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Biodiversity in natural versus

managed forest in Denmark
Morten Christensen

Forest Ecology and Management

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 Fores;;;ology
Management
EISEVIER Forest Ecologyand Management85(1996) 47-51

Biodiversity in natural versus managed forest in Denmark

Mortdn Christensen a** , Jens Emborg b
a University of Aarhus, Department o$Ecology und Genetics, Ny Munkegode, Building 540, DK-8000 irhrhus C, Denmark
b The National Forest ad’ Nature Agency. Haraldsgade 53, DK-2100 Copenhagen 0, Denmark

Abstract

During recent millennia the biodiversity patterns of the Danish forests have changed. Many of the rare or threatened
species in Denmark are related to habitats characteristic of natural forest ecosystems, By comparing the structure and
dynamics of natural and managed forests the following key elements related to biodiversity were identified: woody debris,
forest heterogeneity, (successional) continuity and water. To incorporate these elements into modem forest management, a
change in attitude and practice is needed. Education and training will be an important part of this move towards
sustainability.

Keywords: Mosaic-cycle; Sustainability; Temperate forest dynamics

1. Introduction ash (Fraxinus excelsior L.) and elm (Ulmus glabra

It has recently been realized that protection of
biodiversity will be a major task for future forest
management. As in most of Europe, the Danish
landscape has been heavily influenced by humans
during the last 5-6 millennia. This has changed the
biodiversity patterns of Danish forests. In this article
we try to explain the main reasons for this change
and give some suggestions for forest management to
resolve the problem.
2. The historical background
The surviving semi-natural vegetation in most
parts of Denmark is cool-temperate deciduous forest
dominated by beech (Fugus sylvarica L.), oak
(Quercus robur L., Quercus petraea L. ex Liebl.),
* Corresponding author.
Hudson). The last glaciation of Denmark ended about
13000 years ago, at which time most of the country
was covered by a glacial ice-sheet. After the glacia-
tion, a large-scalesuccessiontook place, new species
successively arrived and some disappeared as the
climate, soil and vegetation changed.
During the first 7000 years after the glaciation,
the country was inhabited by a limited human popu-
lation of fishers, hunters and food-gatherers. From
about 6000 years ago, the land use pattern gradually
changed as the population increased and slash-and-
bum agricultural methods were adopted. The whole
landscape changed as the forests were turned into
agricultural land, a development which continued
throughout the succeedingmillennia.
The forest area gradually decreasedto a minimum
level in 1805 AD, representing less than 3% of the
total area of Denmark.
Since 1805 the forest area has increased to its
present level of about 12%. Most of the area consists

037% I 127/96/$ IS.00 Copyright 0 1996 Elsevier Science B.V. All rights reserved.
PII SO378- 1 127(96)03749-8
48 M. Chrisrensm. J. Emborg/ Forest Ecology and Manugemrnt 85 (1996) 47-51

of plantations, i.e. monocultures of introduced Table I

conifers. The area of forest genetically originating Status of terrestrial species in Denmark (The National Forest and
Nature Agency. 1990)
from the pristine forests is estimated to be about 1%
Total number Number of Red data list
of the total area of Denmark (The National Forest
of species species on spectes primartly
and Nature Agency, 1994). Today, most of these in Denmark the red data restricted to
genetically authentic forests are intensively managed. (approx.) list of 1990 forest habitats
^_____-
In 1993, only about 500 ha were protected as non-in- Macrofungi a 3ooo 903 (30%) 758 184%)
tervention forest. Lichens 900 634 (70%) 218 (34%)
Vascular plants 1200 261 (22%) 63 (24%)
Insects b 3684 1110(30%) 5651519;)
3. Red data book of threatened species in Den- Birds 185 97 (520/r,) 22f23qt
Mammals 50 19 W3%) ! 7 ~68’:: I
mark
Only Basidiomycetes and Ascomycetes with fruit-bodies larger
In 1990, a red data book for Denmark was pub- than 1 mm are included.
lished, considering the rare or threatened species of Only beetles (Coleoptera), butterflies (RhophaloceraL and bumets
(Zygaeidae) and included.
fungi, plants and animals (The National Forest and
Nature Agency, 1991). Out of a total of 9360 species
considered in the list, 3 176 were classified as rare or developmental phases (Fig. 21. With a typical patch-
threatened (Table 1). Of these, 353 had not been size of 500-1500 m2, spatial heterogeneity and
observed during the last 30 years and were thus beta-diversity are high. The natural temperate forest
classified as extinct. About half (1642) of the rare or ecosystem produces a lot of habitat gradients within
threatened species have their natural habitat in the relatively small areas, resulting in a high beta-diver-
forests, indicating the present problems of maintain- sity. Gamma-diversity is also high in natural forests.
ing biodiversity in the managed forests. In Suserup Forest (20 ha), a non-intervention ancient
forest in eastern Denmark, a total number of 601
species of fungi have been recorded between 1900
4. The biodiversity of natural temperate decidu-
and 1995, of which 74 were on the red dam list
ous forest
(Heilmann-Clausen, unpublished data). In Kaas For-
The dynamics of natural temperate deciduous
forests can be described by the mosaic-cycle concept
(Remmert, 1991). The forest cycle describes a con-
tinuous shift between a sequential series of develop-
mental phases (Fig. 1). Each of the phases provides
different life conditions for the organisms in the
forest. At the ‘innovation’ phase, many herbs and
shrubs have optimal conditions. The ‘degradation’
phase provides habitats for saprophytic fungi, wood-
living insects and hole-nesting birds, while the long-
lasting ‘biostatic’ phases provide the continuity and
stability needed for the establishment of tree-depen-
dent mosses, lichens and mycorrhiza fungi. This
explains the extremely high alpha-diversity (Whit-
taker, 1972) found in some patches or phases of the
mosaic-cycle.
Over time, every patch of forest goes through the
phases of the forest cycle, but unsynchronously from Fig. I. The forest cycle of natural temperate deciduous forest
patch to patch, thus creating a shifting mosaic of the (after Emborg, 1996).
 M. Christensen, J. Emborg/ Forest Ecology and Management 85 (1996) 47-51 49

innovation
aggradation
early biostatic
late biostatic
m degradation
Fig. 2. The shifting mosaic of Suserup Forest (10 ha) in 1993, a non-intervention ancient forest in eastern Denmark (after Emborg, I 996)

est (100 ha), an ancient oak-dominated non-interven-

tion forest in western Denmark, a total of 111 species
of epiphytic lichens have been found, of which 31
were red data species (Sochting and Christensen,
harvesting
1989).
early biostatic

I:
5. Tbe biodiversity of managed temperate decidu- ::
ous forest ::
::
The dynamics of the managed forests can be ::
described as a sequential shift between the early or
young phases of the forest cycle, equivalent to about
m!l!!u
aggJndstion
t :I
half of the full forest cycle (Fig. 3). Typically man-
aged beech stands are harvested when they are about innovation

120 years old, i.e. in the ‘early biostatic’ phase. The Fig. 3. The forest cycle of managed temperate deciduous forest.
consequence of this is an almost complete lack of
trees in the ‘late biostatic’ and ‘degradation’ phases
in managed forests. This affects the diversity of forest management. The light open conditions at the
oFganisms restricted to old trees and woody debris forest floor after thinning or clearcutting may even
(e.g. insects, fungi and hole-nesting birds). The di- be beneficial for these organisms.
versity of organisms primarily related to the ‘innova- The structural differences between the natural and
tion’ phase (e.g. vascular plants) is less affected by the managed deciduous forests are reflected in both
50 h4. Christensen, J. Emhorg/Forrst
the alpha- and the beta-diversity of the two forest
types. A typical managed forest in Denmark is com-
posed of a number of even-aged monospecific stands,
each typically within a size range of l-4 ha (Fig. 4).
The spatial heterogeneity is low compared to the
natural forest mosaic, and accordingly the beta-diver-
sity is relatively low. Many species are specialized to
live in the border-zone between two patches (e.g.
birds, insects, or mammals hiding in a closed dark
patch and feeding in an open patch). The large
patch-size in a managed forest reduces the frequency
of such border zones. Further, the long distance
between different patches of the same phase causes
lower alpha-diversity for organisms with a limited
dispersal capability, thus increasing the risk of ex-
tinction (see McArthur and Wilson, 1967).
During the last 150 years up to some 80% of the
forest wetland areas in Denmark have been drained
(Petersen, 1988). This development has further re-
duced the heterogeneity of the forest mosaic and
with this the beta-diversity of the forests. Accord-
ingly, many species related to the wet areas, or to the
gradient from moist to dry. are threatened in the
managed forests. In this way draining has con-
tributed to the reduced diversity of the forests.
u innovation
aggradation
early biostatic
coniferous plantation
Fig. 4. Typical Danish managed forest (IO ha from Ringelmose
Ecology und h4anagement 85 (1996147-51
To summarize, the result of human impact is a
loss of both alpha- and beta-diversity, and hence a
lower gamma-diversity of the Danish forests.
6. Sustainable forestry in Denmark
In short, we conclude that the main biodiversity
problems in the intensively managed forests of Den-
mark are related to lack of woody debris, lack of
floristic and structural heterogeneity, lack of (forest
successional) continuity and lack of permanent water
in the forests. Hence, part of the solution of the
biodiversity problem of the Danish forests is:
1. To set aside non-intervention forest reserves where
natural forest dynamics (i.e. the full forest cycle)
can take place.
2. To improve forest management systems such that
they allow for the continuation of natural forest
dynamics, including forest heterogeneity, the re-
tention of woody material on the forest fhr, and
protection of water bodies and swamps.
To mention one example of management im-
provements: dead wood should be left behind after
cutting or felling operations (Fig. 51. Such efforts do
Forest, Denmark; after M. Christensen, unpublished data, 1%).
 M. Christensen, J. Emborg / Forest Ecology und Munagement 85 (1996) 47-51 51

already to act in order to improve our caretaking of

biodiversity in the Danish forests.
Education and training of forest staff at all levels
(from forest workers to technicians and managers) is
needed to change attitudes and practices after many
years of clearcut-based management. Educational ini-
tiatives should be among the first steps to be taken in
the long-term process towards sustainability.

References
Emborg, J., 1996. The structure dynamics and light conditions of
Suserup Forest, a semi-natural temperate forest in Denmark.
Ph.D. Thesis, The Royal Veterinary and Agricultural Univer-
innovation
sity, Copenhagen.
Fig. 5. The forest cycle of a sustainably managed forest. The basic McArthur, R.H. and Wilson, E.O., 1967. The Theory of Island
principle is to integrate key elements of the full forest cycle in the Biogeography. Princeton University Press, Princeton, NJ.
management system. Petersen, F.R., 1988. Aspects of the Decline in Extent of Mires in
Central North Zealand State Forests During the 19th and 20th
Century. Prize Essay, University of Copenhagen, 99 pp.
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