Forest Ecology and Management 260 (2010) 939–947

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Forest Ecology and Management

journal homepage: www.elsevier.com/locate/foreco

Review

Replacing coniferous monocultures with mixed-species production stands: An

assessment of the potential benefits for forest biodiversity in northern Europe
Adam Felton a,∗ , Matts Lindbladh a , Jörg Brunet a , Örjan Fritz b
a
Southern Swedish Forest Research Centre, Swedish University of Agricultural Sciences, Rorsjovagen 1, Box 49, SE-230 53 Alnarp, Sweden
b
Naturcentrum, Strandtorget 3, 444 30 Stenungsund, Sweden

a r t i c l e i n f o a b s t r a c t

Article history: Conifer dominated plantations in central and northern Europe are associated with relatively low eco-
Received 26 February 2010 logical values, and in some cases, may be vulnerable to disturbances caused by anthropogenic climate
Received in revised form 3 June 2010 change. This has prompted the consideration of alternative tree species compositions for use in pro-
Accepted 10 June 2010
duction forestry in this region. Here we evaluate the likely biodiversity costs and benefits of supplanting
Norway spruce (Picea abies) monocultures with polycultures of spruce and birch (Betula spp.) in southern
Keywords:
Sweden. This polyculture alternative has previously been evaluated in terms of economic, recreational,
Conservation
and silvicultural benefits. By also assessing the ecological implications we fill a gap in our understanding
Plantation
Forest management
of the range of socio-ecological benefits that can be achieved from a single polyculture alternative. We
Biodiversity project likely broad scale changes to species richness and abundance within production stands for five
Risk-spreading taxonomic groups including ground vegetation, tree-living bryophytes, lichens, saproxylic beetles, and
birds. Our research leads us to three key findings. First, the replacement of spruce monocultures with
spruce–birch polycultures in the managed forest landscapes of southern Sweden can be expected to result
in an increase in biological diversity for most but not all taxa assessed, but it is unlikely to improve con-
ditions for many red-listed forest species. Second, modification of other aspects of forest management
(i.e. rotation length, dead wood and green tree retention, thinning regimes) is likely to contribute to
further biodiversity gains using spruce–birch polycultures than spruce monocultures. Third, the paucity
of empirical research which directly compares the biodiversity of different types of managed produc-
tion stands, limits the extent to which policy relevant conclusions can be extracted from the scientific
literature. We discuss the wider implications of our findings, which indicate that some climate change
adaptation strategies, such as risk-spreading, can be readily integrated with the economic, environmental
and social goals of multi-use forestry.
© 2010 Elsevier B.V. All rights reserved.

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 940
2. Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 941
3. Results and discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 941
3.1. Ground vegetation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 941
3.2. Tree-living bryophytes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 942
3.3. Epiphytic lichens. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 942
3.4. Saproxylic beetles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 943
3.5. Forest birds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 943
4. Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 944
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 945
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 945

∗ Corresponding author. Tel.: +46 40 415471; fax: +46 40 462325.

E-mail address: adam.felton@ess.slu.se (A. Felton).

0378-1127/$ – see front matter © 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.foreco.2010.06.011
940 A. Felton et al. / Forest Ecology and Management 260 (2010) 939–947
1. Introduction
There is an increasing body of scientific literature which
addresses the potential benefits of mixed-species production
forests over monocultures (Hartley, 2002; Lindenmayer and Hobbs,
2004; Nichols et al., 2006; Knoke et al., 2008; Jactel et al.,
2009). Potential benefits include improved habitats for biodiversity
(Lindenmayer and Hobbs, 2004; Carnus et al., 2006; Brockerhoff et
al., 2008), advantages in relation to growth rates (Piotto, 2008),
increased recreational value (Norman et al., 2010), improved soil
conditions (Brandtberg et al., 2000), reduced damage to focal
tree species by grazers, pest insects, and fungal pathogens (Jactel
and Brockerhoff, 2007), and reduced risk of damage from wind
(Schutz et al., 2006) and fire (Gonzalez et al., 2006). The poten-
tial for mixed-species stands to provide multiple services to a
wide variety of end users, while concurrently reducing the risk
of abiotic or biotic impacts on production (Knoke and Seifert,
2008), has lead to policy initiatives in central and northern Euro-
pean countries advocating the conversion of some coniferous
monocultures to mixed-species forests (SCCV, 2007; Knoke et al.,
2008).
The actual extent to which ecological, economic and societal
benefits can arise from the conversion of coniferous monocul-
tures to mixed-species forests depends on: (1) the tree species’
being replaced and promoted, (2) the bio-geographical region, and
(3) associated adjustments to management practices. The benefits
from such conversions are therefore entirely context dependent,
and require regional scale assessments to ensure relevance to
local forest managers and policy makers. More importantly, due
to the rarity of interdisciplinary assessments of particular poly-
culture alternatives, once conducted such studies can serve as
internationally relevant case studies of the extent to which multi-
ple socio-ecological benefits are possible from specific polyculture
alternatives. Due to the widespread use of coniferous monocultures
in southern Sweden, and numerous studies of these forest systems
and potential polyculture alternatives (Enander, 2007), this region
provides ideal conditions for such analyses.
The production of timber and other forest products in south-
ern Sweden is primarily based on the rotational clear-cutting of
even-aged stands. Over the last hundred years this form of forest
management and other human associated impacts have dramati-
cally altered tree species composition in southern Sweden (Nilsson,
1997; Lindbladh et al., 2000). Norway spruce (Picea abies, hereafter
spruce) has benefited most from this alteration of the southern
Swedish landscape, to the extent that it composes approximately
60% of forested area (SFA, 2008). Forest managers have promoted
spruce due to its relatively simple management, early returns
from thinning, high wood production, short rotation periods, rel-
atively low susceptibility to browsing pressure from ungulates
(Carbonnier and Hägglund, 1969; Kullberg, 2000), favorable market
demand, and the development of a “spruce culture” among forest
managers (Felton et al., 2010a).
The expansion of spruce dominated monocultures has however
been at the expense of deciduous tree species, which previously
covered much of southern Sweden (Lindbladh et al., 2000). This
has contributed to widespread population declines and increased
extinction risk for many forest dependent taxa in Sweden (Berg et
al., 1994; Fridman, 2000; Gärdenfors, 2005; Chapin et al., 2007).
Spruce monocultures have also led to an increase in the acidity
of some soils, with long-term associated ramifications for biodi-
versity and subsequent land-uses (Nordborg and Olsson, 1999;
Jönsson et al., 2003). Furthermore, there are indications that cli-
mate change will detract from the long-term viability of spruce
monocultures in southern Sweden. A range of climate simulation
models suggest that by the end of this century increased tem-
peratures, altered precipitation and changes to the frequency and
intensity of extreme disturbance events (SFA, 2006; Christiansen
and Hewitson, 2007; SCCV, 2007) are likely to reduce the eco-
logical suitability of spruce monocultures in the south of the
country (Sykes and Prentice, 1996; Sykes et al., 1996; Bradshaw
et al., 2000; Koca et al., 2006). This is due to the tree species’
relatively high susceptibility to drought and wind damage, and
the favored reproductive capacity for two spruce associated pest
species, bracket fungus Heterobasidion annosum (SCCV, 2007) and
spruce bark beetle Ips typographus (Jonsson et al., 2007; Bolte et al.,
2010).
Because spruce monocultures can have negative impacts on the
environment, and in-turn, may be less resistant to abiotic and biotic
disturbances exacerbated by climate change (Jactel et al., 2009),
there is a strong impetus to evaluate alternative tree species for use
in production forestry in southern Sweden (SCCV, 2007). Increasing
the amount of forest allocated to a wider range of tree species in this
region is expected to reduce the vulnerability of the entire system
to climate change by spreading the risk among different tree species
alternatives (Ask, 2002; Fransila et al., 2005; Eriksson, 2007; SCCV,
2007; Sveaskog, 2007; Felton et al., 2010a). One such alternative
involves the replacement of a proportion of spruce monocultures
with mixed-species stands composed of planted spruce and nat-
urally regenerated silver or downy birch (Betula pendula and B.
pubescens, hereafter birch), with birch providing between 30% and
70% of the stand’s basal area.
There are several potential benefits from adopting spruce–birch
polycultures as a risk-spreading alternative to spruce monocultures
in the region (SCCV, 2007). First, birch is the most prevalent nat-
urally regenerated tree species in southern Sweden (Anonymous,
2000; Karlsson, 2001), and over 70% of its sapling populations are
found within conifer dominated forests (Götmark et al., 2005).
Second, experimental trials suggest that mixtures of spruce and
birch can be economically competitive with spruce monocultures,
both in terms of volume production and resultant economic value
(Karlsson, 2001; Agestam et al., 2005). Third, the knowledge base
for spruce–birch mixtures is better than that for other potential
tree species mixtures (Agestam et al., 2005). Fourth, the use of
a broadleaved tree as a component of the mixture increases the
potential recreational value of the stand, thereby providing a non-
market incentive for its adoption (Mattsson and Li, 1994; Bostedt
and Mattsson, 1995; Norman et al., 2010). Fifth, the inclusion of a
broadleaved tree species as a component of the mixture may reduce
the risk of wind (Lekes and Dandul, 2000; Schutz et al., 2006) and
pest species damage to the stand (Jactel and Brockerhoff, 2007;
Jactel et al., 2009), Thus, there are multiple financial and societal
indications as to why spruce–birch polycultures are a feasible alter-
native to spruce monocultures in this region. However, there has
been no detailed assessment of the potential implications for forest
biodiversity.
Here we examine the likely biodiversity costs and bene-
fits arising from the replacement of spruce monocultures with
spruce–birch polycultures in the conifer dominated landscapes of
southern Sweden (Fig. 1). We concentrate on some well-known
taxonomic groups representing a diverse range of trophic levels,
taxonomic affinities, and ecological niches, and for which their dif-
ferent habitat requirements operate at distinct spatial scales (see
Holling, 1992). We assess the peer-reviewed scientific literature
for relevant studies of five forest-associated taxonomic groups for
which contributing authors have extensive research experience;
ground vegetation, tree-living bryophytes and lichens, saproxylic
beetles and birds. We discuss biodiversity value in terms of species
richness, species abundance, and more specifically in regard to taxa
present on the Swedish Red List (Gärdenfors, 2005). We also discuss
the wider implications of our research with regard to the poten-
tial for integrating multi-use forestry goals with risk-spreading
approaches to climate change adaptation.
 A. Felton et al. / Forest Ecology and Management 260 (2010) 939–947
Fig. 1. Map identifying the location of Sweden relative to Europe, with the region of Götaland (southern Sweden) indicated.
2. Methods
We searched electronic databases and the Internet using dif-
ferent combinations of Boolean search-terms. The databases used
were Google Scholar (http://scholar.google.com.se/) and Web of
Science (http://www.isiwebofknowledge.com/). We used, as an
example, the following search-terms for locating relevant stud-
ies on birds: (birch* OR betula OR spruce OR “Picea abies”)
AND bird*. The use of relatively few search constraints increases
the number of results, of which any studies potentially rel-
evant to our inquiries were retained after appraisal of titles
and abstracts. The benefit of this approach is that it avoids
the a priori use of narrowly defined geographical (e.g. Swed*,
Europe*) or management terms (e.g. plantation*, “production for-
est”), which increases the likelihood of relevant studies being
missed. We also obtained papers from colleagues and through
reference lists from published studies, including major review arti-
cles and books on managed production forests and biodiversity.
Furthermore, we obtained information from government stud-
ies and reports. Where possible we refer to studies conducted
within southern Sweden, or the Fenno-Scandinavian region, due
to their increased bio-geographical relevance to the issues being
addressed.
In our assessment of the scientific literature we refer to rel-
evant examples of how some species respond to the types of
environments provided by spruce monocultures or spruce–birch
polycultures, but we purposefully avoid using such observations to
make projections for individual species. Instead we concentrate on
projecting the likely net response of major taxonomic groups using
common biodiversity metrics (i.e. species richness and abundance).
By doing so we avoid embedding the discussion in the diverse and
highly individualistic responses of species to differences in site
productivity, management practices, landscape context, climatic
influences (current and projected), and other disparate or synergis-
tic potential influences. Providing our conclusions as generalized
likely responses for major taxonomic categories, and at the scale
of species richness or abundance, also provides outcomes that are
likely to be more readily disseminated to policy makers and other
interested parties (see Gibbons et al., 2008).
941
3. Results and discussion
3.1. Ground vegetation
Ground vegetation is strongly influenced by the composition
and structure of the overstorey, via its influence on temperature,
light and water, soil nutrients (Saetre et al., 1999), and on the
physical characteristics of the litter layer (Barbier et al., 2008).
For example, the ground layer in mixed spruce–birch stands has
less acidified soil, litter, and humus conditions relative to stands
composed of pure spruce (Brandtberg et al., 2000; Brandtberg and
Simonsson, 2003; Brandtberg and Lundkvist, 2004). Furthermore,
the humus and litter layers of mixed spruce–birch stands in central
and southern Sweden are found to have higher concentrations of
minerals, such as calcium and magnesium (Brandtberg et al., 2000).
The litter layer of spruce–birch stands also possess higher organic
matter quality than under spruce monocultures, as expressed by
carbon:nitrogen ratios (see Brandtberg and Lundkvist, 2004). For
these reasons, there are clear justifications for expecting differ-
ences in the community composition of ground vegetation found
in monocultures of spruce, versus polycultures of spruce and birch.
The ground vegetation of spruce monocultures found in
southern Sweden usually consists of a mixture of herbs, dwarf
shrubs, grasses, ferns and mosses (Nihlgård, 1970; Engelmark
and Hytteborn, 1999), but depends on the age, management, and
soil conditions of the stand considered. Most of these species
are widespread and relatively common throughout the region.
In southern and central Sweden comparisons have been made
of the ground vegetation found within spruce monocultures and
mixed-species stands of spruce and birch (Saetre et al., 1997). The
mixed-species stands had a higher percentage cover and diversity
of herbs, but a lower percentage cover and diversity of bryophytes,
potentially due to the leaf litter from birch.
Other research has assessed the influence that individuals of
birch or spruce trees, and their densities, can have on the ground
vegetation beneath their canopies. Within mature mixed-species
stands in central Sweden, the percentage ground cover of herbs
and some species of moss (e.g. Brachythecium reflexum) decreased
with increasing densities of spruce trees (Saetre, 1999). This was
942 A. Felton et al. / Forest Ecology and Management 260 (2010) 939–947
probably due to the associated decreases in light availability and
soil water content. Likewise, research has been conducted within
primarily coniferous forests, assessing the influence of birch trees
on ground vegetation (Wallrup et al., 2006). Birch trees were associ-
ated with a higher species richness of ground vegetation than their
spruce counterparts, with benefits afforded to species spanning a
range of taxonomic groups including vascular plants, shrubs, and
grasses.
We conclude that species richness and cover of the herbaceous
layer, the herbs and grasses in particular, are likely to benefit from
the increased presence of birch as part of spruce–birch polycul-
tures. However, due to different habitat requirements, we do not
expect any benefits for the 75 species of vascular forest plants
red-listed in Sweden (Gärdenfors, 2005). Likewise, the few red-
listed bryophytes which live on the forest floor, mainly occur in
other forest types than spruce or birch forest (Gärdenfors, 2005).
Also, because the cover of ground-living bryophytes appear to be
inversely related to the percentage cover of birch in spruce stands, it
is possible that any benefits to the herbaceous vegetation may come
at the expense of ground-living bryophyte diversity and abun-
dance.
3.2. Tree-living bryophytes
There are approximately 300 species of mosses and liver-
worts living on tree bark in Swedish forests, of which 23% are
red-listed (Weibull and Rydin, 2005). Studies of these bryophyte
groups have generally focused on the communities found within
natural old-growth forests (e.g. Cleavitt et al., 2009), whereas
few studies have been conducted in production forests (but see
Kruys et al., 1999; Coote et al., 2008; Lohmus and Lohmus,
2008). To our knowledge, no European study has been con-
ducted within polycultures of spruce and birch. Therefore, we used
studies from managed spruce monocultures and temperate conif-
erous/deciduous mixed forests to assess the potential of mixed
spruce–birch stands to influence tree and rock living mosses and
liverworts.
Humidity and substrate pH are important determinants of
the species richness and composition of bryophyte communities
(Barkman, 1958; Wiklund and Rydin, 2004; Lohmus et al., 2007;
Coote et al., 2008; Tinya et al., 2009). For this reason bryophytes
communities appear to benefit from the high humidity associated
with closed forest stands (Barkman, 1958; Coote et al., 2008), as it is
commonly found in spruce monocultures. Counteracting this trend,
the acidic bark conditions associated with spruce (Hallingbäck,
1996; Kuusinen, 1996) and birch (Barkman, 1958) tend to suppress
the diversity of their bryophyte communities. Only 30 species of
bryophyte are found on the bark of birch trees, of which five species
are present on the Swedish Red List (Hallingbäck, 1996). Notably,
all bryophytes found on birch can also be found on spruce (Berg et
al., 2002), whereas there are larger numbers of species (50) which
are found on spruce but not birch. Dead coniferous tree species are
however very important for some bryophyte species (85 species),
including a number of red-listed taxa (20 species) found on their
logs in various states of decay.
It is likely that some species of tree-living bryophyte will be
disadvantaged by the micro-climates found in spruce–birch poly-
cultures. As the bryophyte communities found on birch are a subset
of those found on spruce, the species richness of spruce–birch poly-
cultures may therefore not be higher than that found in spruce
monocultures. However, if bryophyte species exhibit a range of
distinct preferences for the different environmental conditions pro-
vided by spruce and birch host trees, this could result in an increase
in bryophyte species richness due to the increased diversity of habi-
tats provided in spruce–birch polycultures. Whether or not the
strength of this effect will be sufficient to offset the decreased suit-
ability of the micro-climate for some species is unknown, but will
likely depend on the specifics of the bryophyte species pool avail-
able and the percentage cover and spatial configuration of birch
trees within the stand.
3.3. Epiphytic lichens
In Sweden 82% of the 254 red-listed lichen species are forest-
associated species (Gärdenfors, 2005), thereby providing a strong
incentive to assess the relative impact of production forestry alter-
natives on lichen species. There is a rather rich body of literature
concerning epiphytic lichens in old-growth natural or semi-natural
forests, from boreal coniferous as well as temperate deciduous
forests, and often presented in relation to more managed coun-
terparts (Kuusinen and Siitonen, 1998; Nascimbene et al., 2007).
However, the epiphytic lichens of production forests are gener-
ally poorly studied (but see Coote et al., 2008; Hilmo et al., 2009).
As a result we could not find any studies of epiphytic lichens in
mixed polycultures of spruce and birch. To assess the implications
of mixed plantations of spruce and birch for epiphytic lichens, we
used existing studies principally from managed spruce monocul-
tures, and from managed as well as unmanaged birch forests.
Under natural to semi-natural conditions there are approxi-
mately 140 lichen species which can be found in birch forests,
whereas 250 species can be found in spruce forest (Berg et al., 1995).
Relative to most other tree species, there are a relatively small num-
ber of red-listed lichens associated with birch, with no red-listed
species exclusive to birch (Arup et al., 1997; Thor, 1998; Uliczka and
Angelstam, 1999; Berg et al., 2002). Several factors are important
for the establishment and growth of epiphytic lichens in forests,
including humidity, understorey light environments, bark pH, and
bark structure (Hilmo et al., 2009). Under normal conditions both
birch and spruce possess bark with acidic and rather nutrient-poor
substrate conditions, hence the relatively limited lichen flora asso-
ciated with spruce and birch. This overlap in their bark properties
also leads to a corresponding similarity in community composition
(Barkman, 1958), with both spruce and birch possessing mostly
generalist species (Kuusinen, 1996). Notably, as birch can reach
maturity relatively early, with a corresponding rapid development
of rough bark (Uliczka and Angelstam, 1999), stands including this
tree species may provide more suitable substrate conditions for
epiphytic lichens earlier in the rotation than are found in spruce
monocultures.
Spruce plantations and birch forests also differ in their associ-
ated light and humidity environments (Kuusinen, 1996), resulting
in some differences in epiphytic composition (Neitlich and McCune,
1997; Hilmo and Sastad, 2001). In even-aged coniferous monocul-
tures reduced light availability is an important negative influence
on the growth and vitality of photosynthesizing organisms, such
as lichens (Gauslaa et al., 2007). In contrast, birch forests generally
allow more light to reach the forest understorey, which is often ben-
eficial to lichen communities. This combination of factors, substrate
quality and micro-climate, can result in lichen species richness
being higher in mixed tree species stands than in pure birch and
spruce stands (Wannebo-Nilsen et al., 2010).
In summary, spruce–birch polycultures are likely to provide
different habitat conditions for epiphytic lichens relative to those
found in spruce monocultures. This is due to distinctions between
spruce and birch in the rate and type of development of bark crenu-
lation, and in the light and humidity of forest stands created using
these species. These differences may facilitate an increase in species
richness despite the large overlap in their respective lichen com-
munities. Nevertheless, it is unlikely that birch–spruce polycultures
will host epiphytic lichen species of significant conservation con-
cern.
 A. Felton et al. / Forest Ecology and Management 260 (2010) 939–947
3.4. Saproxylic beetles
Saproxylic (wood living) beetles are a species rich group that
have been negatively affected by the intensive management of
forests in Sweden, and associated declines in the availability of dead
wood habitats (Berg et al., 1994; Jonsell et al., 1998; Gärdenfors,
2005). Due to these impacts, over 500 species of saproxylic beetle
are on the Swedish Red List (Jonsell et al., 1998). The importance
of dead wood retention during forest management is particularly
relevant to their populations, as although some species inhabit
living trees, most saproxylic beetles are associated with dead
wood in various states of decay (Dahlberg and Stokland, 2004).
For this reason, assessing the biodiversity value of spruce–birch
polycultures for saproxylic beetles is only meaningful with a con-
comitant expectation that sufficient dead wood will be retained
during the stand’s rotation. This expectation is to some extent
justifiable in Sweden as dead wood and green tree retention is
required after final felling by Anon. (1993) and the certification
schemes of FSC (FSC, 2000) and PEFC (PEFC, 2000), which are
adopted in 80% of all Swedish production forests. It is within
this context that we assess the likely biodiversity benefits for
saproxylic beetles from the increased use of spruce–birch polycul-
tures.
There are distinct differences in the saproxylic beetle commu-
nities associated with the dead wood of coniferous and deciduous
tree genera, and by default, between spruce and birch (Jonsell et
al., 1998; Dahlberg and Stokland, 2004). The distinctiveness of their
fauna results from differences in the physical properties of spruce
and birch wood, and from differences in their fungal flora com-
munities, which help to determine wood properties during the
process of decay. Recent estimates suggest that 466 saproxylic bee-
tle species are associated with birch, the majority of which (304
spp.) are not normally found in spruce (Dahlberg and Stokland,
2004). This provides a strong justification for expecting higher
species richness of saproxylic beetles in spruce–birch polycultures,
than in spruce monocultures. However, as only 8% of species asso-
ciated with birch are found on living trees, rather few species of
saproxylic beetle would benefit from stands of birch during the
first rotation. The principle benefit is expected after final felling
due to the expected increase in dead wood from birch or dying
birch trees. As the majority of birch associated species are found
in wood of dimensions smaller than that associated with normal
production stands, stem diameters should not be a limiting fac-
tor for most species of saproxylic beetle (Dahlberg and Stokland,
2004).
On Swedish clear-cuts it is a standard practice to intentionally
create high-stumps. In a recent study of the saproxylic beetle fauna
associated with these stumps it was shown that 19 species were
unique to birch, a further 17 species unique to spruce, with 30
species found on both tree species (Abrahamsson and Lindbladh,
2006). Another comparative study of fauna associated with high-
stumps found that spruce stumps hosted more species than birch
in total, but a similar number of red-listed species (Lindhe et al.,
2004). In regard to other sources of dead wood, in a study conducted
on 3–5-year-old clear-cuts it was shown that logging residues from
harvested birch hosted significantly more saproxylic beetle species
than spruce residues (Jonsell et al., 2007).
In summary, the use of spruce–birch polycultures should be
beneficial for many saproxylic beetles, both in terms of saproxylic
beetle species richness and abundance. However, the degree of ben-
efit will be largely dependent on the extent to which dead wood of
the different tree species is retained throughout the stands rotation.
If source populations for red-listed taxa are available, sufficient
dead wood retention of birch and spruce could result in provid-
ing valued habitat for saproxylic beetles of significant conservation
concern.
943
3.5. Forest birds
There are approximately 100 bird species breeding in the forests
of southern Sweden (Gärdenfors, 2005). Of these species, approx-
imately 30% have exhibited population declines during the last 10
years, and 18 are threatened (Gärdenfors, 2005). There is a large
body of scientific literature assessing the habitat requirements of
forest birds, and the impact of production forest management on
community composition and species populations. However, we are
not aware of any published studies which compare bird species
composition in mature managed stands of spruce with similarly
aged and managed stands of spruce and birch in Europe.
Very few bird species in Europe are dependent on a particular
tree species (Nilsson, 1997). In contrast, bird species often exhibit a
preference for particular categories of trees (i.e. coniferous versus
broadleaved), or particular compositions of tree species at appro-
priate spatial scales (Bibby et al., 1989; Peck, 1989; Smith, 1992).
For this reason we do not expect the creation of spruce–birch poly-
cultures to be coupled with a general shift towards ‘birch associated
bird species’, as could occur in other taxa (e.g. saproxylic beetles).
One possible exception is the black grouse (Tetrao tetrix), which
can rely on birch leaf buds as a food source during winter (Nilsson,
1997), and may therefore specifically benefit from the increased
prevalence of birch in some regions of southern Sweden. Less obli-
gate associations with birch can also be observed in context specific
circumstances, such as a preference for foraging in birch exhib-
ited by blue tits (Cyanistes caeruleus), great tits (Parus major) and
the Eurasian treecreeper (Certhia familiaris) (Peck, 1989), and the
preference exhibited by some species of woodpecker for birch as
a nest tree species (Hagvar et al., 1990; Poulsen, 2002). Neverthe-
less, we expect that the majority of changes to bird communities
arising from the creation of spruce–birch polycultures will result
from floristic and structural changes associated with growing a
broadleaved species within conifer dominated stands and land-
scapes (Uliczka and Angelstam, 2000).
The bird communities of broadleaved forests are often higher
in species richness and abundance than those found within conif-
erous forests (Adams and Edington, 1973; Archaux and Bakkaus,
2007; Berg, 1997; Tomialojc and Wesolowski, 1990; Uliczka and
Angelstam, 2000), although there are relevant exceptions (Adams
and Edington, 1973; Donald et al., 1998). More importantly, many
bird species exhibit a direct association with broadleaved or conif-
erous forests, thereby causing an important divergence in the
composition of bird communities associated with either forest type
(Bibby et al., 1989; Nilsson, 1997; Roberge and Angelstam, 2006).
This preference is sufficiently strong to enable the categorization
of bird species as conifer specialists, broadleaved specialists, or
even specialists in the use broadleaf–conifer mixtures (Roberge
and Angelstam, 2006). Of direct relevance to our inquiries, sev-
eral studies have shown that the bird communities within mixed
stands of conifers and broadleaves are composed of a rich mixture
of bird species representing both the broadleaved and coniferous
associated fauna (Donald et al., 1998; Hausner et al., 2002).
Researchers have also been able to quantify the specific min-
imum percentages of deciduous or coniferous trees within the
stand or landscape necessary to facilitate the occurrence of some
bird species. For instance, the long-tailed tit (Aegithalos caudatus)
is most commonly found in landscapes with at least 15% cover
of deciduous trees of suitable age classes within the landscape
(Uliczka and Angelstam, 2000). Likewise, coal tit (Parus ater) and
goldcrest (Regulus regulus) typically occur in forest plots with at
least 10% spruce (Nilsson, 1977). As there has been a substan-
tial replacement of broadleaved dominated forests with conifer
monocultures in the southern Swedish landscape over the past
century (Nilsson, 1979, 1997; Enoksson et al., 1995; Lindbladh and
Bradshaw, 1998), the addition of birch to spruce monocultures is
944 A. Felton et al. / Forest Ecology and Management 260 (2010) 939–947
Table 1
Summary of expected changes to species richness and abundance for taxa assessed. Score represents both the direction of change and the level of confidence authors had
in the expected outcome. Outcomes are graded in terms of positive results “+”, negative outcomes “−”, or neutral outcomes “0”, with brackets “( )” indicating that the result
relies on other management activities to ensure the expected probability of outcome (i.e. dead wood retention). Confidence levels are provided as three categories (i.e. +, ++,
or +++) representing “possible”, “probable”, and “highly probable” respectively.
Taxa Species richness
Ground-living herbs +++
Bryophytes—ground-living −−
Bryophytes—tree-living 0 −
Lichens + ++
Saproxylic beetles (+++)
Birds +++
therefore very likely to contribute positively to avian diversity in
southern Sweden.
With regard to projecting potential changes to bird species
richness, abundance, or community composition, some impor-
tant caveats are warranted. There are multiple forest management
issues which will contribute or detract from the potential benefits
to bird communities from promoting spruce–birch polycultures.
For instance, sallying insectivores are known to benefit from
the improved visibility and space provided in deciduous forests
(Hausner et al., 2002). Whether such conditions will be adequately
provided for this guild in the spruce–birch polycultures of south-
ern Sweden is as yet unknown, and would likely depend on the
proportion of birch in the mixture and its spatial distribution. Sim-
ilarly, the retention of mature deciduous trees after cutting can be
of benefit to secondary hole-nesters (species unable to excavate
their own nest holes), such as pied flycatchers (Ficedula hypoleuca)
and marsh tits (Parus palustris) (Soderstrom, 2009). But the extent
to which such factors will influence bird species composition in
spruce–birch polycultures depends on the extent to which green-
and dead-tree retention is adopted during harvesting (Rosenvald
and Lohmus, 2008; Soderstrom, 2009).
In summary, although few bird species can be considered depen-
dent on birch per se, the available evidence indicates that the
addition of more broadleaved tree cover within southern Sweden’s
extensive spruce monocultures should increase avian species rich-
ness and potentially abundance within these stands.
4. Conclusion
Our results indicate that the replacement of spruce monocul-
tures with spruce–birch polycultures should result likely increases
in species richness or abundance for the majority of taxa assessed
in southern Sweden (Table 1). There is sufficient evidence to expect
increases in species richness for birds and understorey vascular
plants, with a possible increase for lichens. The species richness
and abundance for saproxylic beetles are also likely to increase,
but only if adequate amounts of dead wood are retained during the
stands’ rotation. Bryophytes were the only taxa reviewed for which
declines in species richness or abundance was of concern. The avail-
able evidence indicates that this taxonomic group may be relatively
less diverse in spruce–birch polycultures due to associated reduc-
tions in humidity and a possible increase in competitive pressures
from more phytophylic plant species. However, we do not fore-
see such reductions in population density as an overriding issue of
conservation concern, as any decline can be considered relative to
artificially high population densities maintained by the unnatural
forest conditions provided within spruce monocultures. Empirical
research directly comparing the biodiversity of managed produc-
tion stand alternatives would enable these issues to be explored
further (see Stephens and Wagner, 2007).
Although an increase in species richness was projected for the
majority of taxa assessed, few red-listed species were expected to
benefit directly. There are a number of reasons why this was the
Abundance Habitat for red-listed species
+++ 0
−− 0
0
0
(+++) (+)
+ 0
case. First, greater benefits to red-listed taxa would have been pre-
dicted if oaks (Quercus spp.) or beech (Fagus sylvatica) had been
included in the mixed stands. These particular tree species pos-
sess ecological characteristics disproportionately beneficial to a
large number of red-listed species (Berg et al., 1994; Jonsell et al.,
1998), and have experienced dramatic reductions in forest cover
throughout much of southern Sweden (Nilsson, 1997; Lindbladh
et al., 2000; Lindbladh and Foster, in press). Second, unless produc-
tion forest guidelines change in Sweden, spruce–birch polycultures
will be managed under the same basis of rotational clear-cutting of
even-aged stands as occurs in spruce monocultures. Many forest-
associated species in Sweden require habitats that simply do not
persist in forests managed using these techniques (Berg et al., 1994,
2002; Bengtsson et al., 2000; Nilsson et al., 2005). Because of the
causal association between production forestry and the number of
forest species on the Swedish Red List, it is not surprising that few
red-listed species are expected to benefit in spruce–birch polycul-
tures. However, despite these inherent limitations, spruce–birch
polycultures do accommodate more suitable conditions for forest
biodiversity than spruce monocultures, and this raises the poten-
tial for augmenting these gains by altering relevant aspects of forest
management.
For example, the extent to which saproxylic beetle species
richness and abundance will increase in spruce–birch polycul-
tures, is heavily dependent on management decisions regarding
the amount of dead wood and old trees retained during the stand’s
rotation (Berg et al., 1994; Jonsell et al., 1998; Gärdenfors, 2005).
Similarly, rare lichen species favouring older birch trees are unlikely
to increase in species richness unless rotation periods are moved
well beyond the 50–60-year cycles normally employed in birch
production forests, or a significant number of trees are retained
after final felling (Kuusinen, 1996; Kuusinen and Siitonen, 1998;
Wannebo-Nilsen et al., 2010). Likewise, the diversity of understorey
vascular plants (Sieving and Willson, 1998), and the provision of
potentially suitable habitat for a diversity of ground or shrub associ-
ated bird species (Bibby et al., 1989; Berg, 1997; Poulsen, 2002), will
be lost if understorey vegetation is removed via grazing, herbicides,
or thinning (Donald et al., 1998; Hausner et al., 2002).
As such, there are multiple aspects of forest management (i.e.
rotation length, dead wood and green tree retention, thinning
regimes) which could be modified to enhance the biodiversity val-
ues of production forest stands (see Bengtsson et al., 2000; Hartley,
2002; Lindenmayer, 2002; Carnus et al., 2006; Fischer et al., 2006;
Lindenmayer et al., 2006; Stephens and Wagner, 2007; Brockerhoff
et al., 2008; Felton et al., 2010b). Importantly, the application of
such techniques to spruce–birch polycultures may yield larger bio-
diversity benefits than can be achieved in spruce monocultures.
This is because these polycultures begin from a better ecological
starting position, in terms of micro-climatic conditions, and struc-
tural and floristic complexity. In addition, our results indicate that
they should possess a more diverse species pool from which fur-
ther biodiversity goals could be achieved. Notably, the corollary
of this argument will also apply; if the extent of potential biodi-
 A. Felton et al. / Forest Ecology and Management 260 (2010) 939–947
versity gains is larger for spruce–birch polycultures, then so too
is the extent to which biodiversity benefits can be diluted due to
inappropriate or suboptimal forest management.
In summary, the replacement of spruce monocultures with
spruce–birch polycultures in the managed forest landscapes of
southern Sweden can be expected to result in an increase in biolog-
ical diversity for a variety of species representing a diverse range of
trophic levels, taxonomic affinities, and ecological niches. Although
such a transition does not represent a panacea for red-listed taxa,
and some taxa may be adversely affected (i.e. bryophytes), it does
provide a feasible means of improving the biodiversity value of
large numbers of production stands in southern Sweden. Perhaps
most importantly it also means that along with improving the
stands’ recreational value and decreasing climate change associ-
ated risks from wind and pest species damage, biodiversity can
be added to the list of likely benefits associated with facilitating
a transition to spruce–birch polycultures in this region.
This result is relevant to international discussions regard-
ing the potential benefits derivable from adopting risk-spreading
approaches to dealing with climate change impacts on managed
forests (Knoke et al., 2008; Burger, 2009). Diversification of tree
species within stands as part of a risk-spreading strategy appears to
facilitate the provision of multiple services from managed forests,
as is in line with increasing public expectations for forest lands
(Herbohn et al., 2000; Burger, 2009). Thus, at least in northern
Europe, the paradigms of multi-use forestry and risk-spreading
appear to be readily integrated. Further empirical-based assess-
ments of polycultures, both here and in other regions, are needed to
clarify the extent to which mixed-species stands can provide eco-
logical, social, and economic benefits to society, while at the same
time compensating forest managers and owners with reduced risk
from abiotic and biotic disturbances.
Acknowledgements
We thank Annika Felton for helpful comments on earlier iter-
ations of this manuscript. This research would not be possible
without financial and logistical support from the Southern Swedish
Forest Research Centre, SLU, Alnarp.
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