Growth Trends in High-Elevation Forest Ecosystems

High-altitude forest ecosystems often provide invaluable ecosystem services, e.g.

protection against avalanches, landslides, rockfall or flooding (Bebi et al. 2001).
Due to their prevailing limitation by low temperatures and short growing season,
especially mixed mountain forest ecosystems at higher elevations are expected to be
strongly affected by climate warming (Piao et al. 2011; Vayreda et al. 2012; Ruiz-­
Benito et al. 2014) similarly to forest ecosystems in the northern latitudes.
Knowledge of any growth trends or structural changes of mountain forests may
enable forest management to adapt and stabilise these stands and thus avoid decline
of productivity and other forest ecosystem services.
Hilmers et al. (2019) and Pretzsch et al. (2020b) showed, for mixed mountain
forest of Norway spruce (Picea abies (L.) Karst.), silver fir (Abies alba Mill.) and
European beech (Fagus sylvatica L.), how environmental changes can modify the
growth trend of tree species, depending on the altitude. The revealed altitudinal-­
related growth trends were found in European mixed mountain forests but probably
show reaction patterns that may be characteristic for forest ecosystems at higher
elevations under environmental change.
Pretzsch et al. (2020b) sampled increment cores from 1721 Norway spruces,
silver firs and European beeches on 28 long-term experimental plots in mixed-­
mountain forest, in seven European countries from Bulgaria to Switzerland. The
plots were located between 621 and 1569 m a.s.l., having an annual temperature
range between 2.9 and 8.2 °C and an annual precipitation of 794–2767 mm yr−1.
198 H. Pretzsch et al.

Fig. 6.5 Overview of the changes in the level and steepness of the growth curves of Norway
spruce, silver fir and European beech during the last three centuries, according to Pretzsch et al.
(2020b). For all the three species, the steepness of the curve increased the strongest in the case of
European beech and silver fir and the least in the case of Norway spruce

The tree ring series revealed an increase of both the level and the slope of the
diameter-­age relationship for all the three species during the last 300 years, as
shown in Figure 6.5. The trend of steepening the diameter-age relationship from
year 1700 to the present is strongest for silver fir and European beech and the least
pronounced for Norway spruce. In the past (1700), the size growth of Norway
spruce was far ahead of silver fir and European beech, whereas, in the two recent
centuries, the slopes of the growth curves of silver fir and European beech increased,
indicating a trend to a more similar growth vigour.
For Norway spruce, the change in growth trend was not dependent on altitude
(Fig. 6.6a), but, for silver fir (not shown) and especially for European beech
(Fig. 6.6b), we found an altitude-dependent behaviour. In the past (DBH-year 1700),
the growth of silver fir and European beech was the highest at low altitudes and the
lowest at high elevations. Both species changed this pattern from 1700 until present.
Trees with DBH-year 1900 are growing better at high elevations and less at lower
altitudes.
This spatio-temporal pattern suggests temperature increase as the main factor for
significant changes in the growth and interspecific competition at the expense of
Norway spruce in mixed mountain forests. The long-term growth trajectories of
Norway spruce in relation to silver fir and European beech hint at a relative advan-
tage of fir and beech at the expense of spruce. The relative inferior growth trend of
spruce in relation to fir and beech corresponds to a loss of fitness. On the long run,
6 Changes of Tree and Stand Growth: Review and Implications 199

Fig. 6.6 Altitude-dependent changes of the growth of (a) Norway spruce and European beech (b)
in the last 300 years, according to Pretzsch et al. (2020b). (a) In the case of Norway spruce, the
growth was the highest at lower elevations and decreased with altitude in the past (DBH-year 1700
and 1800) as well as at present. (b) In the case of European beech, the growth was also the highest
at lower elevations and decreased with altitude in the past. During the last 200 years, the relation-
ships turned to a superior growth at higher altitudes compared to lower elevations

this may reduce the competitive strength and success of Norway spruce in stand
development and natural regeneration.
Similar altitude-dependent temporal changes were found when analysing intra-
specific synchrony in tree growth response to inter-annual fluctuations during the
last century (del Río et al. 2020). Synchrony within beeches decreased with climate
warming at high altitudes, reflecting a lower dependency of climate conditions in
the last decades. By contrast, Norway spruce synchrony maintained constant at high
200 H. Pretzsch et al.

altitudes but increased remarkably at lower altitudes, indicating that climate warm-
ing is exerting a stronger control on spruce growth at these altitudes.
At lower altitudes, Norway spruce is endangered especially by drought stress and
subsequent bark beetle infestation; at higher altitudes, it is impaired by the growth
acceleration of competing silver fir and European beech in mixed mountain stands.
The climate change-induced promotion of silver fir and European beech will, in
absence of silvicultural activities, gradually replace the role that Norway spruce had
in the past in the high-montane and subalpine zone. At lower altitudes, Norway
spruce will be limited to cold sites in hollow relief where silver fir and European
beech, but not Norway spruce, suffer from late frost.
Regardless of the competitive ability of Norway spruce under increasing distur-
bance regimes, a decrease in wood quality can be expected for this species in the
years to come (e.g. wood density). Being the most relevant timber for softwood
products in Central and Northern Europe, Norway spruce calls for new silvicultural
strategies addressing climate change issues, which include the right choice of prov-
enances and degree of stand density regulation.

6.3.3 Stress Events and Low-Growth Years

Many studies use the long-term course of the annual stem diameter growth of trees
for analysing low-growth years caused by stress events such as drought, late frost or
insect injury (Schweingruber 2012). The studies are often based on dominant trees,
in order to keep the effects of competition by neighbours as small as possible
(Pretzsch et al. 2020a). The normal level of the sampled individuals is assumed as
standard for comparison, and any slump of growth in low-growth years is quantified
by the ratio between the reduced and normal level. This results in information about
relative growth losses. Zang et al. (2011) revealed, by this method, that the ranking
between primary tree species in Central Europe regarding growth losses is Norway
spruce, silver fir, Douglas fir, European beech, Scots pine and sessile oak. By com-
paring the growth reactions in the given years in monospecific stands with neigh-
bouring mixed-species ones, Pretzsch et al. (2013), Grossiord et al. (2014), Thurm
et al. (2016) and Ammer (2019) could show that, in some cases, interspecific neigh-
bourhood can reduce species-specific drought stress effects.
Drought events in Central Europe, among others, in 1976, 2003 and 2015, trig-
gered many studies about the effects of episodic drought on the growth and mortal-
ity of forest tree species (Ciais et al. 2005; Bréda et al. 2006; Allen et al. 2015).
These findings suggest that tree species especially growing at or beyond the border
of their natural range, such as Norway spruce or European larch (Larix decidua
Mill.) in Central Europe, can show severe growth losses and mortality (Kölling et al.
2009; Lévesque et al. 2013). Scots pine (Pinus sylvestris L.) and sessile oak (Quercus
petraea L.) often serve as examples for rather drought-tolerant species (Walentowski
et al. 2007; Zang et al. 2011, 2012), more suitable for forestry in Central Europe
under climate change towards warm and dry conditions. In order to mitigate drought,
6 Changes of Tree and Stand Growth: Review and Implications 201

silviculture aims at the selection of well acclimated species and provenances

(Atzmon et al. 2004; Arend et al. 2011; Zang et al. 2011), at reducing stand density
(D’Amato et al. 2013; Sohn et al. 2016; Bottero et al. 2017), at modifying the kind
of thinning (Rodríguez-Calcerrada et al. 2011; Gebhardt et al. 2014; Pretzsch et al.
2018) or at favouring tree species mixing. The latter, however, is not rated effective
for drought mitigation in general (Grossiord 2019). Indeed, in forest stands, species
diversity is not always positively related to drought resistance. Conte et al. (2018)
observed that stem radial growth and isotope-derived intrinsic water-­use efficiency
were generally higher in pure than in mixed stands of European beech and Scots pine.
Exemplarily, Figure 6.7 shows species-specific stress reactions caused by the
drought year 1976 quantified in relation to the mean growth level in the 3-year-­
period 1973–1975 before the drought stress (reference line = 1.0). The study was
based on tree ring measurement on cores from increment chronologies from 559
trees of Norway spruce, European beech and sessile oak in South Germany, with
half of them sampled in monospecific stands and the other half in mixed stands
(Pretzsch et al. 2013).
For quantifying the resistance, recovery and resilience, indices introduced by
Lloret et al. (2011) were applied, allowing for retracing the tree’s growth reaction on
the episodic drought stress in the years 1976 and 2003. The following general reac-
tion patterns, visualised in Figure 6.7, were found. In pure stands, spruce had the
lowest resistance but the best recovery. Oak and beech were more resistant but recov-
ered less pronounced and thus were less resilient. In mixture, spruce and oak per-
formed like in pure stands, but beech was significantly more resistant and resilient
than in monocultures. Especially when mixed with oak, beech was facilitated. We
hypothesise that the revealed water stress release of beech emerges in mixture because
of the asynchronous stress reaction pattern of beech and oak and a facilitation of
beech by hydraulic lift of water by oak. A potential positive contribution of species

Fig. 6.7 Species-specific stress reactions caused by the drought year 1976 shown in relation to the
mean growth level in the 3-year-period 1973–1975 before the drought stress (reference line = 1.0)
(according to Pretzsch et al. 2013). (a) Norway spruce, European beech and sessile oak in pure
stands. (b) European beech in pure and mixed stands. (c) European beech and sessile in pure and
mixed stands. The courses represent the growth in the dry year 1976 and in the recovery period
(periodical mean of 1977–1979) in relationship to the growth in the reference period (periodical
mean of 1973–1975)
202 H. Pretzsch et al.

Fig. 6.8 Mean (± SE) annual basal area increment (± SE) of all (a) Norway spruce (n = 268) and
(b) European beech (n = 141) from 1998 to 2018. Trees facing rainfall exclusion since 2014 were
excluded (tree numbers refer to the year 2003)

with a deep root system (e.g. oak) towards those with shallow roots (e.g. beech) can
be hypothesised in mixed forests subjected to drought stress (Zapater et al. 2011).
As another example, we show the growth reactions of Norway spruce and
European beech to natural episodic and experimentally extended drought in mature
monospecific and mixed-species stands of Norway spruce and European beech in
the Kranzberg Forest (Fig. 6.8). From the annual diameter growth records since
1998 based on girth tapes, Norway spruce and European beech both reflect the epi-
sodic drought events in 2003 and 2015. The courses of the annual basal area growth
(± SE) from 1998 to 2018 is visualised in Figure 6.8. The long-term trend in this
period is slightly decreasing for Norway spruce and rather parallel to the x-axis for
European beech. This long-term trend, however, is interrupted by slumps of the
annual growth in 2003 and 2015, especially in the case of Norway spruce. European
beech was much more resistant to the drought years. Norway spruce shows a strong
growth reduction in the drought years 2003 and 2015, while European beech shows
just a slight growth reduction in 2003 and even an increase in 2015. It is obvious that
the growth of Norway spruce is severely reduced by about 50–60% compared to the
growth before the drought period. European beech trees do not exceed half of the
growth losses and react much less to drought.
In order to show intra- and inter-species-specific response patterns to drought,
we also compared the behaviour in the drought years 2003 and 2015 with the 3-year
periods before and after the events (Fig. 6.9). Here, we visualise the results just for
2003, as they were similar concerning the relationships between the species and
concerning the intra- and interspecific differences, but more pronounced than in
2015. Figure 6.9a underpins the much stronger effect of drought on the growth of
Norway spruce compared with European beech, in general, without considering
their intra- and interspecific neighbourhoods.
Interestingly, Norway spruce suffered 10–20% less under drought when growing
in the neighbourhood of European beech (see Fig. 6.9b, mean and SE lines for sb).
The growth losses were stronger in the intraspecific neighbourhood of Norway
spruce in 2003. The tree growth of the group with intraspecific competition (group
6 Changes of Tree and Stand Growth: Review and Implications 203

Fig. 6.9 Visualisation of the growth resistance and resilience to the 2003 drought event based on
the annual basal area increment (± SE) (according to Pretzsch et al. 2020a). The pre-drought
growth in the period 2000–2002 is set to 1.0 (1.0 line). The growth in the drought year 2003 and in
the post-drought period 2004–2006 was sketched in relation to this reference level. (a) On average,
the growth of Norway spruce dropped steeply and recovered slowly; European beech was hardly
affected by the 2003 drought. (b) When Norway spruce grew in the interspecific neighbourhood
with European beech (sb, broken lines), it was less affected by drought compared with intraspecific
constellations (ss). (c) When European beech grew in the interspecific neighbourhood with Norway
spruce (bs, broken lines), it was more affected by drought compared with intraspecific constella-
tions (bb)

ss) was significantly lower than the growth in the group with interspecific competi-
tion (group sb).
European beech behaved differently (Fig. 6.9c). European beech growth declined
significantly stronger under drought in the interspecific neighbourhood but recov-
ered quickly (see Fig. 6.9c, mean and SE lines for bs). In contrast, European beech
growing in the neighbourhood of beech trees was much less affected by drought
(mean and SE lines for bb). In dry years, Norway spruce seemingly benefits from
interspecific neighbourhoods. Similar growth-stabilizing interspecific interactions
were reported by del Río et al. (2014).
The results of such studies indicate growth reactions of selected sampled trees.
The diagnosis of growth reactions and growth losses at the stand level can show a
different picture. Small trees may react differently to stress and partially compen-
sate the growth losses of tall trees in low-growth years (Pretzsch et al. 2018). In
addition to the growth of individual trees, stand growth is determined by stand den-
sity, mortality and existing natural regeneration that can strongly modify the growth
under drought stress (Allen et al. 2015; Pretzsch et al. 2015; Sohn et al. 2016).
Stress may also modify height growth, form factor or allometry between root and
shoot; all of these aspects are not considered by relative growth analyses based on
increment cores of individual trees at breast height (Rötzer et al. 2017).
In summary, the dendrochronological analyses of growth oscillation in drought
years may serve as a bioindication and evidence of stress by environmental factors
(Dobbertin 2005; Allen et al. 2010), whereas the growth analyses at the stand level
provide integrated information of growth reactions that are relevant on ecosystem
204 H. Pretzsch et al.

level and for management and planning purposes. Both are useful indicators of
climate-­smart forestry.

6.3.4 Vulnerability Related to High Productivity Level

Compared to the past, forest stand growth has accelerated during the last decades in
many areas of Europe (Spiecker et al. 1996; Kauppi et al. 2014; Pretzsch et al.
2014a). Although proceeding on a raised growth level, stands are encountering a
series of drought years. But, even if tree and stand growth by this may be reduced
below the present level, they still exceed the past level, represented, for instance, by
the yield tables. The negative deflection from the currently increased level is natu-
rally interpreted as stress exposure. However, at the same time, the absolute level is
still higher than in the past. This may evoke contradictory assessments regarding
forest health and vitality.
For showing this concurrency of long-term upward trend and episodic stress
events, we present results from an ongoing study of monospecific and mixed-­species
stands (triplets) of Norway spruce, Scots pine, European beech and sessile oak
across Europe (Pretzsch et al. 2020c). Figure 6.10 depicts the stand volume growth
of monospecific stands of the respective species in Germany in comparison with the
corresponding yield tables by Wiedemann (1943), Jüttner (1955), Assmann and
Franz (1963) and Schober (1967).
In the case of Norway spruce (Fig. 6.10a), the observed (oscillating course) and
expected growths (unimodal growth curves) were the closest to each other. In the
case of European beech (Fig. 6.10b, d), the observed growth strongly deviated from
the yield table predictions. Most of the stands showed an increased growth level and
an increasing growth trend since one or two decades. Recent drought years just
cause an oscillation on a luxury hypertrophy level and only occasionally cause a
slump of the growth trajectories below the yield tables used as standard for compari-
son. Also, Scots pine (Fig. 6.10c, e) strongly exceeded the level of the yield tables
and strongly increased in growth. In the last few years, the upward trend is, from
time to time, interrupted by low-growth years. The most positive deviations from
the yield tables and upward trends were found for the stand growth of sessile oak
(Fig. 6.10f); the drought years in 2003 and 2015 had hardly any effects on the
annual course of growth of sessile oak. The growth of the mixed-species stands also
proceeded above the yield tables and showed an increasing trend (not shown). But,
compared with the monospecific stands, the inter-annual oscillation in mixed-­
species stands was lower. So, species interactions not only stabilise growth at tree
level (Fig. 6.9) but also contribute to greater temporal stability at the stand level (del
Rio et al. 2017).
On average, the growth of the analysed stands of spruce, pine, beech and oak was
48% above the historic level represented by the yield tables (Pretzsch et al. 2020c);
this reflects a considerable change of forest growth and potential for wood utilisa-
tion compared to the previous century. Growth loss due to drought was highest in
6 Changes of Tree and Stand Growth: Review and Implications 205

Fig. 6.10 Courses of the absolute annual volume growth (merchantable wood with diameter 7 cm
at the smaller end including bark) of monospecific stands of the triplets of (a and b) Norway spruce
and European beech, (c and d) Scots pine and European beech and (e and f) Scots pine and sessile
oak in the period 1997–2018 compared with the courses of the yield tables for moderate thinning
by Wiedemann (1943), Jüttner (1955), Assmann and Franz (1963) and Schober (1967)
206 H. Pretzsch et al.

Fig. 6.11 Growth of traditional yield table vs. current reality in schematic (a) absolute and (b)
relative representation. Growth predicted by yield tables (yield classes I and IV), current growth
trend and low-growth events caused by drought years (growth decrease marked by filled circle). In
the marked drought year, the stand growth would lie under the level of the upward trend but still
above the level of the yield tables (1.0 line)

Norway spruce stands, lowest in sessile oak stands and medium in European beech
and Scots pine stands. Mixed-species stands performed slightly better in drought
years but not significantly. The current growth courses of the analysed stands can be
understood as a new standard. In the recent drought years, the stand growth was
reduced on average by 28%; in the most favoured or unfavoured cases, it was
reduced by 19% and 37%, respectively, referenced at the new standard.
The aggregated picture applying for sites with medium to good growing condi-
tions, i.e. for many sites in the Central European lowlands, is given in Figure 6.11.
Figure 6.11a shows the course of growth predicted by the yield tables (curves for
yield classes I and IV), the current annual course of growth (sawtooth curve) and the
low-growth collapse in a drought year (minimum value marked by a dot) in sche-
matic representation at the absolute scale. Figure 6.11b shows the same relation-
ships at a relative scale. Both graphs reveal that the growth in the drought year
declines below the level of the current upward growth trend, but it is still above the
level of the yield table. This means that the growth is reduced by drought stress but
may still lie considerably above the growth predicted by the yield tables.

6.4 Acclimation, Adaptation and Recovery

6.4.1 Acclimation

Trees and stands may physiologically or morphologically acclimate to drought

stress (Cinnirella et al. 2002; Lapenis et al. 2005; Reich et al. 2016). Here, we show
an example of trees’ acclimation to extended drought stress (Fig. 6.12). In a rainfall
exclusion experiment, Goisser et al. (2016) and Pretzsch et al. (2020a) analysed the
6 Changes of Tree and Stand Growth: Review and Implications 207

Fig. 6.12 Visualisation of the resistance to the 2014–2018 rainfall exclusion based on the single-­
tree annual basal area increment (± SE). Pre-drought growth in the period 2011–2013 is set to 1.0
(1.0 line, solid black); the growth during years of the rainfall exclusion is related to the pre-drought
level. (a) On average, growth of Norway spruce dropped steeply and recovered slowly; growth of
European beech dropped less and even increased above the level of the pre-drought period after
4 years. (b) When Norway spruce grew in the interspecific neighbourhood with European beech
(sb, broken lines), it was less affected (20–30%) during the rainfall exclusion in the first years and
recovered remarkably compared with spruce trees in intraspecific constellations (ss). (c) When
European beech grew in the interspecific neighbourhood with Norway spruce (bs, broken lines), it
performed better in the first years of drought but then relapsed the growth of beech trees in intra-
specific neighbourhoods (bb)

courses of the annual basal area increments within a 5-year period of water exclu-
sion over all summer periods. Figure 6.12a shows the behaviour of Norway spruce
and European beech in monospecific and mixed stands. Both species strongly
reduced their growth in the first year after the rainfall exclusion. Norway spruce
continued to decrease but stabilised in 2016–2018. European beech stabilised even
earlier and rearrived at the base level in 2018. Since 2015, Norway spruce grew on
average significantly less than European beech on the treatment plots. This under-
pins the species-specific drought tolerance reviewed by Niinemets and Valladares
(2006). The asymptotic growth trend of Norway spruce and the upward growth
trend of European beech after multi-year drought can be interpreted as acclimation.
Especially in the case of Norway spruce, the neighbourhood seems to matter for
the growth behaviour imposed by experimentally extended drought stress
(Fig. 6.12b). Norway spruce growing in the neighbourhood of spruce trees dis-
played a much stronger growth decrease than those in vicinity to European beech.
In the case of European beech, both trees in inter- and intraspecific neighbourhood
behave similarly in the first 3 years (Fig. 6.13c). Since 2016, we found significant
differences between the two groups, i.e. beech trees in intraspecific competition
outgrew those neighboured by spruce trees.
In summary, the experimentally induced drought from 2014 to 2018 caused a
strong growth reduction in the first year, followed by a slight acclimation to the dry
conditions. European beech acclimatised and recovered better than Norway spruce;
Norway spruce acclimatised better in the neighbourhood of beech trees. For Norway
208 H. Pretzsch et al.

spruce tree mortality was fivefold, while for European beech it is similar to the long-­
term mortality rate on the untreated plots.

6.4.2 Adaptation

The intraspecific genotypic and phenotypic variability plays an important role in

terms of adaptation. Figure 6.13a illustrates how a high genetic diversity can con-
tribute to adaptation, as each genotype shows a distinct optimum curve to climate
conditions.
Similarly, there can be substantial differences in expression of the optimum
curve among different provenances of a species (Benito Garzón et al. 2011) through
adaptation to varying climate conditions (Aitken et al. 2008).
The Scots pine provenance trial Schwabach 304, established in 1927, provides
an example for provenance-dependent behaviour of growth (Fig. 6.14a). This exper-
iment revealed that the production of the provenance “Bamberg” was inferior until
an age of 50 years compared to the provenances “Schwabach” and “Unterfranken”.
The provenance Bamberg seems to be better suitable for the changing growing con-
ditions since the last century. However, it turned to the most productive provenance,
possibly indicating higher adaptation to the specific site conditions. In the case of
the Douglas fir provenance trial Kösching 95 (established in 1961, first survey 1961,
last survey 2015), all provenances performed very similarly during the juvenile
stand development. With increasing age, however, the difference in total stand
growth between the weakest and strongest performing provenance becomes a
remarkable 500 m3 ha−1 (Fig. 6.14b). This trial also includes plots of Norway spruce,

Fig. 6.13 Growth of different genotypes depending on their real ecological niche in schematic
representation (niche reduced to one axis for simplifying the representation). (a) Unimodal rela-
tionship between site conditions for different genotypes. A greater genetic diversity can contribute
to a better adaptation to climate change. (b) Growth of two species with different phenotypic
plasticity. A higher phenotypic plasticity can reduce the impact of climate change and favour
“adaptation” in a broad sense of the term. (c) Shift in species growth response along the ecological
niche caused by epigenetic preconditions. Different growth responses to environmental conditions
by epigenetic changes can be inherited in the new generation and contribute to tree species
adaptation
6 Changes of Tree and Stand Growth: Review and Implications 209

Fig. 6.14 Long-term observations can reveal changes in the performance ranking of different spe-
cies’ provenances as well as deviations of their growth and yield characteristics from the prediction
by yield tables. (a) On the Scots pine provenance trial Schwabach 304, initially low-performing
provenances finally have superior total production. After 20 years, all the three provenances sur-
pass the level of the yield table. (b) Douglas fir provenance trial Kösching 95, where some Douglas
fir provenances significantly excel the other and the total production of the Norway spruce refer-
ence plots in advanced age. The yield tables for Scots pine by Wiedemann (1943, yield classes I–
III) and for Douglas fir by Bergel (1985, yield class I) and Norway spruce by Assmann and Franz
(1963, O 40) were used as reference

which performs similarly to Douglas fir initially but lags behind most of the Douglas
fir provenances at advanced ages. The deviations from the reference yield tables are
negligible in the beginning but accumulate strongly with increasing stand age.
The long-term changes in ranking and trend, depicted in Figure 6.14, underpin
that the choice of silvicultural options (e.g. provenance, thinning and tree species)
should not be based just on early tests but on long-term observation. Real-time
series of observations cannot simply be replaced by artificial time series, which try
to derive the development over age by measuring and combining stands of different
ages on the same site.
Generally, a higher genetic diversity implies more possibilities of adaptation to
changing conditions, but species adaptation through selection during the regenera-
tion phase is a long-lasting process. Therefore, other aspects, such as species phe-
notypic plasticity to environmental conditions or the above-mentioned ability of
acclimation, are important for adaptation to climate change, in a broader perception
of the term “adaptation”. More plastic species or provenances are expected to resist
better to climate changes in large parts of their distribution areas (Fig. 6.13b).
However, plasticity is not always linked to genetic diversity (Mutke et al. 2010);
therefore, it does not always guarantee good performance.
Epigenetic changes can also have important effects on species adaptability
(Fig. 6.13c). Current research scrutinises to what extent climate change can cause
heritable epigenetic changes and adaptation of trees beyond the genetic adaptation
by sexual reproduction (Franks Hoffmann 2012; Verhoeven et al. 2016). That
210 H. Pretzsch et al.

environmentally induced epigenetic changes may be inherited by future generations

is of special relevance for long-lived organisms, such as trees (Bräutigam et al.
2013). Rico et al. (2014) and Shanker et al. (2014) report on epigenetic changes
under drought stress, and Bossdorf et al. (2008) discussed how to consider such
high important aspects in future experiments in ecology.

6.4.3 Recovery

The recovery capacity after cessation of disturbance is an important characteristic

of tree growth to better understand growth reaction patterns under changing envi-
ronment. Beyond the already revised growth response to extreme annual events
through the analysis of resistance, recovery and resilience indices by Lloret et al.
(2011), here, we present two examples of growth response to longer disturbances,
such as SO2 pollution and O3 exposure, and the recovery afterwards.
Growth Reactions of Norway Spruce and Silver Fir to SO2 Pollutants The stem
diameter growth trajectories of Norway spruce and silver fir in Fig. 6.15 show the
stress reaction and recovery caused by SO2 pollutants (1950–1985) and dry years
(1976, 2003). The trajectories are based on 118 trees per species sampled by incre-
ment boring in 22 mixed-species stands across Germany. The stands cover a wide

Fig. 6.15 Average course of the stem diameter increment (id mm yr-1) of about 100-year-old silver
fir (n = 118) and Norway spruce (n = 118), in Bavaria and Lower Saxony. The graph shows the
period 1950–1985 with increased SO2 pollution, the species-specific growth trend after 1985 with
reduced SO2 pollution and the drought years 1976 and 2003 (Uhl et al. 2013)
6 Changes of Tree and Stand Growth: Review and Implications 211

range of site conditions, within and beyond the natural distribution of both species,
in Bavaria and Lower Saxony. In Bavaria, the tree age of Norway spruce and silver
fir was 110 and 114 years, respectively, and in Lower Saxony it was 72 and 88 years,
respectively. Since 1900, both species first show a rather similar and steady level of
growth. Silver fir then shows a growth depression from 1950 to 1980 and a growth
recovery afterwards. The growth of Norway spruce is superior to silver fir in the
period 1950–1980 and turns to be inferior after 1980.
The growth of silver fir was strongly reduced by the high SO2 pollutants in the
period 1950–1980 (Uhl et al. 2013). After reduction of the SO2 load in the atmo-
sphere by installation of filters in power stations (Joos 2006; Elling et al. 2009), the
stem diameter growth of silver fir increased continuously in the last 20–30 years
despite the advancing age. The finding that the growth of silver fir dropped less and
recovered faster than the growth of Norway spruce in drought years means that the
vitality changed in favour of silver fir in the last decades. Consequently, in the very
widespread mixed-species stands of Norway spruce, silver fir and European beech
in European mountain regions, silver fir gained in competitive strength and produc-
tivity compared to the other species (Pretzsch et al. 2015; Bosela et al. 2018; Hilmers
et al. 2019).
Growth Reactions to Chronic O3 Stress and Recovery After Temporal O3
Exposure Increased O3 concentration in the air can have significantly detrimental
effects on the health and growth of trees in urban areas and forests (Matyssek und
Sandermann 2003). O3 fumigation experiments show that ozone exposure can
reduce the photosynthesis (Botkin et al. 1972; Karnosky et al. 2007; Wittig et al.
2009), the carbon assimilation (Chappelka and Samuelson 1998; Sitch et al. 2007),
the growth (Wipfler et al. 2005; Pretzsch und Schütze 2018) and allometry of trees
(Dickson et al. 2001; Grantz et al. 2006; Pretzsch et al. 2010). The effect of O3 on
growth can be analysed by tree ring analyses and comparison of exposed trees with
controls (Pretzsch und Schütze 2018).
The free-air ozone fumigation experiment in the Kranzberg Forest showed how
80-year-old Norway spruce and European beech trees reacted to double-ambient O3
exposure (2000–2007) and also how they behaved after the stop of the temporary
exposure (Matyssek et al. 2010; Häberle et al. 2012). During the phase with double-­
ambient O3 exposure (2 × O3), the annual basal area increments of Norway spruce
and European beech decreased by 24% and 32%, respectively (Fig. 6.16). In the
period 2008–2016, after the stop of the exposure, both species recovered. The basal
area increments of the previously O3-fumigated trees recovered and raised by 14%
and 24% above the level of the control trees (1 × O3). So, they nearly caught up with
the control trees regarding diameter growth. However, the resistance and resilience
of the previously fumigated trees to drought and late frost were lower compared with
the control trees. This underlines that measures for pollution control may result in a
quick recovery from chronic O3 stress and emphasises the relevance of O3 control
for the health and the carbon sink function of trees and forests (Matyssek et al. 2010).
212 H. Pretzsch et al.

Fig. 6.16 Pattern of growth responses of mature Norway spruce and European beech to temporary
2 × O3 ozone fumigation, according to Pretzsch et al. (2018). Free-air twice-ambient ozone fumi-
gation (2 × O3) reduced diameter increment of mature Norway spruce and European beech within
the 2 × O3 fumigation period from 2000 to 2007. In the post-fumigation period, growth of the
former fumigated trees can recover and even exceed the growth of the control trees. 1 × O3 means
ambient ozone concentration and represents the control tree (black line); 2 × O3 means twice-­
ambient ozone-fumigated trees (red line)

6.5  iscussion: Implications for Environmental Monitoring,

D
Forest Ecology and Management

The reviewed tree and stand growth behaviour by reasons of changing environments
is highly relevant as any change of forest growth and structure has strong impacts
on the provision of ecosystem goods and services. In the following, we discuss the
implications for environmental monitoring, forest ecology and management.

6.5.1 Environmental Monitoring

As tree and stand growth strongly depends on environmental conditions, their

observation delivers a unique contribution to environmental monitoring. Thus, the
tree and stand growth measured on long-term experiments and the tree growth rates
retrospectively derived from increment cores and stem discs can provide evidence
of environmental changes (Schweingruber 2012). Growth records from trees or
stands without active forest management are of special value, as their growth
courses are not superimposed by silvicultural management effects. Long-term
observation helps to quantify the human footprint on nature (Pretzsch et al. 2014a).
6 Changes of Tree and Stand Growth: Review and Implications 213

In temperate Europe, there is a patchwork of forests with positive, neutral and

negative growth reactions to the ongoing change of environmental conditions. In
regions with sufficient water supply, growth strongly accelerated due to favourable
effects of global change, such as nitrogen deposition, extension of the growing sea-
son (McMahon et al. 2010) and rise of the CO2 concentration (Ewald et al. 2004).
In drier regions, forest growth is reduced by drought and accompanied biotic calam-
ities (Martin-StPaul et al. 2013). In fact, many forests show growth slumps in dry
years but are currently growing at higher levels than before, requiring a long-term
reference for impact assessment (Fig. 6.11).
The demonstration of forest growth and decline and the practical consequences
on long-term experiments can motivate measures for environmental care and pollu-
tion control (Matyssek and Sandermann 2003; Uhl et al. 2013). The growth trends
and events on environmental conditions can also be used for assessing the climate
smartness of various forest and stand types and contribute to developing indicators
of stability and climate smartness for monitoring and decision-making purposes
(Bowditch et al. 2020; Santopuoli et al. 2021).

6.5.2 Forest Ecology

The accelerated tree growth and forest aging and the projected sink saturation
require conformance of all associated organisms, including humans. Plants and ani-
mals inhabiting these habitats depend on specific phases in stand development and
structure; faster growth means interference in species living conditions and demands
for higher mobility. In other areas, growth decline may call in question the future of
tree species and the associated plants and animals. In the lowlands, we find horizon-
tal mosaics and, in the mountains, elevation zones with changing growth, changing
competitive strength of the species and a transition to species with ecological niches
better suitable for both the continuously and abruptly changing growth conditions.
If we interpret tree growth trends as an indication of fitness (Dobbertin 2005), we
may consider that some species, such as Norway spruce and Scots pine, especially
when cultivated beyond their natural range in dry lowland areas, lose competitive-
ness. In many cases their growth is still on a rather high level but with a decreasing
tendency, especially caused by drought years that became more frequent in the last
decades.
Long-term observation of tree growth shows that trees can recover from episodic
and chronic stress and that they are also able to acclimatise to a certain degree to,
e.g. drought stress (Pretzsch et al. 2020a). Future research will show the potential of
acclimatisation and of adaptation over successive generations. Natural regeneration
may pave the way to a continuous adaptation by natural selection of the fittest.
Permanent plots surveyed with a combination of high resolution and coarse size will
be required to track forest demography in time and space (e.g. tree mortality, age
structure) and understand scale and pattern of forest development (e.g. species
diversity, vegetation shift).
214 H. Pretzsch et al.

The shown growth trends suggest new experiments in monospecific and mixed-­
species stands, in order to better understand the effects of environmental changes on
forest resilience. Of importance are, e.g. experiments analysing the provenance
selection and adaptation to climate change, the impacts of tree species mixing on
growth stability, the potential for acclimation and adaptation of tree species, the
effects of various silvicultural measures (e.g. thinning), the interactive consequences
of disturbances and the effects of nutrient addition on stress resistance, resilience
and recovery.

6.5.3 Forest Management

Management options and decisions regarding the choice of species and provenance,
stand regeneration, spacing, thinning and rotation length depend on information
about the current tree and stand growth. Susceptibility of presently used species to
biotic or abiotic damages will determine the selection of new provenances and for-
eign species (Atzmon et al. 2004; Arend et al. 2011; Zang et al. 2011). By early
stand regeneration and transition to natural regeneration, the stand acclimation and
adaptation to drought may be promoted (Aasamaa et al. 2004). Spacing and thin-
ning may be intensified in order to use the accelerated growth or to reduce stand
density and drought stress (Sohn et al. 2016). Long-term experiments show, in many
regions, an increase of the maximum stand density; this means that keeping stands
at the traditional density levels may cause severe growth losses as the optimal den-
sity will be under matched. The accelerated growth may suggest a shortening of the
rotation period (Pretzsch et al. 2014a).
All these measures depend on information about growth trends and short-term
growth reactions to adjust forest management and maintain sustainability, in par-
ticular, under changing environmental conditions. More rapid tree growth can result
in earlier harvest threshold diameters and rotation due to increased stand productiv-
ity, which can raise the annual harvest. Recent growth trends allow foresters to
maintain much higher growing stocks. However, strong thinning, which uses past
conditions as a guideline, might reduce stand density, such that the actual growth
potential is not fully realised. In addition, a shortened rotation period can mean
reduced risk in terms of forest damage, including windthrow, bark beetle infestation
and/or snow breakage. On the other hand, at sites with decreasing growth trends and
decay symptoms, urgent adaptation strategies are required, such as species diversi-
fication, density regulation, promotion of natural regeneration or even, in some
cases, species or provenance assisted migration.
For a specific stand mean tree size, standing stock and mortality rate can be
achieved one or more decades earlier. This leaves age-based experience values,
widely used yield tables and other models and many traditional management guide-
lines to become obsolete. Trade-offs exist between shortening rotation period to
maximise carbon sequestration and lengthening tree growth to reduce biodiversity
loss. Growth records may further serve for updating forest growth models not only
6 Changes of Tree and Stand Growth: Review and Implications 215

for monocultures but also for more complex mixed-species stands. Such models
should provide forest management with appropriate growth and yield information
for calculation of the annual sustainable harvest. The underlying experimental and
observation plots may serve as demonstration and training plots on how to manage
forest under environmental changes and as silvicultural assistance to the transition
from high-risk monocultures to close-to-nature forest systems, towards climate-­
smart forestry.

6.6  he Importance of Long-Term Experiments

T
for Fact-Finding

As most of the results presented in this chapter are based on long-term growth and
yield experimental plots, we finally discuss the importance of long-term experi-
ments for fact-finding. The founding fathers of forest yield science, such as
Danckelmann and Schwappach, were convinced that trees and forests require long-­
term observation, and for this purpose they established the first forest research sta-
tions, beginning in 1870 (Ganghofer 1881; Milnik 1999; Landesforstanstalt
Eberswalde 2001). Most of our scientific knowledge of tree and stand dynamics,
and practical decision, is based on long-term experiments. However, long-term
experiments are often under scrutiny, endangered and sacrificed for cost reduction.
The reasons are as follows: forest areas with long-term experiments have to be left
out from regular forest operations, long-term experiments are costly and waiting for
some results more than a couple of years hardly fits into these fast-moving times.
A misleading argument for discontinuance is that forest inventories established
in the last decades at national or enterprise level render long-term experiments
superfluous. Long-term experiments and temporary plots are rather complementary,
more than redundant (Nagel et al. 2012). Long-term experiments provide far-­
backward-­reaching time series of growth and yield for selected sites; they also
include extreme variants, such as unthinned stands or solitary growing conditions,
which are hardly represented by inventories. Unthinned plots can be important as
reference (maximum density, site-specific productivity), and solitary plots are rele-
vant for understanding tree behaviour (aging, wood quality) under strong thinning
and the trade-off between tree and stand level performance. Temporary plots and
forest inventories, in contrast, include mainly routinely managed stands (mean stand
density, common silvicultural treatment) and provide a representative overview of
the current growth behaviour.
Forest inventories can provide large-scale representative data, and the informa-
tion potential can be exploited by big data methods (e.g. Liang et al. 2016). However,
inventories may hardly provide information about the stand history and intermedi-
ate yield; a lack that can be partly remedied by permanent inventory plots, when
they get measured repeatedly and gradually, provide longer time series. Long-term
experiments, in contrast, can reveal the cause-effect relationship of various
216 H. Pretzsch et al.

treatment options on tree and stand behaviour, as they are established under con-
trolled continuous (ceteris paribus) conditions. Inventory plots indicate correlations
but provide no evidence for casualties, as they can vary in many traits beyond the
factor in question. Long-term experiments strive for general rules, laws and under-
standing, while inventory plots aim at regional information for practical purposes.
Simulation models can make use of both long-term plots for developing the thin-
ning, mortality and regeneration algorithms and inventory data for calibration of the
site-growth relationship, model initialisation and risk assessment.
Large-scale and long-term monitoring plots may be useful to characterise eco-
logical processes (succession, outbreak, windthrow, etc.), thus providing models
with data from heterogeneous conditions and including temporal variability of cli-
mate. Monitoring of large plots helps to emerge the effect of processes that are
spatially autocorrelated (dispersal, competition, facilitation or mortality).
Monitoring of permanent plots helps intercept the impact of phenomena that are
temporally discrete (e.g. extreme events). During the careers of research topics,
such as impact of acid rain, nitrogen deposition or climate change or global change
on forests, repeated observation on permanently marked plots provided information
far beyond the purpose they originally were established for. They represent stands
with known history regarding establishment, silvicultural treatment and distur-
bances. They offer time series of stand development for biomonitoring, develop-
ment of silvicultural treatment, modelling and demonstration and training.
Current course of growth may reveal site- and species-specific reaction to vari-
ous disturbances and may contribute to a less emotional but more objectified discus-
sion of the human influence on tree growth and forest dynamics. While the public
debate about forest ecology and health changes its focus typically minute by min-
ute, long-term experiments provide a differentiated and consolidated information
base about where and to what extent not only forest growth is influenced by humans
but also that pollution and climate control can help.

Acknowledgements The authors would like to acknowledge networking support by the COST
(European Cooperation in Science and Technology) Action CLIMO (Climate-Smart Forestry in
Mountain Regions – CA15226), which received funding under the EU Framework Programme for
Research and Innovation HORIZON 2020.
The first author wishes to thank the German Science Foundation (Deutsche
Forschungsgesellschaft) for funding the projects “Structure and dynamics of mixed-species stands
of Scots pine and European beech compared with monospecific stands. Analysis along an ecologi-
cal gradient through Europe” (# DFG PR 292/15-1) and “From near-death back to life: Mixed
stands of spruce and beech under drought stress and stress recovery. From pattern to process” (#
DFG PR 292/22-1). We would also like to thank the Bavarian State Ministry for Environment and
Consumer Protection for funding the project “Pine (Pinus sylvestris) and beech (Fagus sylvatica)
in mixed stands: Suitable partners to ensure productivity on dry sites in times of climate change
(KROOF II)” (# GZ: TKP01KPB-73853) and the Bavarian State Ministry for Nutrition, Agriculture
and Forestry for funding the project “W047” (# GZ: 7831-28160-2018). We further thank the
6 Changes of Tree and Stand Growth: Review and Implications 217

Bayerische Staatsforsten (BaySF) for supporting the establishment of the plots and the Bavarian
State Ministry for Nutrition, Agriculture and Forestry for permanent support of the project W007
“Long-term experimental plots for forest growth and yield research” (# 7831-26625-2020). Thanks
to all the project partners of the mentioned projects for providing data on tree and stand growth
across Europe. Thanks also goes to anonymous reviewers for their constructive criticism.

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