Advanced Review

Historical, ecological, and

governance aspects of intensive
forest biomass harvesting in
Denmark
Inge Stupak* and Karsten Raulund-Rasmussen

Intensive forest harvesting has increased in Fennoscandia over the last few dec-
ades. Similar developments may follow throughout Europe as renewable energy
replaces fossil fuels. The international literature suggests that intensive harvest-
ing could increase ecological risks to yield, carbon stores, soil fertility, and biodi-
versity, but geographically specific knowledge is sparse in many countries, and
results do not extend beyond 5–30 years after harvesting. We use Denmark as a
case for discussing future directions. Forest history is described, and research on
ecological effects and their inclusion in governance is reviewed. Denmark was
almost completely deforested by the early 1800s, but centuries of afforestation
have resulted in a current forest cover of 14.3%. Research commonly uses stem-
only harvesting as a reference against which to compare intensive harvesting
impacts, but pristine forests would be a more useful reference for ecological pro-
cesses and biodiversity. However, pristine forests are almost non-existent in
Europe, and non-intervention, self-regulating forests provide an alternative.
Governance and positions of non-governmental organizations in Denmark focus
more on general forest management impacts and conservation of light-
demanding biodiversity associated with historic coppicing and grazing than on
intensive harvesting. The energy sector drives the development of new govern-
ance to verify forest biomass sustainability, but the knowledge base for such veri-
fication is limited. As part of a larger solution, we suggest establishing a network
of non-intervention, self-regulating forests that can serve as a reference for long-
term research and monitoring of intensive harvesting impacts. This would allow
the application of adaptive management processes, supported by new best man-
agement practice guidelines. © 2016 John Wiley & Sons, Ltd
How to cite this article:
WIREs Energy Environ 2016. doi: 10.1002/wene.206

REVISITING INTENSIVE FOREST

*Correspondence to: ism@ign.ku.dk
Department of Geosciences and Natural Resource Management,
Faculty of Science, University of Copenhagen, Frederiksberg,
Denmark
Conflict of interest: The authors have declared no conflicts of inter-
est for this article.
Additional Supporting Information may be found in the online ver-
sion of this article.
BIOMASS HARVESTING IN
DENMARK
International research and development on use of
tree parts other than stems for industrial purposes
started in the 1960s1–3 and, as a consequence of the
oil crisis in the early 1970s, quickly focused on bio-
energy end uses. In Finland and Sweden, public
grants supported research to find more feasible and
profitable technologies for extracting low-value

© 2016 John Wiley & Sons, Ltd

 Advanced Review
woody biomass resources, including logging residues,
small trees, and stumps.4,5 In Denmark, whole-tree
harvesting and logistics systems were developed that
were especially suited for flat landscapes, dry sandy
soils, and short transport distances.
Scientific studies on ecological impacts of inten-
sive harvesting emerged as technologies for biomass
removals progressed. Early studies focused mainly on
the consequences for soil fertility,6–10 with less on
biodiversity.11 A large body of research has accumu-
lated since then, especially in Sweden, Finland, the
USA, and Canada, and several reviews of the interna-
tional literature have been published.12–17 It is
now well-established that harvesting of residual for-
est biomass can be associated with various ecological
risks.
It is critical to the successful and sustainable
deployment of bioenergy that the challenges of eco-
logical risks be dealt with through an open dialog
among scientific, commercial, government, and civil
society actors, and this discussion must include the
governance systems initiated to address these chal-
lenges.18 Discussions on ecological risks were first
raised in Denmark at a conference in 1976 and again
in the first half of the 1980s when an intense
exchange of opinions among professionals and a lit-
erature review19 led to the publication of best man-
agement practice guidelines.20 The review relied
heavily on literature from neighboring countries
because knowledge of the ecological impacts of har-
vesting residual forest biomass in Denmark was
scarce.19
After 30 years, the intensification of the forest
harvesting is ongoing, and whole-tree harvesting is
now a common practice in most forests; however,
neither the original review nor associated best man-
agement practice guidelines have been updated. Our
objective is therefore to reconsider the short- and
long-term ecological effects of intensified harvesting
within a Danish context and discuss and then suggest
future directions for research, monitoring, and gov-
ernance by:
• describing Danish forest and wood energy his-
tory, because it is important to view current
practices and their impacts in the light of past
practices and their long-term impacts;
• assessing the scientific knowledge of the ecologi-
cal impacts of intensified forest biomass har-
vesting in Denmark and elsewhere, because the
body of Danish and especially international
research literature has grown considerably since
the mid-1980s; and
© 2016 John Wiley & Sons, Ltd
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• examining how ecological effects of intensive
harvesting are addressed by existing policies,
public and private governance mechanisms, and
by non-governmental organizations’ (NGOs)
positions in Denmark, because research is often
not applied, which creates a gap between avail-
able knowledge and what is practiced.21
Our case study is specific to Denmark, but other
countries face similar challenges. Intensified forest
biomass harvesting and production is one of the tools
needed to reduce emissions of fossil fuel carbon, and
Denmark shares with many other European countries
a history of gradual felling of pristine forests and
century-long exploitation of secondary forests;22,23 it
also shares the fact that little national research has
been dedicated to harvesting regimes other than con-
ventional stem-only harvesting.
FOREST AND WOOD ENERGY
HISTORY
Denmark (longitude 8–13 E, latitude 55–57 N) has a
land area of 42,923 km2.24 It has an oceanic temper-
ate climate with an annual average temperature of
7.7 C, which is uniform across the country, and
annual precipitation varies from less than 550 mm to
more than 900 mm (1961–1990 climate normals).25
Till sediments were deposited in the ice-covered east-
ern and northern parts of the country during the
Weichsel glaciation. In the southwest, the soils origi-
nate from glaciofluvial sandy sediments and Saale till
deposits and consist mostly of nutrient-poor sandy
soils. The potential vegetation of about 90% of Den-
mark is mixed broadleaved temperate forest domi-
nated by European beech (Fagus sylvatica).26 The
current forest area is 615,000 ha, corresponding to
14.3% of the total land area.27
It is generally believed that most of Denmark
was covered with forest before the Neolithic era,
which is when humans started to farm the land,28,29
and hence, it is assumed that forests have been uti-
lized since the first humans settled more than
10,000 years ago. Data on trends in forest cover over
past centuries are not available, but deforestation
rates probably accelerated during the last
1,000 years, especially since medieval times and up
through the following centuries, when the population
grew and increased pressure on forest resources.30,31
Large parts of central and western Jutland were left
as unfertile, treeless heathland by the end of the 18th
century, and new private owners often felled trees
without replanting when the Crown sold forests to
 WIREs Energy and Environment Aspects of intensive forest biomass harvesting

create income.32 It is likely that the remaining forest
use was intensive because of the grazing, cutting, and
firewood collection that resulted in soil degrada-
tion.30 Firewood prices in Copenhagen indicate a
high demand and thus a high pressure on forests;
they rose threefold during the 18th century and
doubled between 1780 and 1800.32 Fritzbøger33 also
concludes that the general demand for firewood was
considerable, with deadwood likely cleared from for-
est floors. A recent historical assessment suggests that
a minimum level of deadwood in Danish forests was
probably reached between 1800 and 1950 because of
the combination of small forest area and high fire-
wood demand.34 The average amount of deadwood
in Danish forests is currently 6 m3 ha−1.27
The historical minimum forest area of about
4% was reached in the beginning of the 19th cen-
tury33 before increasing because of intensive afforest-
ation programs. The transition to a net afforestation
phase can be attributed partly to the first Danish For-
est Act of 1805, although various other regulations
and initiatives preceded the Act.32 Afforestation of
heathlands in Jutland began in the late 18th century
but especially accelerated with the formation of
Hedeselskabet (a land reclamation society for
improving heathland fertility) in 1866.32 Afforesta-
tion was also a welcomed means of job creation dur-
forest resources. Energy consumption became
increasingly based on fossil fuels after World War II,
and the use of wood fuels reached a historical mini-
mum in the early 1970s.34 The oil crisis of the early
1970s let to an increase in household heating with
firewood, farms started using straw, and the wood
industry used their wood wastes for energy (Figure 1
(a)).34,38 Modern large-scale use of wood chips as a
fuel in district heating and combined heat and power
production started in the mid-1980s, as it did in Swe-
den and Finland.4,5,39,40
The oil crisis highlighted the need for improved
national energy security, and governmental policies
and financial incentives were put in place to increase
the use of fuels other than oil.41 Energy security and
rural employment policies increasingly went hand in
hand with policies to combat climate change after
the adoption of the Climate Convention in 1992, and
biomass was considered to be a renewable, carbon-
neutral energy source.42 Wood-based energy produc-
tion in Denmark was 83 PJ in 2013, which was 12%
of total production (703 PJ).37 Domestic forest bio-
mass has been the main source of energy wood until
the 2000s. Wood pellets have thereafter increasingly
(a)
30
Energy content (PJ year–1)

ing the recession in the 1930s and remained so until Wood chips
the 1960s, when high employment rates and a boom- 25 Wood pellets
Firewood
ing agricultural industry slowed afforestation activ-
20 Wood waste
ities again.35 Conifers, especially mountain pine
(Pinus mugo), were planted in the first rotation 15
because the challenge was to simply get trees to
grow. More productive (but still hardy) exotic conif- 10
erous species were also introduced, including Nor- 5
way spruce (Picea abies), which was followed later
by fast-growing North American tree species.33 Third 0
and fourth rotations are now being planted, and the 1970 1980 1990 2000 2010

focus is moving from wood production towards

(b)
establishing forests with greater natural values 30
Energy content (PJ year–1)

(including planting of native species), especially in

25
state forests. The proportion of coniferous forests
reached 60% of the forest area over the 20th cen- 20
tury, but conversion to broadleaves began in the
2000s so that there are now almost equal propor- 15
tions of conifer and broadleaf stands in Denmark.27
10
The average annual afforestation rate since 2000 is
about 1300 ha yr−1,36 with two-thirds of the area 5
planted with broadleaves.27
The introduction of peat and fossil fuels for 0
1970 1980 1990 2000 2010
energy production in the late 18th century also con-
Year
tributed to the recovery of Danish forests.32 Fossil
fuels gradually reduced the demand for firewood, F I G U R E 1 | Wood fuels in the Danish energy production
which was probably the most severe pressure on 1972–201337; (a) domestic wood fuels and (b) imported wood fuels.

© 2016 John Wiley & Sons, Ltd

 Advanced Review wires.wiley.com/energy

been imported to meet increased demands 3.0 (a) Industrial wood

(Figure 1(b)). Firewood
2.5
Prior to 1980, stems were harvested for the Energy wood
wood industry, and only a minor part of branch resi-

106 m3 year–1
2.0
dues from broadleaved stands were removed for fire-
wood. It then became common during the 1.5
1980–1990s to harvest and chip whole trees from 1.0
thinnings and logging residues from clearcuts in
coniferous plantations for use in district heating and 0.5
combined heat and power production (Figure 2
0
(a) and 3(a)). Logging residues began to be increas- 1990 1995 2000 2005 2010 2015
ingly harvested in broadleaved stands in the mid-
2000s, partly replacing traditional firewood collec- 0.6 (b)
tion (Figure 2(b) and 3(b)). Wood harvested as fuel 0.5
per unit area of forest land averaged 2 m3 ha−1 yr−1

106 m3 year–1
in 2012, which is about 50% of the total harvested 0.4
volume (Figure 3(a) and (b)); this is a considerable
0.3
amount in both absolute and relative terms com-
pared to other countries (Figure 4). Forest biomass 0.2
harvesting in Denmark is thus very intensive, even if
0.1

(a) 0
1990 1995 2000 2005 2010 2015
Year

F I G U R E 3 | Annual harvesting of industrial wood, firewood, and

other energy wood from Danish forests 1990–2012; (a) conifers and
(b) broadleaves.43 Note the different scaling of the axes in (a) and (b).

7 Industrial wood
Energy wood
6
(b)
5
m3 ha–1 year–1

0
k

ce

Be s
m

en

d
SA

da
an

nd
ar

an

U
iu
an

ed

na
U
m

r la

lg

nl
Fr

Sw

Ca
en

Fi
er

he
G
D

et
N

F I G U R E 4 | Average amount of harvested wood in selected

F I G U R E 2 | Intensive forest harvesting in Denmark to produce
woodchips for use as fuel in district heating or combined heat and
power production, (a) whole-tree harvesting and chipping of a thinning
in a Norway spruce plantation (Photo: Simon Skov), (b) branches stacked
after a felling in European beech (Photo: Karsten Raulund-Rasmussen).
countries for one hectare of managed forest in 2005, distributed
according to industrial or energy end use as recorded by the Global
Forest Resource Assessment by the United Nations Food and
Agriculture Organization (FAO).44 Managed forest is obtained by
subtracting primary forest from the total forest area.

© 2016 John Wiley & Sons, Ltd

 WIREs Energy and Environment
harvesting intensity can only really be quantified by
relating absolute annual harvest to the total annual
increment of the site45 and if the relative proportions
of the crown and stem biomass removed can be
assessed.
Biomass is expected to play a significant
role in reaching the national goal of an energy system
free of fossil fuels by 2050,46 but specific bioenergy
targets have not been adopted, especially because
of uncertainty about future competition for interna-
tional biomass resources and their sustainability
characteristics. Wood energy production has never-
theless increased rapidly in the last few decades
(Figure 1 and 3), and there is considerable interest
in increasing domestic biomass production, even
if imported biomass will likely play a significant role
in the future. Several strategies for increasing forest
biomass supply, including afforestation and greater
management intensity in existing forests, have been
recommended: conversion to more productive species
and provenances, shorter rotations, increased plant-
ing densities, use of nurse crops, cutting a larger part
of increment surplus in thinnings, and changing the
balance between wood used for fuel versus other pro-
ducts;47 harvesting of residues after clearcutting and
whole trees from thinnings were not suggested, how-
ever, because these sources are almost fully utilized
and have little further potential.
ECOLOGICAL EFFECTS OF
INTENSIVE FOREST BIOMASS
HARVESTING
An assessment of the sustainability challenges associ-
ated with imported wood fuels was made in 201348
as part of the Danish government’s bioenergy analy-
sis conducted to inform future bioenergy policies;49
however, the sustainability challenges associated with
Danish biomass fuels have not been assessed since
1985.19 At that time, there were no relevant Danish
field trials that addressed the effect of whole-tree har-
vesting on growth or ecosystem processes. An evalua-
tion of possible ecological impacts was especially
based on emerging Swedish research, from which it
was concluded with caution that growth reductions
resulting from whole-tree harvesting are likely to
occur on poorer soils, but the complexity of inter-
related causes in Denmark would likely not be identi-
cal to those suggested by Swedish research; for exam-
ple, induced nitrogen deficiency caused by nutrient
removals in biomass likely play a major role in
growth reductions in Sweden, but this is less likely to
occur in Denmark because atmospheric nitrogen
© 2016 John Wiley & Sons, Ltd
Aspects of intensive forest biomass harvesting
deposition is substantial and greater than in Sweden.
No data were available for calculating nutrient bal-
ances for indicating which other nutrients might
become critical in Denmark. Best management prac-
tice guidelines based on the review were published20
and contain a key recommendation that cut trees
should be left in the forest to dry and shed nutrient-
rich needles before chipping and removal.
Given the amount of new knowledge generated
since 1985, we reassessed the ecological impacts of
intensified forest biomass harvesting in Denmark in
light of the global literature by considering five main
concerns: growth, ecosystem carbon, soil nutrients,
hydrology and water quantity and quality, and biodi-
versity. We describe the Danish context for each
issue, review Danish research on the ecological
impacts of intensified forest harvesting, and then we
supplement this with selected results from other
countries to fill knowledge gaps because the Danish
body of work is still relatively small.
Growth
Increasing interest in energy biomass led to large
efforts being made in Denmark over the last 5 years
to collect field data to create functions that describe
biomass content and allocation in different tree
components.50–53 Combining knowledge of biomass
in tree components with growth models allows simu-
lation of the development in stem, crown, and below-
ground biomass for monocultures of different species
and forest management practices (for example initial
spacing, thinning, rotation age), and how much bio-
mass can be theoretically removed. However, empiri-
cal data for actual effects of stem and crown removal
on future site productivity only exists for two Danish
experimental series. The first series includes two
experiments established in approximately 25- and
35-year-old Norway spruce plantations in western
Jutland to determine the effect on future growth of
removing on-site dried whole trees and green whole
trees in thinnings with and without threshing before
removal to increase the drop of needle and twigs.54
Four years after thinning, harvesting green whole
trees reduced stem volume growth by 10 and 17%
(p < 0.05 on both sites) compared to harvesting dried
trees and by 9 and 18% (p < 0.05 on one site) com-
pared to harvesting stems and coarse branches after
needles and twigs were threshed from branches and
left in the stand. There were no significant differences
among treatments during the next 6 years of growth
or over the entire 10-year growth period. In the sec-
ond series consisting of two experiments, removal of
logging residues after clearcutting reduced 25-year
 Advanced Review
stem volume growth in the next rotation on one site
compared to plots where residues were left and dis-
tributed evenly;55 however, no significant effect
among treatments was found on the other site. This
suggests that treatment effects can be significant on
some sites but not on others.
One of the most comprehensive experimental
series in the international literature consists of
22 Norway spruce and Scots pine stands across
boreal Finland, Sweden, and Norway that were
whole-tree thinned when 20 to 71 years old.56,57 The
thinning treatment was repeated after 10–13 years at
11 of the sites, and growth was assessed over 20 years
on seven of the sites. Results varied widely among
sites, but green whole-tree thinning reduced the over-
all growth 10 and 20 years after the first thinning by
4% (p = 0.059) and 8% (p = 0.075), respectively,
compared to stem-only thinning in Scots pine;56 the
corresponding figures for Norway spruce were 5 and
13% (p < 0.05). It is noteworthy that the negative
effect was generally greater after the second whole-
tree thinning,57 although other experiments did not
find these effects after 20 years when only one
whole-tree thinning was performed.58 The Nordic
series also showed that growth following compensa-
tory fertilization generally more than made up for
losses in the first 10-year period but not in the second
10-year period. One of the Norway spruce stands on
a fertile site in southern Finland was clearcut (21 and
11 years after each of the two thinnings), at which
time the whole-tree harvesting treatment was
repeated. There was no evidence 10 years after clear-
cutting that two whole-tree removals at thinning fol-
lowed by the removal of stems and logging residues
in the final felling reduce growth in the next rotation
of Norway spruce.59 However, the potential effects
on growth may not yet be evident because nutrient
uptake by trees in the first decade after stand estab-
lishment is relatively small14,60 compared to the years
when maximum annual growth increment is reached.
Meta-analyses and similar approaches using
data from a wider geographical range suggest that
the effects of intensive harvesting on growth vary
widely and include both negative and positive
responses, probably because of differences in climate,
soil characteristics, tree species, microclimate, com-
peting vegetation, management history and practices,
and stand development phase at which measurements
are taken.14,61,62 A review of results for boreal and
temperate forests concluded that intensified biomass
harvesting has the greatest potential to influence tree
survival and growth in the early years after clearcut-
ting,62 with some examples of reduced tree growth
later in stand development (up to 20–25 years after
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intensive harvesting) attributed to possible decreases
in nitrogen and phosphorus availability. However,
growth effects can also be variable at this stage of
stand development.63 To our knowledge, no results
from field trials exist beyond 20–30 years after the
first intensive harvesting event. A modeling study by
Morris et al.64 indicates, however, that productivity
may be negatively affected in the long term but also
that changes are not necessarily irreversible.
Evidence from Denmark is still scarce, but we
conclude with caution that negative growth responses
will mainly occur on poor soils, especially if very
intensive harvesting is conducted repeatedly in thin-
nings and again after clearcutting. Nutrient availabil-
ity is greater on loamy, relatively nutrient-rich soils,
and negative growth effects are therefore less likely,
at least in the short term (see also ‘Soil nutrients’). It
is likely that negative effects can be counteracted
through the addition of organic or mineral fertilizer.
Ecosystem Carbon
Average carbon densities of living biomass and dead-
wood in Danish forests are about 74 and 1.3 Mg C
ha−1, respectively;65 average forest floor pools are
18 Mg ha−1 (varying from 1 to 164 Mg C ha−1), and
average 0–100 cm soil pools are 156 Mg C ha−1
(varying from 23 to 682 Mg C ha−1).66 Depending
on the pool, carbon densities vary with site produc-
tivity, species, stand age, moisture conditions, and
soil texture.66–69 A large proportion of aboveground
living carbon pools is in the stem: 70–80% in stems
in beech51 and 50–80% in stems in Norway
spruce,50 with the proportion of stemwood relative
to the aboveground pool increasing with stand age.
The impacts of intensified harvesting on forest
carbon pools are not well-studied in Denmark, but
results from the experiments mentioned earlier in
which logging residues were removed after clearcut-
ting show that forest floor carbon pools were signifi-
cantly lower after 25 years compared to plots with
no logging residue removal on one site,55 but no sig-
nificant differences were found on the other site.
There were no significant differences for the top 30-
cm of mineral soil on either site. The authors suggest
that soil carbon pools will not be significantly
affected if growth is sustained; after 25 years, stem
volumes in the residue removal treatment on the first
site were significantly less, but there were no differ-
ences on the second site.
There is very little stump harvesting in Den-
mark, although it is sometimes discussed as a future
option. Studies in Sweden and Finland find decreased
soil carbon pools after stump harvesting,59,63,70 but
 WIREs Energy and Environment
this carbon decrease was mitigated in two field trials
in southern and northern Sweden where a concomi-
tant increase in tree carbon pools was found 22 and
24 years after stem + stump harvesting + deep soil
cultivation70 compared to a stem-only harvest and
manual patch scarification reference treatment.
Improved growth may lead to a larger input of
organic matter and carbon to the soil in the long
term, but a comprehensive review of the effects of
stem-only harvesting, thinnings, and whole-tree and
stump harvesting on soil-organic carbon pools con-
clude that the effects on soil carbon pools are
species-, site-, and practice-specific and that firm con-
clusions regarding impacts in boreal and northern
temperate forest ecosystems cannot yet be made.71
No significant effects of stem or stump harvesting
may be detected if the study takes place very soon
after.72
To our knowledge, no field trials extend
beyond 20–30 years, and hence, modeling is the only
option for estimating long-term effects. One simula-
tion study predicted that a change in management
from intensive clearcutting to less-intensive partial-
cutting systems would increase the long-term steady-
state pools of carbon in living and deadwood73.
Another modeling study predicted that soil carbon
pools will decrease when a previously unmanaged
forest is intensively managed in relatively short rota-
tions.64 It is also possible that soil carbon pools will
decrease when natural disturbances are taken into
account in non-intervention reference scenarios,74
although the decrease in soil carbon pools will likely
occur at a decreasing rate over time.75
Harvesting aboveground biomass will inevita-
bly decrease carbon input to deadwood and the for-
est floor, and harvesting belowground biomass will
inevitable decrease carbon input to mineral soil rela-
tive to no harvesting or less-intensive harvesting
regimes. However, several assessments have not been
able to document decreases in soil organic carbon,
possibly because of large variation, difficulties in
detecting small changes in large pools, and a too-
short measurement period. The extent to which
aboveground biomass inputs are being transferred to
the soil is not known.
Soil Nutrients
Approximately 75% of aboveground nitrogen
(N) and phosphorous (P), 60% of potassium (K) and
magnesium (Mg), and 50% of calcium (Ca) are
located in the crown of Norway spruce,76 which is
the opposite of carbon, for which the largest pool is
in the stem. It is an often-mentioned rule-of-thumb
© 2016 John Wiley & Sons, Ltd
Aspects of intensive forest biomass harvesting
that whole-tree harvesting in boreal coniferous for-
ests increases N and P removals 2–3 times compared
to stem-only harvesting.14 The corresponding figures
are lower for leafless broadleaves, but the relative
impact on nutrient cycling of harvesting residues
could still be significant. Furthermore, net export of
base cations (Ca, Mg and K) will accelerate soil
acidification.
There are no field trials in Denmark that have
directly measured the effect of intensified harvesting
on soil nutrient pools or acidification, but several
international studies exist. A study of response ratios
of stem-only and whole-tree harvesting from a large
number of experiments from other Nordic and
European countries and North America shows that
site variability of responses is large. Generally, how-
ever, negative impacts on soil nutrient pools and
acid–base status 5–20 years after intensified biomass
harvesting are more frequent in the forest floor than
in the mineral soil when compared to stem-only har-
vesting.62 A meta-analysis by Achat et al.77 found
that the supply of P and K in particular may become
critical under intensified harvesting regimes, but site
variability was also large. Notwithstanding responses
to nutrients, it has so far been difficult to link these
to changes in tree growth.
Nutrient budgets have also been used in differ-
ent countries, including Denmark, as indicators of
potential long-term impacts on soil fertility. Nutrient
budgets were developed for a Norway spruce stand
on a fertile 78 and a nutrient-poor site in western Jut-
land.79 Leaching rates and soil pools of P, K, Ca, and
Mg indicate that mineral soil weathering can ade-
quately support growth and stem harvesting on the
fertile site and perhaps even very intensive harvesting,
at least in the short term. However, nutrient
removals from fertile sites are high, and while aver-
age annual N deposition rates of 20–35 kg ha−1 yr−1
80,81
will likely fully compensate N removals of stem-
only harvesting, this is not necessarily the case for
very intensive harvesting regimes. Large inputs of Mg
with sea salt deposition, as indicated by generally
high leaching rates of this element, ensures that an
Mg deficiency is unlikely to occur in Denmark.79,82
Sea salt spray input also compensate for K and Ca
removals in Norway spruce within a distance of
about 20–30 km from the sea, regardless of harvest-
ing regime.79 Surplus N from atmospheric deposition
may cause acidification and induce concomitant
leaching of essential cations, such as Ca, Mg, and
K. Rich supplies of N to sites of medium and poor
fertility have been shown to increase the risk of P, K,
and Ca becoming limiting to growth.83 This is espe-
cially critical on relatively poor podzolic soils where
 Advanced Review
insufficient mineral weathering may lead to long-term
fertility problems.
Absolute weathering estimates have not been
made for Danish soils, although these are critical
when nutrient budgets are used to assess nutritional
effects. Modeled estimates exist in, for example, Swe-
den84 and Canada,85 but there are often significant
variations in estimation methods and models.86,87 In
a study from boreal Sweden, Klaminder et al.86 cau-
tiously suggested that the long-term cumulative
release of base cations from weathering is of the same
magnitude as accumulated harvesting losses. How-
ever, they also considered that the precision in weath-
ering estimates are lower than needed to distinguish
between effects of different levels of nutrient
removals. In another study, nutrient removals in
three harvesting-intensity scenarios are compared to
soil organic and exchangeable pools, and primary
soil mineral stocks of nutrients in 15 northern hard-
wood stands in the White Mountain region of the
northeastern USA,88 and it was found that accurate
estimates of weathering rates are critical for reaching
any conclusions. Based on Callesen & Raulund-
Rasmussen,89 we suggest that the contribution from
weathering can only be assessed in relative terms, as
also suggested in an extensive examination of weath-
ering rates derived from different models.87
Recycling of ash from biomass combustion is
often seen as a measure to at least partially, compen-
sate for the nutrient removals with intensive harvest-
ing, especially in areas such as Denmark where N is
less likely to limit growth.90 Legislation with criteria
for the use of wood ash for soil amendment is in
place in Denmark,91 and commercial solutions to
application logistics exist,92 but in practice, only
small amounts are currently recycled to forests. On-
going research projects are examining heavy metal
movement and toxicity when larger amounts of
wood ash with higher contents of cadmium than cur-
rently allowed in operations are applied.93 Current
legislation allows the application of 3 tons of wood
ash three times over a 75-year period, with a maxi-
mum concentration of 20 mg cadmium kg−1.91
We conclude that intensified harvesting will
accelerate nutrient depletion. Ca and P might be the
most critical elements in Denmark, whereas N is
probably less problematic because of the high rates
of atmospheric deposition. The most vulnerable
soils will be sandy podzolised soils in western Den-
mark. These soils are relatively acidic, which indi-
cates that weathering is insufficient to compensate
for further acidification resulting from intensive
harvesting.
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Water Quantity and Quality
Precipitation in Denmark is 700–1000 mm yr−1 in
the west and 500–700 mm yr−1 in the east. Potential
evapotranspiration is 500–600 mm yr−1 throughout
the country,94 and potential groundwater formation
in the east is significantly lower than in the rest of the
country. Fortunately, loamy soils in eastern Denmark
have a higher water-holding capacity, and often, the
rooting depth is greater than on the sandy soils in
western Denmark. An increase in annual precipita-
tion over recent decades has generally led to
increased groundwater levels, although not in eastern
Denmark, where the increasing precipitation levels
only stopped but did not reverse the trend in
decreases that prevailed in the 1990s.95 Locally and
regionally, water levels in streams, bogs, and lakes
may still be negatively affected by high extraction
levels,92 especially in eastern Denmark, where the
water surplus is less and population densities are
greater.
The water balance of forests and percolation of
excess water from the root zone are mainly con-
trolled by greater canopy interception, evaporation,
and reduced inputs to the soil in forests compared to
open land. No Danish research examines the effect of
intensive harvesting on hydrology, but generic mod-
eling for various forest management alternatives sug-
gest a modest decrease in water surplus for
production forests with high growth rates throughout
the forest rotation.96
The quality of seepage water may be influenced
by intensified harvesting. Net base-cation export can
cause acidification of both soil and seepage water,
and, in extreme cases, aluminium might be mobi-
lized, which has been demonstrated in a small num-
ber of water abstraction studies on poor podzolic soil
in western Denmark.97 Alternatively, the increase in
the export of N caused by the removal of harvesting
residues or whole trees may, at least theoretically,
reduce the nitrate content of seepage water.81,98 The
potential benefits of this would be especially impor-
tant in areas with high N deposition, including
Denmark.
It is obvious that human activities have severely
influenced natural hydrological cycles since Neolithic
times. Deforestation and reduction of forest cover
have, in several locations, led to increased water sur-
plus, perhaps creating more wetlands, increasing
streamflow or percolation to groundwater, and lead-
ing to faster surface water runoff and soil erosion on
steeper slopes.99,100 On the other hand, afforestation
with exotic conifers plus the conversion of broadleaf
to conifer stands over the past 200 years may have
 WIREs Energy and Environment
reduced the flow of surplus water from forests.101
Current forest policies to increase the amount of
broadleaves may reverse this situation, but agricul-
ture still plays a dominating role regarding both
water quantity and quality in Denmark95 as crop-
lands cover 61% of the total area.24
In conclusion, intensive harvesting per se will
not reduce water yield. Increased nutrient export
may reduce the nutrient surplus and thereby the
leaching rate of nitrate and associated cations such as
Ca and P. The resultant effect on water quality is
likely to be mixed and presumably site-specific, with
reduced N leaching where it occurs, but more acidity,
especially on less fertile soils.
Biodiversity
Denmark is home to about 30,000 species,102 which
is about 19% of the total number of species in
Europe. About 12% of the species in Denmark are
on the European Red List.103 Forests are considered
to be among the most species-rich habitats in Den-
mark because many threatened or vulnerable species
are typically dependent on deadwood, old trees, or
open moist areas and meadows in forests. Plantations
of exotic tree species do not contribute significantly
to biodiversity at a national scale.104
There has been little focus in Denmark on the
effects of intensified forest biomass harvesting on bio-
diversity, perhaps because wood chip harvesting
started in coniferous plantations, which are consid-
ered biodiversity-poor. It is also possible that the
additional effects of very intensified harvesting are
perceived as insignificant compared to the base-line
impacts of ‘traditional’ forest management, which
often involves removal of deadwood and understory
vegetation, ditching, planting of exotic species, and
use of heavy machinery.104 It has been pointed out
also in Sweden that centuries of ‘traditional’ forest
management has already significantly reduced the
amounts of coarse deadwood in the forest and chan-
ged the distribution of its qualities.105
Saproxylic organisms are among the most
threatened species in Europe because they depend on
the wood that is the feedstock of the forestry indus-
try.106 Danish surveys of deadwood-inhabiting
organisms mainly focus on macrofungi in beech
forests.107–109 The surveys show that species richness
in logs is positively correlated with log volume and is
the greatest for logs at a medium stage of decay. Inci-
dence of red-list species is the greatest for logs at
more advanced stages of decay and greater for bro-
ken than for uprooted trees. Positive correlations are
also found with stand factors such as maximum tree
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Aspects of intensive forest biomass harvesting
age, deadwood abundance, and the ratio of advanced
decay to fresh woody debris.
More rigorously designed field trials on the
effect of intensified forest harvesting on biodiversity
have been carried out mainly in North America and
boreal Fenno-Scandia. These experiments usually
focus on selected groups of organisms110,111 or spe-
cific indicators112,113 or attributes relevant for certain
species groups, such as deadwood.114 Framstad
et al.105 reviewed the ecological effects of intensified
harvesting in Sweden and concluded that the biggest
threat to biodiversity might arise from stump harvest-
ing because they are the most important coarse
woody debris component in managed forests for
insects such as saproxylic beetles.115 They considered
that adequate amounts of fine woody debris will
remain in the landscape to support the biodiversity
associated with it, even if more of it is harvested.
Dahlberg et al.116 concluded that residue harvesting
poses only a very limited risk of regional extinction
of saproxylic species, even if 70% of logging residues
in Norway spruce stands is removed on 50% of
Swedish clearcuts; in practice, about 60–80% of the
logging residues can typically be removed.54 At a
stand level, greater biodiversity values are more fre-
quently associated with sparse broadleaved dead-
wood, old snags, high stumps, and old, standing
trees.105,117–119 Ranius et al.120 concluded, however,
that even if intensified forestry production is met
with compensation measures to achieve both eco-
nomic and biodiversity goals, then it will sometimes
be necessary to avoid intensive harvesting practices.
Framstad et al.105 also argued that forest manage-
ment has generally led to more dense forests with
fewer species, especially broadleaves, less coarse
woody debris, and less natural disturbances and that
the impacts of intensified harvesting must be assessed
within the context of the impacts of forest manage-
ment in general.
Meta-analyses across several taxa are emerging
and assessing the effect of intensive harvesting across
individual species or specific species groups. In the
USA, Verschuyl et al.113 found that thinning has no
significant or even a positive effect on three diversity
attributes (‘diversity,’ ‘guild abundance,’ and ‘species
abundance’) among a wide range of taxa, including
birds, mammals, reptiles, amphibians and inverte-
brates. Riffell et al.112 compared the influence of
removing or adding downed coarse woody debris
and snags, either artificially by harvesting or as natu-
ral disturbances, on the same three diversity attri-
butes. The effect of low levels of downed woody
debris and snags was negative for birds and, to some
extent, for mammals and invertebrates, while there
 Advanced Review
was no apparent general effect on reptiles and
amphibians. Petersen et al.121 used a spatial
approach to study biodiversity based on information
of the species distribution of many different taxo-
nomic groups in Denmark. Their preliminary conclu-
sions suggest that 75,000 ha or 12% of the Danish
forest area must be set aside as non-intervention for-
est to properly promote and protect biodiversity in
Denmark.
As with other ecological impacts, we do not
know of primary investigations that have looked at
long-term effects of intensified harvesting on biodi-
versity. We also do not know of primary research
that addresses impacts on entire food webs, even
though this would give us a deeper understanding of
observed effects and the ability to manage these. It
has been suggested that the research focus should be
on species’ functions, with individual species being of
subordinate importance to function.122 However,
wide-ranging conclusions seem risky given our very
limited knowledge about the linkage between species
and ecosystem functions.123
We conclude that saproxylic species will
mainly be negatively influenced by intensive harvest-
ing, although the effect might be minor compared to
the significant impacts of modern forest management,
including introduced species, even-aged monocul-
tures, ditching, and clearcutting systems. Biodiversity
is, however, a complicated area of study that includes
multiple species interactions, even though this is not
reflected in current studies that typically address indi-
vidual species or species groups one at a time.
GOVERNANCE AND NGO
POSITIONS
It is not possible to put a market value on many of
the ecological values, and governance is therefore a
means to protect such values. Laws, guidelines, and
private certification schemes have been established in
Denmark to achieve sustainability goals for forests,
and in December 2014, an industry agreement was
reached on sustainability standards for wood chips
and wood pellets used in the energy sector. The credi-
bility of these legal requirements and private stan-
dards rely, among others, on the extent to which
they are based on science. NGOs’ policies also often
address the protection of ecological values in forests.
These organizations may influence the development
of governance through mandated public hearings
when new legislation is put in place, as founders or
stakeholders of private regulatory initiatives, or they
may exert their influence through public campaigns.
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In this section, we examined the extent to which
there is agreement between impacts of intensified for-
est harvesting and management as identified by sci-
ence and how they are included in governance
addressing the forestry and energy sectors and by
NGO positions.
The Forest Sector
The first forest act in Denmark was passed in 1805
and was revised in 1935, 1989, 1996, and
2004.124–127 The Forest Act of 1805 introduced the
concept of ‘protected forest’ (PF), which means that
the land has to remain forested forever. The Act
required ‘responsible forest management’ in the PF
areas, with an emphasis on continuous wood produc-
tion and prevention of wind erosion. Harvesting was
restricted; regeneration after clear felling became
mandatory; and grazing was prohibited.125 The For-
est Act of 1935 also addressed natural values, largely
with the same focus as the Forest Act of 1805. Later,
the concept of ‘good and multifunctional manage-
ment’ was introduced in the Forest Acts of 1989 and
1996. A requirement that ‘site fertility’a must be
maintained was first codified in the Act of 1989, and
the Act of 1996 added that ‘natural capital to sup-
port nature values’b must be maintained and specifi-
cally addressed biodiversity. The concept of
‘sustainable forest management’ was finally intro-
duced in the Forest Act of 2004 as a follow-up on
international commitments.128 It stipulated that for-
est management must take into account economic as
well as ecological and social values. The Forest Act
of 2004 still requires good and multifunctional for-
estry, but protection of nature and habitats for spe-
cies is given higher priorities than before.
In the light of the ongoing intensification of for-
est harvesting, it is noteworthy that maintenance of
‘site fertility’ and ‘natural capital’ is no longer specifi-
cally included as a requirement in the Act of 2004.
Fertilization to maintain site fertility is in principle
allowed in PF areas but must be reported to the
authorities. Forest fertilization with N is generally
not allowed.129 Fertilization with sludge and ash
from biomass combustion is allowed without docu-
mentation of fertilizer needs as long as legal require-
ments on fertilizer quality and maximum amounts
are met.91,130
The General Guidelines for State Forest Man-
agement130 advise that fertilization with P, K, and
micronutrients can take place when it is needed to
avoid a reduction of site fertility and that N fertiliza-
tion may also be considered in stands where intensi-
fied biomass removal is carried out to fulfil the
 WIREs Energy and Environment
Biomass Agreement of 1993 and 1997 (Table S1,
Supporting Information). The Biomass Agreement
stipulated the use of 1.2 million tonnes of straw and
0.2 million tonnes of wood chips as fuels in power
production.131 This goal was reached around 2010,
and the agreement is no longer applicable. The Gen-
eral Guidelines for State Forest Management is a
compilation of several policies, strategies, and pre-
scriptions developed through the 1990s and 2000s,
and it is unclear if parts are carried over from previ-
ous forest legislation. It is noteworthy that the 1985
guidelines for harvesting of forest wood chips19,20,132
are not included in the General Guidelines for State
Forest Management.
For promotion and protection of biodiversity,
the Forest Act focuses on (1) fulfilling the EU Natura
2000 directives,133,134 and (2) supporting biodiver-
sity associated with historical management systems,
such as forest grazing and coppicing. These historical
management systems were once a threat to forest
wood production but are now viewed as making a
positive contribution. Coppicing and grazing are wel-
comed on 10% of the PF area, and forests can be
converted to other open natural areas on another
10% of the FP area (Table S1). The Forest Act does
not specifically address the importance of deadwood
for biodiversity, but the General Guidelines for State
Forest Management does indirectly by stipulating a
joint goal for non-intervention forests, forest grazing,
and coppicing of at least 40,000 ha or 6.5% of the
current forest area27 by 2040. The guidelines pre-
scribe further that 3–5 trees ha−1 should be protected
as veteran trees in managed broadleaved stands
(Table S1).
State forests and some private forests are also
certified by the Programme for the Endorsement of
Forest Certification (PEFC) and the Forest Steward-
ship Council (FSC) and must fulfil the requirements
of these systems. The requirements of these private
systems largely reflect the abovementioned require-
ments of public regulations and guidelines, with some
nuances. PEFC Denmark requires that the balance
between wood harvest and increment must be moni-
tored (Table S1).135 FSC requires sustainable harvest-
ing levels and, in more general terms, a focus on
enhancing ecological functions, even if specific indi-
cators address regeneration methods, native species,
and ditching rather than site fertility.136 In line with
the Forest Act, FSC allows the use of organic fertili-
zers but only when the need for fertilization is well-
documented. PEFC generally requires that the use of
fertilizers, including organic fertilizers, should be
phased out. A minimum of 7.5–10% of the forest
should be managed for the protection of biodiversity,
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Aspects of intensive forest biomass harvesting
and part of this should or could be non-intervention
forest. Old trees should be left undisturbed, and 3–5
trees ha−1 (FSC) or 5 trees or 10 m3 ha−1 (PEFC)
should be left in managed stands as veteran trees sub-
ject to natural death and decay.
We conclude that both public and private forest
governance in Denmark broadly require sustainable
forest management, including intensified manage-
ment practices. However, considerations of soil fertil-
ity and soil carbon are weak, and it is largely the
responsibility of individual forest owners or man-
agers to identify the practical management implica-
tions of these considerations. In practice, this is an
almost unregulated area in the governance of sustain-
able forest management. The Danish PEFC was
aligned with the new international PEFC 2010 stand-
ard in 2012, and changes are not expected in the
near term. For FSC, the Danish standard from 2004
is in the process of being updated to align with the
new international FSC 2015 standard and new inter-
national generic indicators (IGI), but it is not yet
clear what the changes will be.
The Energy Sector
Intensification of forest harvesting has not prompted
any greater changes in forest-related governance, but
the increased importation of wood fuel increased the
concerns of the energy sector if the wood fuels come
from sustainable sources. The energy sector recom-
mends that sustainability requirements for solid bio-
fuels should be adopted at the EU level,137 as they
have been for liquid biofuels.138 So far, the European
Commission has recommended that member states
establish their own requirements,139 with the UK, Bel-
gium, Denmark, and the Netherlands being furthest
along in the process. In Denmark, the energy sector
reached an industry agreement with the government
in December 2014 on sustainability standards for
wood chips and wood pellets used in the energy sec-
tor, which must be fully implemented by 2019.140
The sustainability requirements included are based on
those of the UK and the Danish timber procurement
policies for the public sector (Table S1).141
At least two of the largest Danish energy pro-
ducers, Dong energy and HOFOR, plan to imple-
ment the agreement using a new private certification
system, the Sustainable Biomass Partnership
(SBP).142 The SBP has been developed by the major
energy-producing companies in Europe with the goal
of addressing the sum of all requirements in EU
member states.143 The SBP system has two pathways
for verifying its sustainability indicators: (1) biomass
that has already been certified by FSC or PEFC or
 Advanced Review
(2) biomass with no FSC or PEFC certification,
which must fulfil the requirements of the SBP Feed-
stock Compliance standard.143 This standard is also
based on the UK requirements,141 which again are
based on requirements of FSC and PEFC sustainable
forest management standards.
The SBP Feedstock Compliance Standard
requires, for example, that forest management main-
tains or improves soil quality and the forest’s capabil-
ity to act as an effective carbon sink or store in the
long term.143 How an effective carbon sink and store
can be combined with intensive harvesting is not spe-
cified, except that impacts of residue removal should
be identified, mitigated, and monitored. A mechan-
ism to feed monitoring results back into operational
practice must also be included. Other indicators
require that forests and other areas with high conser-
vation value are identified, mapped, and that poten-
tial management-related threats to such areas are
addressed. Key ecosystems and habitats must also be
conserved or set aside in their natural state. Other
requirements are in line with requirements for bioli-
quids:138 biomass must not be sourced from con-
verted areas that had a high carbon stock before
January 2008, but do not have this any longer, or
from forests converted to production plantation for-
est or non-forest land after January 2008. Biomass
removed and produced from forests converted into
open natural areas, as allowed according to the Dan-
ish Forest Act §10 (Table S1), may potentially be
declared non-eligible according to SBP criteria that
require that biomass not be sourced from areas
deforested after January 2008. However, the stand-
ard has not yet been adapted to Danish conditions.
The majority of public and private stakeholders
in Europe recommend that the EU adopt sustainabil-
ity requirements for solid biofuels, but national
authorities and forest groups, especially from forest-
rich countries, oppose this.137 They argue that cur-
rent EU policy frameworks are sufficient and that
new requirements will be too costly, too burdensome,
and a barrier to reaching renewable energy targets.
In conclusion, the energy sector’s demand for
verification of biomass sustainability is currently the
driver of new governance addressing intensified for-
est biomass-harvesting practices. The private certifi-
cation system SBP will probably play an important
role, and it appears more ambitious than Danish for-
est governance, even if it is still uncertain how it will
be implemented in practice. The SBP standard proba-
bly needs further adaptation and interpretation in a
Danish context because of potential conflicts and a
lack of clarity at a practical level, but it is likely that
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the system will address all major concerns that have
so far been raised by science.
NGO Positions
A detailed search of NGO homepages revealed that
these organizations often have positions on the use of
wood for energy and intensified forest harvesting,
even if it does not appear to be high on their agendas.
Motivated by climate change benefits, the use of wood
for energy is not rejected by any organization, but
Greenpeace, for example, does not support the use of
primary forest biomass but only residues from wood
processing and discarded wood products (Table S2).
At the other end of the spectrum, the Danish Ecologi-
cal Council supports the use of all woody biomass as
long as inefficient combustion is avoided. The Danish
Society of Nature Conservation also supports the use
of forest biomass for energy but has specific forest
management requirements, including that the area of
non-intervention forest should be increased considera-
bly as compensation. Apart from these examples,
NGO viewpoints are often in agreement with the gov-
ernment’s near-to-nature management strategy for
state forests,144 with the added opinion that this strat-
egy should also be applied to private forests.
The ‘green think tank’ CONCITO has been the
most active in a lively debate in the public media on the
climate benefits of wood fuels.145 They consider that the
climate benefits of using wood pellets and wood chips in
energy production are very uncertain and encourage
politicians to reconsider their direct and indirect support
for bioenergy. Forest researchers have opposed this by
pointing out that productive forests mitigate carbon
emissions more than non-intervention forests and that
climate benefits are only questionable when biomass
harvesting damages a forest ecosystem’s ability to
sequester carbon (i.e. decreases site fertility).146
In conclusion, there is an overlap between state
forest policies and some NGO positions, indicating
some level of direct or indirect interaction between
public and private regulation and NGO policies. Only
Greenpeace has stricter requirements than can be sup-
ported by even the most extreme scientific results and
that are stricter than in the range of requirements
found in private and public governance systems of the
forestry or energy sector. However, judging from par-
ticipation in the public debate, the specific ecological
consequences of intensive forest harvesting are gener-
ally not an issue that is high on the agendas of Danish
NGOs, except for CONCITO’s intensive involvement
in the debate on climate benefits.
 WIREs Energy and Environment
FUTURE PERSPECTIVES
Danish landscapes are highly manipulated, with for-
est ecological processes and biodiversity being
strongly affected by both historical forest utilization
and current forest management practices. Forest har-
vesting has re-intensified over the last few decades,
but national research results on the impacts of this
management are still scarce. What exists, including
the international literature, has only paid limited
attention to the fact that stem-only harvesting is an
arbitrary reference that might itself have impacts on
short-term growth, long-term site productivity, eco-
logical processes, and species diversity. Most studies
of growth effects14 and carbon and nutrient pools62
include harvesting of green whole trees or logging
residues as treatments and use stem-only harvesting
as the base-line reference. In Danish experiments,
harvesting of pre-dried whole trees has also been
used as a reference,54,147 corresponding to a ‘busi-
ness-as-usual’ base-line in Denmark over the last two
decades. Another forest reference included by Nord-
Larsen54 was harvesting of stem and coarse branches
with needles and twigs being threshed from branches
and left in the stand; at that time, this was considered
a potential integrated operations alternative to pre-
drying. However, compared with such base-line
references, it is not likely that a significant difference
in growth and site productivity of the yet-more-
intensive harvesting of green whole trees will be seen.
Stem-only harvesting is also frequently used as
a reference in studies of effects on biodiversity,110,114
even though such studies more often consider that
traditional forest management may already have had
an impact on biodiversity.105,118,119 Other biodiver-
sity studies compared harvesting with or without best
practice guidelines,111 low or high levels of coarse
woody debris (due to its removal or addition),112 or
thinned compared to unthinned forest.114 The
unthinned references have also been used in tradi-
tional thinning trials, where it will have the highest
possible basal area or stand volume of living trees at
any stage of stand development for given site condi-
tions, species, and initial stem number.148 The non-
intervention forest reference is also common through-
out the increasing amount of literature on the climate
change mitigation benefits of forests,149–152 but these
also can be the product of intensive forest manage-
ment systems with very different ecosystem processes
and species composition compared to pristine forests.
This would be the case for even-aged monoculture
plantations of exotic, high-yielding species such as
Norway or Sitka spruce in Denmark or forests that
are managed to favor pioneer species in the USA114
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Aspects of intensive forest biomass harvesting
on land that was historically covered with pristine
forest.
Studies on forests and their climate mitigation
benefits also use simulations of more natural forests
as references, sometimes including natural distur-
bances. For example, Taeroe et al.152 used measure-
ments from a 100-year-old set-aside forest to
simulate the development of a near-natural forest ref-
erence, and Seely et al.74 simulate a mixedwood for-
est stand subjected to a catastrophic wildfire on a
150-year fire cycle for comparison with monocultures
managed in clearcutting systems. However, there is
much disagreement on how carbon pools develop in
non-intervention forests149,153 and on what pristine
forest landscapes might have looked like if these
landscapes are used as a reference in relation to pat-
terns, structures, and biodiversity.28 Siitonen154
described the scientific discourse as a conflict between
those supporting the ‘closed-forest hypothesis’ and
those supporting the ‘wood-pasture hypothesis.’ The
‘closed-forest hypothesis’ was put forward by vegeta-
tion scientists more than 50 years ago, and it was
widely accepted by forest ecologists and conserva-
tionists until challenged by Vera.155 Vera proposed
that the natural vegetation is a mosaic of grasslands,
scrubs, trees, and groves and that large free-living
mammals including aurochs, wisent, and wild horse
were the driving force that determined forest struc-
ture through their effect on tree regeneration.
According to Siitonen,154 the ‘wood-pasture hypothe-
sis’ has gained popularity among conservationists,
lichenologists, and invertebrate specialists, even if the
existing palaeoecological data indicate that open-
canopy forests only became common after human
settlement, i.e., over the last 5,000 years.156
This discourse is also reflected in discussions of
Danish policies related to natural habitats for which
the key goal is the creation of more open areas in for-
ests to benefit species that depend on both light and
shelter, such as butterflies.157 It is believed that these
goals will be achieved by management activities such
as the re-introduction of large herbivores to forests,
typically domestic animals but also possibly wisent
and elk. Some see this as cultural conservation of
former (damaging) agricultural practices,126 and
others see it as a return to natural forest dynamics
after centuries of exclusion of animal husbandry
from forests.125 It is thought that animal husbandry
might have had the same effects as wild megaherbi-
vores that have been extinct for centuries or millen-
nia.104 A similar discourse exists for the open
heathlands, which would slowly turn into forest or
scrubland if trees were not removed at regular inter-
vals or browsed through grazing.158,159
 Advanced Review
Some take a more relative stance and hold that
forests at any point in time are products of both their
preceding natural and anthropogenic disturbances;160
future forest conditions will likewise be the result of
both future natural disturbance yet to come and the
way in which we chose to manage them and all
their past disturbances. Following this logic, ‘non-
intervention’ forest might be a useful term for iden-
tifying the type of reference forest that we are seek-
ing. By ‘non-intervention’, we mean forests that
possess intrinsic self-regulating mechanisms that
ensure their ability to respond to destabilizing dis-
turbances with resistance and resilience.161,162
Within this context, the intensity of forest manage-
ment can be understood as ‘the degree of external
regulation necessary to compensate the lack of
intrinsic self-regulation’.161
Suserup forest in Denmark has been left
unmanaged for about 100 years, except for compul-
sory cuttings during World War II,163 and is an
example of what could be considered a self-
regulating forest. Although left unmanaged for a cen-
tury, it is still uncertain if carbon stocks and species
composition have reached their long-term steady
states. Average forest carbon pools were probably
significantly greater in pre-Neolithic times than they
are today. Throughout the Holocene period, until
about 100–200 years ago, deforestation and overuti-
lization led to lower aboveground forest carbon den-
sities, which are presumed to have had an impact on
inputs of organic matter to the soil and the amount
of carbon stored in the soil. Living and deadwood
carbon pools respond relatively quickly to changes in
utilization and management, but soil carbon recovery
is usually very slow.164 The protection of Suserup
and reduction in pressure on other forests during the
Industrial Revolution may have resulted in some
recovery in soil carbon pools. Although we cannot
know with certainty, it is likely that today’s intensi-
fied biomass utilization is a continuation or even an
acceleration of a trend in declining soil carbon pools
over the past millennia since human settlement began
in Denmark. In extreme experiments carried out on
fallow land, with no organic matter and carbon
inputs to the soil from vegetation, only a low level of
very stable carbon fractions remains in the long
term.165
Nutrient cycles may also be affected by the leg-
acy of past utilization. Field studies show that N
leaching is significantly affected by former land use
for at least 30 years, particularly in forests on former
agricultural land.101,166 Modeling studies also indi-
cate that intensive harvesting may influence growth-
determining ecosystem processes for centuries.167
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There is also abundant field evidence that forest man-
agement history has a significant influence on biodi-
versity. Danish studies of wood-inhabiting
macrofungi indicate the fundamental effects of forest
history on community structure,108 as in studies in
other European countries.106,168 Overall species com-
position may also change through succession for cen-
turies after intensive forest management ceases and
no utilization takes place.163,169 Pairwise compari-
sons from tropical forests suggest that there is no
substitute for ‘primary forests’ (forest with no clearly
visible indications of human activities44) in support-
ing a high level of biodiversity.170 However, this con-
clusion may not be universal: (1) there is a lack of
studies in temperate and boreal forest ecosystems,
(2) results of comparisons with degraded forests may
differ from comparisons to forests managed accord-
ing to the principles of sustainable forest manage-
ment, and (3) management systems may be
developed that support new values and ecological
services.
The scarcity of protected, non-intervention for-
ests of suitable age in Denmark and throughout
Europe is problematic if self-regulating forests are to
be used as an additional base-line reference.23 Stem-
only harvesting is a relevant business-as-usual refer-
ence, but we propose that action should be taken so
that we leave our descendants the legacy of a
national network of protected, non-intervention,
self-regulating forests. A long conversion phase may
be needed, especially in regions where plantation for-
ests of exotic species dominate, but the network
could serve as a reference for forest management
research and monitoring in the long term. Knowl-
edge generated could contribute to the development
of more rigorous definitions of sustainable forest
management and a better understanding of what
data need to be collected to better monitor and doc-
ument the sustainability of forest management prac-
tices. Building on a Danish ‘legacy’ concept for
operational forestry, Thiffault et al.171 developed a
protocol for monitoring the effects of intensified for-
est harvesting on growth and selected ecological
parameters. This use of randomized and replicated
treatments within monitoring plots installed as part
of forest operations and adapted for Danish needs
could serve as a starting point for developing a cost-
effective national system for monitoring impacts of
forest management practices. Ideally, this kind of
system should be integrated within the National For-
est Inventory.
We also propose that forest-harvesting intensi-
ties be defined and assessed quantitatively in a way
that increases relevance to ecosystem processes rather
 WIREs Energy and Environment Aspects of intensive forest biomass harvesting

than as the balance between harvested stem volume 3. Secondary forests managed primarily for feed-
and stem volume growth.45 Additional quantitative stocks for commercial forest products, includ-
measures could usefully include the total amount of ing wood fuel; and
biomass removed relative to the potential net primary 4. Short-rotation forests or coppice systems of
production of the site and biomass quality in terms highly productive species managed in systems
of nutrient concentrations and biomass properties, with 3–30-year rotation ages, primarily for bio-
such as mass and dimensions, that are relevant to mass production for energy.
biodiversity objectives. Regionally verified biomass
equations could be used to convert stem volumes to
If more forests are converted to non-intervention for-
biomass and give meaningful estimates and distribu-
ests, then forests and plantations in the other three
tion of biomass. With knowledge of nutrient con-
groups should ideally, together with other landscape
centrations and biomass-harvesting intensity, the
elements, help support green infrastructures (habitats
removal of biomass, carbon, and nutrients could
and corridors) while at the same time meeting pri-
also be estimated. Indicators of forest management
mary management goals.
regime should also include system inputs such as
Throughout Europe, intensified forest-biomass
fertilizer, drainage, or genetically improved plant
harvesting and production is one of many tools
material.45
needed to avoid the further mobilization of fossil fuel
The proposed monitoring system and quan-
carbon. Within this context, definitions of sustainable
tification of harvesting intensities and manage-
forest management practices must more widely rec-
ment regimes could be applied through an
ognize that the ecological functioning of intensive
adaptive forest management approach, which can
forest management systems are far from those of
be used to improve practices even when full sci-
pristine or self-regulating forests. We call for more
entific knowledge of all relevant impacts under
conscious and clear recognition of the degree to
changing conditions cannot be obtained.17 The
which forest management affects the ecological func-
adaptive approach will help us move on, monitor
tioning and species composition of forests and to
impacts as we go, and improve our management
which observed effects are the results of land-use his-
based on quantified impacts and benefits through,
tory. There is currently very limited knowledge avail-
for example, regular revisions of best manage-
able in face of the needs suggested by current forest
ment practice guidelines.
management practices and perspectives. Achieving
Guidelines are tools that facilitate the imple-
contrasting goals within the same forest or area may
mentation of more sustainable practices.172 How-
fail when trade-offs become unachievable, such as
ever, forests can be managed for multiple goals, and
managing forests for both conservation of biodiver-
if these cannot be contained within a single set, then
sity and maximum climate change mitigation bene-
different guidelines specific to different objectives
fits. Clearer classification of forests based on
may be required. We propose that best management
management goals coupled with knowledge from
practice guidelines be developed for Danish forests to
research and monitoring are needed for political and
help manage at least four types of forests and associ-
societal discussions on the composition and distribu-
ated goals:
tion of forest types that are needed to best achieve
the many forest management goals.
1. Forests with long continuity and natural eco-
system dynamics set aside as non-intervention,
self-regulating forests;
NOTES
2. Secondary older forests, typically more than
a
150 years old, accepting management to In Danish: Fredskov.
b
achieve non-commercial goals, such as biodi- In Danish: Dyrkningsgrundlaget.
c
versity conservation or recreation; In Danish: Naturgrundlaget.

ACKNOWLEDGEMENTS
We gratefully acknowledge the Velux foundation for providing funding for this work under the 2011 grant,
‘Consequences of large scale bioenergy production in Denmark on biodiversity, forestry and agriculture in the
context of sustainable development’ (http://veluxfoundations.dk/). We also thank our colleague Brian Titus for
careful revision of the language.

© 2016 John Wiley & Sons, Ltd

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