The impact of forest management on biodiversity

by

Johnny de Jong, Jonathan W. Humphrey and Hans Peter Ravn

2.1. Introduction
Forests and other wooded land cover roughly 30 % of the total land area of Europe and deliver a wi
range of social, economic and environmental benefits including key components of biodiversity (FAO,
2004; CEC, 2006; EEA, 2006). In recent decades, global and European agreements on sustainable
multi-functional forestry have led to the development of policies and targets for the conservation and
enhancement of forest biodiversity in Europe (CEC, 2006).

Under the 6th Environment Framework (CEC, 2001), member states in the European Union (EU-25)
have agreed to halt the loss of biodiversity in Europe by 2010, and a recent review of progress in
achieving this goal has indicated positive trends for forests (EEA, 2006). In particular, forest area is
not decreasing, forests are growing older and thus more valuable for conservation, a high percentage
of forest area in some countries has now received independent certification indicating that sustainable
management is in place, and 25% of the forest area is now protected to retain biodiversity and
landscape values. However, there is still a need to address issues such as the impact of habitat
fragmentation, harvesting of old-growth forest, climate change and pressure for intensification of
forest utilisation leading to simplification of forest biotopes in some countries (EEA, 2006).

One of the most difficult challenges faced by the forestry sector is to deliver improvements in the
economic outputs (timber and other materials) from forests whilst not unduly compromising
biodiversity (Angelstam et al., 2004). In addition, there is an increasing realisation that biodiversity
conservation is unlikely to be achieved by pursuing a strategy that focuses solely on protecting small
areas of key biotopes or the needs of a few priority species and targeting economic activity in other
places (Andersson et al., 2004; Bruinderink et al., 2003; Watts et al., 2005).

To date there has been no attempt to study the impacts of forestry activities on sustainability across the
whole Forest-Wood-Chain (FWC). In this section the impacts of FWC activities on biodiversity is
reviewed in a European context. The aim is to present an evaluation and synthesis of the known effects
of specific forest management operations (Table 1.2) on biodiversity and to rank the importance of the
different effects pointing out the most important and influential management operations.

2.2. Concepts and indicators

The term “biodiversity” or biological diversity has been defined as “the variability among living
organisms from all sources including, inter alia, terrestrial, marine and other aquatic systems and the
ecological complexes of which they are part; this includes diversity within species, between species
and of ecosystems [sic]” (UN Environmental Programme, 1992). The term is frequently used in
conservation discussions. In general terms the most important abiotic factors determining the
distribution and abundance of different taxa across Europe are climate, soil type and geomorphologic
conditions. Also biotic factors such as competition, predation, parasites, diseases etc are important
(Krebs, 1985). In forests other important factors are the structural complexity (e.g. variation in
vegetation structure, gaps, edges), occurrence of important substrates (old trees, big trees, dead wood
etc.), and tree species composition (Esseen et al., 1992).

The variation of complexity, substrate and tree species composition is a result of different types of
disturbances, and all together these factors create a big variation of forest biotopes in which different
species are adapted to.

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Here we define biotope as the type of environment, often described as a combination of vegetation
type, tree species composition, structure (the physical features of the environment, e. g. open, semi-
open or edges) and management, e.g. semi-natural pastures, old-growth blue-berry spruce forest, or
open meadows. Sometimes “Habitat” is used as synonym to “Biotope”, but here we define “Habitat”
as the range of environment in which a species occur (Krebs, 1985). Within the same type of biotope
the tree-species composition, structure and management regime may vary, as well as the number of
habitats for different species. Over recent years, attempts have been made to develop classification
systems for forest biotopes (Larsson, 2001; Barbati et al., 2006) and these systems give a framework
and context for understanding the impacts of forest management on biodiversity. However, not only
qualities in the forest biotope but also the arrangement of these biotopes in the landscape are important
for biodiversity (Angelstam, 1997).

In order to understand how to conserve biodiversity in practical terms it is important to recognise the
need for specific goals. Normally, the goal is not to get as many species as possible, but to conserve
the species occurring naturally (i.e. not introduced by humans). Total measure of biodiversity is
scientifically interesting in order to understand biodiversity pattern, but it is not used for conservation
purposes. Instead a number of different indicators of biodiversity have been suggested (Lindenmayer
et al., 2000). In reality the conservation discussion mainly focuses on the red-listed species which
often have very specific biotope, or substrate requirement. The underlying assumption is that if we
focus on the more demanding red-listed species also all other species with more general requirement
will be conserved.

Red-listed species are used both for assessing forest qualities and for evaluating management, or
conservation methods. Instead of using all red-listed species a subset of species are used as indicators.
The idea is to use a nested pattern, in which occurrence of one species indicates occurrence of many
other species. There are many suggestions of indicator species, however there are only few examples
of scientifically investigated nestedness pattern (Nilsson et al., 2001). Another problem is that
occurrence of species, or abundance not always are good indicators e.g. if population viability or
source-sink pattern is unknown (Van Horne, 1983). One type of indicator is the umbrella species
(Simberloff, 1998), which means that a number of species have similar requirements of substrate or
biotope complexity, even though there are no other ecological links between the species. Woodpeckers
have been suggested as umbrella species (Martikainen et al., 1998), and as good indicators of
naturalness of forests (Angelstam and Mikusinski, 1994).

Species identification is often a problem, and because of that many other types of indirect indicators
are used, such as abundance of dead wood, tree-species composition, occurrence of specific substrates
etc (Nilsson et al., 2001). One problem with these indirect measurements is that you also have to know
how much quantity is needed of the specific substrate or biotope for species survival. Some threshold
values have been suggested, e.g.20-40m3/ha for abundance of dead wood in temperate conifer forest
(Humphrey et al., 2004, de Jong et al., 2004) and area of suitable biotopes, but for most species we
have no data on the limiting factors and threshold values. Further, the threshold value might vary
within regions and during the seasons (Wiktander et al., 2001).

Another method for biodiversity assessment without making species surveys is to use Habitat
Suitability Index (HSI). Instead of detailed knowledge of species occurrence this is based on the
composition of habitats in the landscape (Angelstam et al., 2004). However, when the HSI is created
detailed knowledge about habitat selection, habitat use, dispersal pattern and other factors for some
indicator species occurring in the landscape must be known. In addition landscape structure is also of
key importance for species survival (Andrén, 1994; Fahrig and Merriam, 1994; Villard et al., 1999).
During the 1980s landscape ecology became a scientific discipline of its own with big influence on
conservation biology. Habitat fragmentation has been pointed out as one of the most negative factors
behind species extinction (Fahrig, 1997).

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Habitat fragmentation contains two components affecting biodiversity: Habitat loss and isolation.
Habitat loss means that the suitable area is decreasing and fewer individuals can use the resources. The
population size decreases, and finally if the habitat loss continues the population will not be viable due
to genetic or demographic factors. The habitat loss might also result in a patchy landscape. If these
patches are small and isolated the population on each patch will go extinct, even though the total
number of individuals in the landscape is big. Species survival depends on habitat area, habitat
isolation, occurrence of migration routs through the matrix (corridors), and quality of the matrix. In the
forest landscape it is obvious that clear-cuts creates a fragmented forest. However, it is important to
remember that also other types of management create fragmented forests. Forests with high quality for
biodiversity are islands in a well managed forest. In Sweden, for example, the forest area and the
forest volume have increased considerably during the 20th century, but meanwhile the fragmentation
has increased due to more intensive management. That fragmentation is a real problem has been
demonstrated in many empirical studies (Saari et al., 1998; Komonen et al., 2000). On the other hand
there are a number of species, including some species specialised on specific substrates of dead wood,
which are able to disperse long distances or survive on clear-cuts with high abundance of dead wood
and which not experience forest patches as real islands (Ås, 1993).

The dispersal capability varies a lot among different species. Species with low dispersal capability will
be the first one affected by fragmentation. If low dispersal capability is combined with low persistence
(i.e. low possibility to survive during critical periods) the extinction risk increases. However, also
species with low dispersal capability are moving around, and by creating a good infrastructure species
might survive and explore new areas with suitable habitat. Therefore it is relevant to talk about
continuity of suitable habitats on the landscape level (Angelstam, 1997; Sverdrup-Thygeson and
Lindenmayer, 2002).

Many studies, both empirical and theoretical, have demonstrated that extinction due to habitat loss and
fragmentation often is a slow process (Hanski 2000). Until a certain limit populations of species will
survive even though their habitats disappear. However, when this limit is passed the extinction might
be rapid. To identify this threshold value of remaining habitat for species survival has been an
important area of study in conservation biology (Fahrig, 2001). Several studies indicate that the
probability of extinction increases dramatically when less than 10-30 % of the original habitat area
remains (Andrén, 1994).

2.3. Forest Management – methods, approaches and effects on biodiversity

Forest management includes for example: clear-cutting, drainage, soil scarification, plantations, pre-
commercial thinning and thinning. Often it results in even-aged monocultures. Long-term
consequences of forest management in the landscape include decreasing areas of old-growth forest,
decreasing number old trees, dead wood and other for biodiversity important structures (Linder and
Östlund, 1998; Andersson and Östlund, 2004). However, the consequences of forest management on
biodiversity can vary considerably depending on which methods are used, and in many cases forest
management and species conservation can be combined.

2.3.1. Tree species choice and methods of regeneration

When the forest regenerates naturally, the next generation of trees is a result of the available seed
sources and natural competition within and between species. When seeds or seedlings are planted man
is involved in the selection. The traditions in European forestry on tree species selection are highly
variable between the regions. In intensively driven forests where yield in cubic meters has priority,
exotic genetic varieties or species is often the rule. Some tree species – e.g. Norway spruce, lodge pole
pine and Sitka spruce – have been turned into the main tree species outside their natural vegetation
zones. Even where broadleaved trees such as beech are the natural vegetation the seeds used for
planting may have been selected from an exotic origin. In some areas of Europe where the former
natural tree vegetation has been removed by man, there have been attempts to re-establish this
vegetation sometimes using exotic species. For example, tree planting experiments in sub-arctic parts

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of Europe the tree species may have been collected on the southern hemisphere. E.g. Nothophagus spp
on the Faroe Islands (Ødum, 1979). The change of tree species will affect biodiversity as well as the
homogenous structure of the plantation. In general dense coniferous plantings will allow no vascular
plants or other vegetation to survive on the forest floor and very few insect species will survive in
these areas. Further on acidification of the soil will influence the microarthropod fauna. The number of
insect species associated with various tree species has been analysed in several studies. The number of
species of the major plant feeding orders of insects (Lepidoptera, Coleoptera and most groups of
Hemiptera) associated with British trees is closely correlate with the number of records of their
Quaternary remains (Southwood, 1961; Kennedy and Southwood, 1984). In Britain the highest
number of insect species is found on oak, willow, birch and hawthorn, whereas in Russia the highest
number is found on Pine (Southwood, 1961).

Diversifying the tree species composition of plantations can be extremely beneficial to biodiversity.
For example in the UK, naturally created gaps in upland spruce forests are often colonised by
broadleaved trees and mixed conifer/stands are becoming increasingly common (Humphrey et al.,
1998; Mason, 2006). Increasing the broadleaved area and number of native broadleaves species in
conifer plantations is generally beneficial to biodiversity (Patterson, 1993; Humphrey et al., 1998).
The diversity of fungal (Humphrey et al., 2000) lichen and invertebrate communities (Humphrey et al.,
1998) has been shown to increase in response to increasing broadleaves.

Intra-specific variation in different trees species may also be of importance for dependant diversity.
For example genetically modifying trees for resistance to pests and diseases can impact on the value of
that tree species as host for a variety of organisms (Carnus et al., 2006). Therefore when dealing with
stands of site native species in particular, the conservation of biodiversity is often best served by using
natural regeneration which helps to retain autochthonous genetic variability (Peterken, 1993).

2.3.2. Site preparation

2.3.2.1. Physical manipulations
The main site preparation methods used prior to both afforestation and reforestation are tillage,
ploughing and scarification. Site preparation is important for several reasons, e.g. it has a negative
effect on weeds competing with the planted seeds or seedlings, and exposed mineral soil around the
new plant has a negative effect on the pine-weevil, Hylobius abietis. Scarification is beneficial for
some vascular plants adapted to disturbances (Pykälä, 2004; Haeussler et al., 2002). The species
composition, species richness and abundance of vascular plants are all affected. Haeussler et al. (2002)
demonstrated that species richness of vascular plants peaked after moderately severe site treatment,
and that the removal of soil organic layers resulted in a higher abundance of species regenerating from
seeds. However, some other organisms are negatively affected. Bellocq et al. (2001) demonstrated that
arthropod diversity declined with increasing post-harvest site disturbance especially collembolans and
mites – which is important for keeping the soil fertile by making adventitious pore structure. Drainage
of wet habitats such as peatland, fens and swamps has in the past led to loss of wetland biodiversity,
e.g. in northern Scotland, planting on deep peat led to erosion and loss of habitat for wading birds
(Lavers and Haines-Young, 1997). In the forest of Grib skov in Denmark, Rune (1997) documented
extensive reduction in wet areas over the last 100 years with a dramatic change in the flora as a
consequence.

2.3.2.2. Chemical treatments

The use of chemical control methods in forestry is in general limited in comparison to other growing
systems (agriculture and horticulture). However where the rotation is intensive in time (energy forest,
Christmas tree production) or space (nurseries) the pesticide usage is also intensive. Control methods
always have side effects on non-target organisms. In forestry at large these side-effects are considered
limited. The closer production and management methods resemble intensive agriculture the more we
could expect the same negative consequences on biodiversity as known from e.g. agriculture. For
example, the collembolan species Folsomia quadrioculata has in the Boxworth growing system

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experiment shown to be negatively correlated to more intensive pesticide usage (Greig-Smith, 1992).
This species is abundant in forests soils and is essential for good soil structure.

2.3.2.3. Prescribed burning

In some regions, prescribed burning is used to reduce competition from vegetation on tree
establishment. However, it can also have benefits for biodiversity. In former times wild fires were the
most important factor affecting the abundance of dead wood in the northern boreal forests (Ehnström,
1997). Through prescribed burning it is possible to create more favourable conditions for the
organisms especially adapted to the post-burning situation. This occurs where trees are left on clear-
cuts before burning. Many rare and threatened insect species benefit from prescribed burning and
burnt trees that it creates (Wikars, 1992). Also several bird species are favoured by the variation in the
landscape created by fires (Dale, 1997). Mychorrhiza fungus has been shown to respond to fire by
fructification (Vrålstad et al., 1998). Some species regarded as pests are also attracted to fire, e.g. the
longhorn beetle Monochamus sutor and the wood wasp Urocerus gigas may cause economical
damage on the wood. Also Hylobius abietis is attracted to burned areas. The fungal pathogen Rhizina
undulata gets virulent when exposed to temperatures 35-45°C (Petersen, 1971).

2.3.3. Stand management and harvesting

2.3.3.1. Forest management alternatives
The five management alternatives described in Table 1.3 all have different consequences for
biodiversity as described below:

1. Forest nature reserve. Conservation of biodiversity is the key objectives for this type, which
means leaving the stand without doing any management. Stands in which the organisms are
adapted to natural disturbances (e.g. large areas in the boreal forests of northern Europe) that are
left without any management very soon (within 30-40 years) develop high nature values, mainly
due to increased amount of dead wood.
2. Close to nature forestry. In order to conserve biodiversity it has been suggested that forestry must
mimic the consequences of natural disturbances (Bengtsson et al., 2000). This idea has partly been
adopted by forestry and includes leaving some dead wood, big trees, small biotopes with high
value for biodiversity (Larsson and Danell, 2001; Ferguson and Elkie, 2003) and the result have
been evaluated in a number of studies (Väisänen et al., 1993; Esseen et al., 1997; Kaila et al.,
1997; Hazell and Gustafsson; 1999; Vanha-Majamaa and Jalonen, 2001; Hautala et al., 2004;
Ekbom et al., 2006). It is however obvious that it seems to be impossible to combine conservation
of some species with economic sustainable forestry and this is an important reason for including a
network of forest reserves within the managed landscape (Ökland, 1994, 1996; Niemelä, 1997).
However there are also some stands in which the organisms are adapted to cultural disturbances
(e.g. deciduous forest in the hemi-boreal or nemoral zones) the biodiversity might decrease
without management intervention.
3. Multi-purpose forestry. The purpose here is both to increase forest production and conserve
biodiversity. This includes leaving all tree-stumps, leaving some old or big trees, leaving dead
wood, in some regions it also mean to conserve a high proportion of deciduous trees. If the goal is
to optimise forest production it is always negative to leave areas for free development except
where thinning increases the risk of windthrow e.g. in north west Europe (Quine, 2000).
4. Intensive even-aged forestry. In this management alternative, the purpose is to create a
monoculture with high forest production. Rotation lengths can be quite short (20-40 years) and
harvesting takes place by clear-cutting. The consequences of this type of management for
biodiversity is invariably negative at the stand level, except where the intensive plantation has
replaced a land use of even lower biodiversity value such intensive agricultural fields (Humphrey,
2003).
5. Wood-biomass production. This management altrenative is an extreme version of (4) where
rotation lengths are even shorter. In most cases the cut trees are collected and used for bioenergy.
This alternative is always negative for biodiversity compared to the other four options.

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2.3.3.2. Silvicultural regime
In the management alternatives 2, 3, and 4, there are a range of different silvicultural regimes available
to managers which have differential impacts on biodiversity. These are reviewed in general below.

2.3.3.3. Clear-cut system

Clear-cutting is the most dramatic change in the forest succession. The consequences of clear-cutting
on biodiversity might be positive or negative. The result depends on which species or species group
that are considered, and how the cutting has been carried out in relation to the natural disturbances in
the area. For species adapted to old forest with small scale disturbances and long continuity of tree
cover (e.g. many cryptogam species), clear-cutting results in habitat loss and fragmentation of
remaining suitable habitat. However, some of these species are able to persist during the regeneration
phase, and species with good dispersal ability are less affected. Species adapted to large-scale
disturbances might benefit from clear-cutting provided that suitable habitat and substrate are created.
This means that in some types of forest managed under the Close to nature management alternative,
some clear-felling may be appropriate to conserve biodiversity (Quine et al., 1999). In some natural
forest the fire is the main disturbance creating large areas of open forests. Some of the species, but not
all, adapted to post-fire biotopes are able to survive on clear-cuts. In some cases, clear-cutting is
combined with bio-fuel harvest. This will decrease the structural diversity at the site which decreases
the possibilities for some ground living species to survive the open biotope succession phase (Åström
et al., 2005).

Vascular plants is one example of a species group which is less affected by clear-cutting, or which
even might benefit from clear-cutting. Early succession stages of forests are important for many plant
species, and the abundance might increase considerably (Lindholm and Vasander, 1987; Humphrey et
al., 2003). In a study of plant communities in Canada Haeussler et al. (2002) demonstrated that species
richness was 30-35% higher 5-8 years after logging compared to the old forest. The result was
confirmed in Finland by Pykälä (2004) who concluded that the number of species was almost double
in clear cuts compared to mature herb-rich forests. As a consequence of increasing abundance of some
herb species on clear-cuts several mammals benefit, such as rodents and cervids. Also some generalist
predators such as red fox, wolves and lynx benefit from increasing abundance of rodents and cervids.
Some of the most negatively affected species are the pine marten (Brainerd et al., 1995), squirrels and
some species of bats (de Jong, 1995, Ekman and de Jong, 1996). However, most species of bats
benefit from increasing edge-area. The response on bird species varies considerably. In short predators
feeding on rodents or generalist predators are favoured by a more open landscape benefit by clear-
cuttings (Petty, 1998). Also many other species common in the agricultural landscape associated with
open or semi-open grassland and bushes are favoured by clear-cutting (Humphrey, et al., 2003), while
species adapted to permanent tree cover or natural wildfire or water disturbances decrease. Bird
species in the latter group are often non-migratory, e.g. wood-peckers (Mikusinski et al., 2001).

Amphibians are severely affected by clear-cutting. During some parts of the year amphibians are
connected to water, but many species spend a lot of time in terrestrial biotopes. Several studies have
demonstrated a total elimination of salamanders due to clear-cutting (Petranka et al., 1993; Petranka,
1994). In general many species of amphibians requires humid condition and occurrence of dead wood.
However, by using adapted management near aquatic biotopes it might be possible to combine
forestry with clear-cuts and conservation of amphibians. Invertebrates and cryptogams adapted to old-
growth forest with natural disturbances, with high degree of specialisation, low dispersal ability and
low persistence belongs to the most negatively affected species in the managed forest. Most of the red-
listed species in forest belongs to this group and in general clear-cut is the main threat. Because of low
dispersal ability fragmentation also affects some generalist arthropods such as spiders and ground
living beetles (Miyashita et al., 1998; Abildsnes and Tømmerås, 2000).

One well studied consequence of clear-cutting is the edge effect. A new edge means new climatic
conditions and interactions with new species for the species living in the forest. The result depends on
the composition of the edge (structure and species composition) and sun and wind exposure. The

6
increased wind exposure often results in higher abundance of dead wood near the edge which is
positive for many species adapted to disturbances and using dead wood. One example is beetles of the
family Scolytidae of which several species plays important roles in the boreal forest ecosystem
(Weslien, 1992, 1994). Several other insect families are very abundant in the edge biotopes (Helle and
Muona, 1985; Ferris and Carter, 2000), which also favour birds eating insects. In the boreal region
many species of birds and mammals are attracted to edge biotopes (Hansson, 1994). However, for
many other species edge-effects are mainly negative due to the climatic changes and increased
competition (Spence et al., 1996; Esseen and Renhorn, 1998). For bryophytes it has been found that
the climatic consequences are more dramatic for south-facing edges compared to north-facing edges
(Hylander, 2005).

In the boreal region clear-cutting has been compared with fire disturbances. By mimicking post-fire
biotopes as much as possible it might be possible to increase the species number on the clear cuts
(Similä et al.; 2001). In North-America Reich et al. (2001) compared clear-cuts and wildfire areas of
different succession phase and found no difference in plant species diversity. Even though there is a
structural similarity, there are also some important differences (Delong and Tanner, 1996; Bergeron et
al., 2002). A fire disturbance creates a lot of dead wood and a more varied structure and the ground
cover is burned off (Bergeron, 2004; Harper et al., 2002, 2004). To leave some trees and biotopes on
the clear cut does not completely compensate for fire disturbances, also restoration of biotopes by
using fires is important (Niemelä, 1997).

2.3.3.4. Continuous cover systems

Continuous cover silvicultural systems encompass varying types of non-clear fell management (Mason
et al., 1999), and include shelterwood, group, and selection systems (Mathews, 1999).

Small-scale cutting (e,g, small groups <0.25 ha or individual trees) is an alternative harvesting method
to clear-cutting. At some sites it is difficult to regenerate the forest when using clear-cutting and thus
selective cutting might be a solution. Selective cutting is also used for conservation reasons. Small
gaps created by thinning and small group felling allows the development of larger trees and the
provision of habitat conditions for a range of species group such as fungi, red squirrels, bryophytes
and hole-nesting birds (Humphrey, 2005). This will benefit species requiring continuous tree-cover
such as ectomycorrhizal fungi. These species are naturally adapted to low intensity fires of which most
tree individuals survive, or to refuge biotopes (Dahlberg, 2002). However, in contrast Kropp and
Albee (1996) demonstrated that thinning affects mycorrhizal fungi. The total number of species was
reduced with some species negatively affected while others were positively affected.

Not only is continuous tree cover important, but so also is the continuity of other resources such as
large trees and deadwood. Selective cutting regimes can be particularly valuable if some trees are left
in perpetuity to create large diameter standing and fallen deadwood (Peterken et al., 1992; Humphrey
et al., 2002; Humphrey, 2005).

In a study in Sweden, Bader et al. (1995) demonstrated that the consequences for wood-inhabiting
fungi differ a lot depending on how the selective cutting is carried out. For these species the
abundance and quality of dead wood in the new stand is most important. Among saproxylic beetles
especially species living in hollow trees requires continuity of substrates (Nilsson and Baranowski,
1997). There are a number of other studies demonstrating the importance of continuity of different
substrates which in general is rare in managed forests (e.g. Mycetolophidae, Ökland, 1994).

2.3.3.5. Thinning and pruning

All silvicultural regimes rely on some form of pre-commercial thinning and commercial thinning if the
goal is to maintain or increase forest production. The methods adopted are also important for
biodiversity. There are however, very few experimental studies on the consequences for biodiversity
of thinning or cleaning. However there are many studies comparing biodiversity in old-growth forests
and managed forests (Söderström, 1988; Gustafsson and Hallingbäck, 1988; Andersson and
Hytteborn, 1991; Siitonen and Martikainen, 1994; Martikainen et al., 2000). One important difference

7
between old-growth forests and managed forests is thinning. Old-growth forests have long continuity
and natural disturbances which are prevented in managed forests). Pre-commercial thinning (or
cleaning) and thinning change the composition and structure of the stand, and it has been shown in
several studies that increased complexity and variation due to retention harvesting increase the
conservation value (Work et al., 2003). In conservation mainly big trees have been in focus, however,
for some invertebrates and fungi fine woody debris is very important (Kruys and Jonsson, 1999;
Nordén et al., 2004).

The reason why big trees are such an important structure for many red-listed species is that this
resource is rare. Concerning fine woody debris the situation is totally different. Even though some
species are specialised on fine woody debris few of these are red-listed. Until now fine woody debris
has not been a limiting resource. However, with increasing interest for bio-fuel there is a risk that also
fine woody debris will be a limiting resource and that the number of species falling into the red-listed
category will increase.

2.3.3.6. Rotation length and crop rotation

Often when a tree is harvested it is less than halfway through its natural life cycle. The senescence,
dying and decay of a dead tree is omitted in rational forestry. The tree is cut when the annual increase
in growth starts to decrease. However, most species living on wood use old trees, dying or dead trees
(Samuelsson et al., 1994) and by removing the tree before it is biological mature the habitat for a
number of organisms will be removed.

In many growing systems crop rotation is considered good management practise since negative
influences by pests and diseases are diminished in this way. However this method is also negative for
organisms of no economic importance. For example by interrupting a long continuity of beech forest
by introducing Norway spruce the fungal flora will change dramatically (Flemming Rune, pers.
comm.). Peterken et al. (1992) and Humphrey (2005) have recommended extending rotations in
temperate plantations to benefit species associated with late successional forests such as lichens. In
contrast there is also evidence that species can adapt to plantation forestry and ecologically short
rotations. For example Humphrey et al. (2000) found that Sitka spruce plantations supported a range
of red data book fungi.

2.3.4. Other management options

2.3.4.1. Road construction, fencing, other infrastructure
Transporting attacked and diseased wood out of the forest is an important countermeasure against
pests. Consequently road building in forests is one of the most important actions to handle problems
with pests and diseases. During the “battle against bark beetles” in Norway 1978-1980, most resources
were dedicated to road construction (NOU, 1979). However, for other species with limited dispersal
abilities roads, tracks and other infrastructures may act as barriers eliminating or limiting migration.

Jepson (1994) presented information on boundary permeabilities to certain ground beetles. He found
that 3 m dirt road reduces permeability to 50%, 1 m gravel track reduces permeability to 15%, 0.5 m
paved road reduces permeability to 23% and 5.7 m railway embankment reduces permeability 10-17%
compared to the original biotope. Fry and Robson (1994) showed that even a 1m high hedge
significantly reduced landscape permeability to butterflies. Wildlife bridges have been made to
facilitate high-way crossing of deer, toads/frogs and dormouse, Muscardinus avellanarius. Wildlife
fences are made to keep out deer from new plantations. Not only culture plants are influenced but also
herb diversity will be different on the inside of the fence compared to the outside.

2.3.4.2. Management of edges, buffer zones and open space for biodiversity
Open space and edge habitats are key features for biodiversity in managed forests and can include
areas of unimproved scrub, tree-line/montane scrub, grasslands, crags, bogs, heaths, and limestone
pavements (Ferris and Carter, 2000). Wet areas, such as lakes, ponds, streams, fens, bogs and marshes

8
provide valuable aquatic habitats (Lindemayer et al., 2006). Species suited to open ground and forest
edges abound in some parts of Europe because of a long history of cultural management of ‘open’
forest with glades and rides, such as wood-pastures (Vera, 2000). Way-leaves, roads, rides, and
thinning racks can form a network of open ground and edges to allow species movement inside and
outside the forest (Ferris and Carter, 2000). Management of edges and open areas is usually required
to maintain diversity; vegetation succession if left unchecked may lead to loss of valuable species. For
example, many birds depend on the maintenance of a diverse edge structure (Fuller and Browne,
2003). Butterflies require nectar sources and food plant associated with edges and open areas (Tudor
et al., 2004). Management actions can include: thinning ride edge trees and encouraging the natural
regeneration of native shrubs/plants to increase nectar sources and produce a graded edge structure;
maintaining the diversity of successional habitats such as scrub woodland (e.g. by flailing), grassland
(e.g. by grazing/mowing) and heathland (e.g. by period burning); encourage mosaics or gradations of
open ground and woodland for species such as black grouse (Cayford, 1993).

Buffer zones are important features in managed forests as they offer protection to sensitive biotopes
such as aquatic ecosystems. Riparian areas form some of the most important buffer zones in managed
forest (Potvin and Bertrand, 2004) as they form the interface between aquatic habitats and woodland.
Riparian zones can have high biodiversity value as they contain a diversity of habitats and act as
important corridors for the movement of wildlife (Petersen et al., 2004). Riparian woodland also has
an important role in improving floodwater storage and providing resources for aquatic communities.
The most important management consideration in the riparian zone is the density and distribution of
trees, and therefore shading, and how this relates to natural bank features. Maintenance of bank
processes and habitats supports a wide variety of wildlife. Some riverbanks may be relatively species-
poor as a result of heavy shading by trees, for example densely planted non-native conifers. Riparian
buffer zones are also prone to invasion by non-native invasive species such as Himalayan balsam
(Hejda and Pyšek, 2004). These are often pioneer species able to thrive in dynamic habitats within
riparian zones.

2.4. Synthesis
Forest biodiversity is a result of different types of natural and anthropogenic disturbances creating a
high variability of habitat, structures and substrates. This variability is also related to soils, climate and
biogeographic zone. The impact of FWC chain activities on biodiversity has to be evaluated in relation
to biophysical context and the different forest types found in different parts of Europe. Thus it is
difficult to generalise across Europe as a whole. Furthermore, the consequences of different
management methods are difficult to generalise because it depends a lot on how the management is
carried out, and different species respond in different ways to management.

However our review has highlighted some broad management issues which are important for
biodiversity regardless of context. These issues relate quite closely to the five guiding principles of
Lindenmayer et al. (2006) for biodiversity conservation in multi-purpose forests.

1. the maintenance of connectivity (i.e. addressing the effects of habitat fragmentation);

2. the maintenance of landscape heterogeneity;
3. the use of natural disturbance regimes to guide human disturbance regimes (i.e thinning and
cutting regimes)
4. the maintenance of stand structural complexity (including taking account of the impact of
associated specific stand management operations such as drainage; cultivation etc);
5. the maintenance of aquatic ecosystem integrity;

With regard to the sustainability of the Forestry Wood Chain it is important also to consider the
specific role of the intensive forest management and lingo-culture production alternatives (4 and 5 in

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Table 1.3), which are somewhat out with the envelope of multi-purpose forestry since there is an
overriding focus on timber or biomass production.

2.4.1. Intensive forest management

As far as we know there are few examples of rare or threatened species which are able to survive in
very intensively managed forest plantation systems. In general the number of species is very low and
is dominated by a few species with general habitat requirements. Normally, at the stand level the
conservation value of these areas is very low. On the landscape level the significance of intensively
managed forest plantation depends on how large and area they cover and what kind of habitats have
been removed. If they cover large areas and natural forest has been removed it might be a threat
against many species. On the other hand if small areas are covered and mainly intensive agricultural
fields are used for the plantation there are no threats against biodiversity. Some people even argue that
intensively managed forestry plantation benefit biodiversity because by concentrating timber
production to these areas other more natural forests can be less intensively managed.

2.4.2. Habitat loss, fragmentation and restoring habitat networks and

connectivity
Some of the biggest threats to biodiversity are habitat loss, leading to fragmentation, leading to
dispersal- and genetic problems. Examples of important habitat types in the natural forest which are
lacking or fragmented in the present forestry landscape are uneven-aged stands, old-growth forests and
forests with continuous tree cover. In general, species at most threat from habitat loss and
fragmentation have very special habitat requirements and poor dispersal ability. These requirements
can be difficult or even impossible to combine with forestry. Some examples are species demanding
very high abundance of dead wood, old trees, big trees or specific natural disturbances. The only
possibility to conserve these species is to conserve their habitat and the processes creating the right
conditions. Normally this is only possible by strict protection through nature reserve or national parks.
In order to conserve these types of species a network of such protected areas in the landscape covering
different types of habitats is important. The area needed for protection depends on a number of factors:
type of land-use in other parts of the landscape, the spatial distribution of high-quality forest patches,
and the life-history of the species in the protected area.

The dispersal capabilities of threatened species can also be enhanced by management to increase
landscape heterogeneity and improve the “permeability” of the matrix between habitat patches.
Invariably the matrix forms the bulk of the managed forest and there is scope here for modifying
management to improve hospitability of the matrix to red data species. For example, leaving legacies
on clear-fells or doing variable density thinning. In theory, catering for red list species should ensure
that wider diversity is also catered for.

Habitat restoration plays a key role in reversing fragmentation, but is only useful if some habitat
qualities still exist and there are source of species in the landscape. Therefore restoration is most
efficient near hot-spots and for connecting different hot-spots to each other. However, if the continuity
of qualities is lost and the species is extinct, restoration is ineffective and it will take very long time
until the species return (if ever). In order to find hot-spots many different types of indicators have been
suggested. However, very few have been scientifically evaluated. To find suitable restoration level
threshold values of the abundance of critical resources have been suggested, but so far too few
examples exist to make this operational, and most examples are only valid in a specific group of
species in a specific geographical situation. Further development and scientific evaluation is needed.

2.4.3. Landscape planning to improve heterogeneity and connectivity

There is no single prescription for a stand that will maximise biodiversity, different species groups
have different requirements therefore landscape level solutions are required. Nor is it always possible
to combine forestry for timber production and conservation on the stand level. In addition, many
species are wide ranging and rely on threshold amounts, and/or specific spatial configurations of
habitat at the landscape scale to ensure persistence over time.

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In essence, to conserve biodiversity there must be a focus on creating and managing a landscape in
which species can find suitable habitats and dispersal possibilities over a long-time frame. Forestry
and other forms of land use alter the distribution and condition of habitats, but on the landscape level it
is possible to conserve habitat and key structures whilst retaining economic productivity. Different
tools can be combined such as: strict reserves, voluntarily reserves, conservation agreements, adapted
management, and forestry with conservation considerations. Often some kind of economic
compensation is paid to the landowner. In these examples also other values, such as cultural history
and social values have been added. Basically there are two main approaches:

1. Establishment of large areas of national parks or nature reserves (often 20-30% of the land area) in
which there is no forestry, combined with intensive management outside the reserves.
2. A combination of relatively small nature reserves and different types of extensive or intensive
forestry. In order to create an ecological infrastructure the area of conserved habitats and general
conservation considerations are related. The larger area that is conserved the smaller is the need
for general conservation considerations and vice versa.

There is also some evidence to suggest that management regimes should be varied at the landscape
scale simply to improve general structural heterogeneity. However, careful planning is required to
ensure a balance between this aspiration and the need to ensure conservation of specific priority
species and spatio-temporal connectivity of their habitats. One of the key considerations is the spatial
and temporal arrangement of harvest units and the different types of silvicultural regimes (e.g. clear
cutting versus low impact regimes; short rotations versus long rotations etc.). Natural disturbance
regimes can offer a template for informing the spatial and temporal arrangement of different types of
management.

2.4.4. Mimicking natural disturbance regimes

Both natural and anthropogenic disturbances have varied during history, and the result of this is a
dynamic ecosystem, which can be very difficult to restore or conserve. Lack of management at the
same time as natural disturbances are prevented (e.g. fire suppression in nature reserves) might result
in a loss of biodiversity. However, as knowledge of the importance of natural disturbance for
conserving biodiversity has increased there are an ever increasing range of examples where natural
disturbance is being used to guide the spatial and temporal distribution of silvicultural regimes across
the landscape as well as informing the creation of structural diversity within stands. In the former the
general approach is to reflect a gradient of increasing severity and scale of disturbance, and to mimic
different types of disturbance. In large landscapes this can lead to the use of prescribed burning of
varying intensity in different parts of the landscape and a mix of clear- cutting and low intensity
silviculture to reflect variability in the spatial distribution of stand-replacement disturbance regimes
and gap-phase disturbance regimes.

2.4.5. The maintenance of stand structural complexity

By using some adapted forestry methods it seems to be possible to combine conservation with forestry
for the majority of species including several of the red-listed species. Most important is to mimic the
structures and habitat that naturally are created by natural disturbances, which means retaining key
elements of stand structural complexity (Lindenmayer et al., 1996). These elements include structures
such as dead wood, old trees, big trees, under storey vegetation etc. Structural complexity can be
catered for throughout the “normal” crop rotation or by extending rotations. There is good evidence
that clear-cutting combined with structural retention can cater for a wide range of different taxi.
Similarly, variable density thinning can be used in young and middle-aged stands to improve structural
diversity and spatial heterogeneity, and retain dead and dying trees. Smaller biotopes such as wet
flushes, boggy areas etc can also be left undisturbed in larger stands whilst also carrying out stand
management operations.

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Measures to improve structural diversity within stands can affect economic yield, in that the most
efficient harvesting method may not used and some trees are left etc. However, there are many
examples from different areas in Europe showing that this kind of forestry can still be profitable. Often
mechanised operations such as harvesting can be directed to avoid key biotopes and this can save time
and money. For example, tree growth can be reduced in wet areas and hence the crop may be of
reduced value and not worth harvesting.

Whilst the soil and vegetation disturbance associated with harvesting operations and cultivation prior
to establishment can have a negative impact on both above and below ground biodiversity, these
impacts can be relatively short-lived compared to more permanently damaging activities such as
drainage or road building. Roads can be planned to avoid damaging key biotopes, and current forestry
and nature conservation legislation offers protection for aquatic habitats in most countries.

2.4.6. The maintenance of aquatic ecosystem integrity

Lindenmayer et al. (2006) emphasize the importance of maintaining aquatic ecosystem integrity, and
in our review we highlighted the importance of buffer strips for protecting aquatic ecosystems from
the impact of forest operations. As mentioned in the previous section, aquatic ecosystems should be
protected by legislation due to their importance in maintaining and delivering ecosystem services such
as flood control, drinking water, fish production as well as biodiversity. However, careful inventory
and planning is needed to ensure that the conservation of aquatic ecosystems is maintained alongside
forestry operations.

2.4.7. Biodiversity indicators

The indicators selected by the EFORWOOD project for assessing the impacts of different forest
management scenarios on biodiversity are listed in Table 1.1. Here we focus on the indicators in the
second column in Table 1.1. The rationale for selection has been given in the document PD2.2.1 and
can be summarized here. Tree species composition is measured in terms of the number of tree and
shrub species. In general, as this number goes up, there is an increase in the number of different niches
for dependent flora and fauna. This indicator should be of particular value in assessing changes to the
Multifunctional forest management alternative (Table 1.3), where diversifying tree species is often a
key objective of management. Forest continuity is an indicator of long-term provision of habitat
required by a range of rare forest species. Continuity can be measured by the area and historical
continuity of woodland key habitats (e.g. in the Close to nature forest management alternative) and
also at the large scale by the overall area of forest nature reserve. Deadwood is a key indicator in all of
the forest management alternatives except the wood-biomass alternative, where by definition there can
be no deadwood. Presence of key habitat such as remnants of old-growth and wet forest is an
important indicator in the Close to nature and Multi-functional forest types. Due to the statutory
framework in most countries, authentic water regime (measured by changes in ditches and irrigation)
is a key indicator in all the forest types in Table 1.3.

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