Larsen Et Al. Ecology of Tree Species and Selection

Ecology of tree species and species selection
J. Bo Larsen, Karsten Raulund Rasmussen and Ingeborg Callesen
Introduction
The key principle of nature based silviculture is to incorporate ecosystem processes such as
succession, natural regeneration, and self-differentiation in management. A prerequisite is that the
species used can regenerate naturally i.e. are adapted to the local site and soil conditions and they
can grow together in harmony in the forest community.
The Forest development types include various types of tree-species compositions based on the
knowledge of soil and climate demands for each individual species, as well as the sociological part
it plays in a forest stand.
Below you will find an ecological ‘profile’ for each of the 19 most important North-European
species. Each species being characterized by:
1. Native range and role in the forest community
The natural distribution of the species is examined and displayed on a map. Further, its successional
role in the forest community and the associated species are described.
2. Climatic requirements:
The climatic requirements of the species are examined especially regarding frost, drought, wind and
salt. Furthermore, the potential adaptability to the expected climatic changes (higher temperatures
and winter precipitation, lower summer precipitation, and increased storm frequency) is discussed.
3. Site requirements:
Site requirements (nutrient and water supply, root development, etc) and soil conditions of the
species are described. A site classification diagram states the site requirements of the species
according to nutrients and water availability. The nutrient conditions are classified from 1 (very
poor), to 6 (very rich). The water conditions are divided in soils with free drainage and soils with
limited drainage.
Soils with free drainage never have reduced oxygen pressure to such an extent that it affects even
the most sensitive species. They are classified according to water supply degree, root zone capacity
and summer precipitation at a scale from 1 (very dry) to 6 (very fresh/moist).
1
Soils with limited drainage are classified from 7 to 9 depending on depth to the poorest layers of
drained stratum. Limited drainage covers pseudogley as well as gley. Soils with extremely bad
drainage (type 9) might have an increased humus layer or simply developed a peat layer. Peaty
soils are explicitly described, if the species are often used at such sites.
In the ecological diagrams the water and nutrition conditions are grouped as follows:
• Optimal (green) where the species is healthy, steady growing and productive corresponding to
the best site conditions.
• Acceptable (grey) where the species normally is healthy but characterized by slower growth
with a tendency to stagnation and sensitive to storm corresponding to limited but acceptable site
conditions.
• Unsuitable (red) where the species is not recommendable neither according to growth, health
nor stability.
Beech Soils with free drainage Limited

4 moderate fresh
drainage
3 moderate dry
7 80 - 160 cm
8 40 - 80 cm
6 very fresh
1 very dry
9 < 40 cm
5 fresh
2 dry
1 very poor
Nutrition
2 poor
3 moderate poor
4 moderate rich
5 rich
6 very rich
Ecological diagram for beech (Fagus silvatica)
4. Matching species to sites and to forest development types:
Here the species is described according to its role in nature-based forest management and to its
possibilities according to different forest development types.
A more detailed description of each of the species according to range, race, planting in Denmark,
seed supply, variation of provenance and recommendation of seed sources (provenances) is found in
(Larsen 1997): Tree species- and provenance selection in a sustainable forestry, Dansk Skovbrugs
Tidsskrift, 82, 1 - 252, 1997.
5. Provenances recommended:
The selection of the right species is a prerequisite for robust and functional forests. Almost as
important is the choice of genetic material within the species – the selection of provenance. A list of
recommended provenances for Danish growing conditions based on Larsen (1997) is displayed.
2
Tree species
Alder (Alnus glutinosa (L.) Gaertn.) 4
Ash (Fraxinus excelsior L.) 6
Beech (Fagus silvatica L.) 8
Birch (Betula pendula Roth. and Betula pubescens Ehrh.) 10
Cherry (Prunus avium L.) 12
Hornbeam (Carpinus betulus L.) 14
Lime (Tilia cordata Mill.) 16
Mountain ash (Sorbus aucuparia L.) 18
Norway maple (Acer platanoides L.) 20
Pedunculate oak (Quercus robur L.) 22
Sessile oak (Quercus petraea Liebl.) 24
Sycamore maple (Acer pseudoplatanus L.) 26
Douglas fir (Pseudotsuga menziesii (Mirb.) Franco) 28
Grand Fir (Abies grandis Lindl.) 30
Larch (Larix dicidua Mill., Larix kaempferi Carr. and Larix x eurolepis) 32
Norway Spruce (Picea abies (L.) Karst.) 34
Scotch pine (Pinus silvestris L.) 36
Silver fir (Abies alba Mill.) 38
Sitka spruce (Picea sitchensis (Bong.) Carr.) 40
3
Alder (Alnus glutinosa (L.) Gaertn.)
Native range and role in the forest community:
Alder is widespread all over Europe and in some parts of Asia Minor and North Africa (the Atlas
Mountains). In Scandinavia it is growing up to 66° N. It is a typical lowland species connected to
river valleys and swamp forests. In southern parts of Europe it is a mountain tree and found at 1800
m in the Alps.
Alder is light demanding especially in the juvenile phase. It grows fast in the youth and seldom
reaches more than 100-120 years of age. Due to its high demand for light it cannot compete with
longer living, more shade tolerant species. However it is superior on poorly drained sites. On
permanent water logged sites alder is growing in even aged stands. On less ground water influenced
sites it grows in mixture with other broadleaved species. On fertile wet soils it is found together
with ash and oak and on poorer sites in mixed stands with birch.
Climatic requirements:
Alder is rather frost resistant and is seldom damaged by frost. It tolerates permanent water logging
and with roots in direct contact to ground water it can reach high age. Alder is expected to be robust
to climate change. The potential area will probably diminish if climate changes are followed by less
rainfall.
4
Site requirements:
Alder prefers meso- to eutrophic, humus rich soils. It is growing along stream sides and permanent
wetlands. You will not find it on raised bogs. But it can grow on wet soils and lowland moors with
stabile water conditions. The root system is densely branched and might be able to penetrate
compact soils. Compared to ash nutrient requirements of alder are low and its capacity to fix
nitrogen can improve nutrient poor sites. Dry and very poor sites should be avoided. On such sites
alder can function as a nursing tree, but it often dies after 20-30 years. Due to its biodegradable
leaves and nitrogen assimilative root tubercles, alder has a soil improving capacity.
Red alder Soils with free drainage 4 moderate fresh Limited

drainage
3 moderate dry
7 80 - 160 cm
8 40 - 80 cm
6 very fresh
1 very dry
9 < 40 cm
5 fresh
2 dry
1 very poor
Nutrition
2 poor
3 moderate poor
4 moderate rich
5 rich
6 very rich
Matching species to sites:
Alder is especially bound to the groundwater level. Due to draining of forest wetlands during the
last centuries, natural sites of alder have more or less disappeared. If the natural hydrology of these
areas was re-established, alder could regain a considerable range. It is only represented in a few
forest development types. Most outstanding in forest development types: Ash with alder (31), where
it can be quite dominating on the very wet sites. Further, it can develop on smaller (restored) water
logged sites mixed with oak, ash, birch and Scots pine. Finally, alder is attached to the historical
forest development type Coppice forest (91) as well as to the Woodland meadow (93).
Provenances recommended:
Alder seems to build local adapted populations, and it is especially not advisable to move alder
from south to north. Foreign seed sources are not recommended.
1. Denmark: Selected seed stands

2. Denmark: Other local seed sources
5
Ash (Fraxinus excelsior L.)
Ash is naturally distributed in most of Europe. It is ranging from Ireland, England and Wales,
southern part of Norway, Sweden and Finland to northern part of Spain, Sicily and Greece. It grows
along the Black Sea, in the Caucasus and in the Elburz mountains in Iran. Up north ash trees are
growing in the lowlands, while in Central- and E Europe it can grow up to altitudes of 1600 m. In
Denmark ash is common almost anywhere. Larger stands are found along smaller riverbanks and in
beach forests in SE Jutland, Funen and Bornholm, locally also on Lolland, Falster, and South-
Zealand.
Ash is shade tolerant in the first development stages, but a prerequisite for further growth into the
canopy is development of permanent openings/gaps. Ecologically ash is a gap specialist. With an
ample, regular and easily wind-dispersed seed production, a steady regeneration is secured
everywhere there is a canopy gap. Like sycamore maple, it is therefore often a part of the Middle
European broad-leaved forest dynamics, characterized by small scale disturbances.
Correspondingly, it is often seen with sycamore maple in well drained beech dominated forests. It
associates as well with pedunculate oak, lime and hornbeam on nutrient rich, badly drained sites. In
very wet soils it is seen with alder and pedunculate oak.
6
Ash is rather sensitive to frost, but can nevertheless be established without shelter in not too frost
exposed sites. Good water supply is a must either from rainfall or better through ground water
contact. Expected climatic changes with not too severe summer drought can be profitable for ash,
since Denmark is positioned close to its northern limit.
Site requirements:
Ash has relatively large site amplitude. It is developing well in deep grounded, nutritious soils, but
it thrives also on poorer sites when its water requirement is fulfilled. It grows well on sites with
pseudogley and gley, in chalky soils, and in well drained and not too acid, peat soils. Due to
competition from the much more competitive beech, ash is often found far away from its optimum.
Dead leaves are easily decomposed and the species is a typical mull former.
Ash Soils with free drainage Limited

4 moderate fresh
drainage
3 moderate dry
7 80 - 160 cm
8 40 - 80 cm
6 very fresh
1 very dry
9 < 40 cm
5 fresh
2 dry
1 very poor
Nutrition
2 poor
3 moderate poor
4 moderate rich
5 rich
6 very rich
Due to a large ecological amplitude and good regeneration ability ash will be a good deal more
important in nature-based forest management, than it has ever been in classic forestry, where it has
often been deported to poorly drained sites. It plays an important role in forest development type
Ash and alder (31), and a co-dominant role in Beech with ash and sycamore (12) and Oak with ash
and hornbeam (21). It is also participating in Coppice (91) and Grazing forests (92).
Growth and especially form can to a great extend be influenced by provenance selection. Ash is
easy to breed, and there are several seed orchards, some of which have rather limited number of
clones.
1. Denmark: Selected seed stands and seed orchards
2. Germany: Selected stands (Gebiet 81101, 81102, 81103)
3. Holland: Selected stands
7
Beech (Fagus silvatica L.)
Beech is widely distributed in West and Central Europe’s temperate zone. Its northern range goes
through the southern parts of Norway and Central Sweden and its southern range as far south as
Sicily. In N Europe it is a low land tree, often mixed with ash, sycamore maple, oak, elm, cherry
etc. In the south and south-eastern parts of Europe it is growing in the mountains associating with
silver fir and Norway spruce and occasionally with larch. Beech is a shade tolerant species giving a
lot of shadow. It is a typical climax species attached to late succession stages and is often dominant
in natural forest sites. It regenerates best in already existing forests with shelter, shadow and high
humidity.
Beech is sensitive to severe winter frost. Like a typical climax species it bud-bursts early and is
often damaged by spring frost; at the most frost beaten places (afforestation and clear-cutting)
nursing trees or pre-cultures are therefore necessary. In the growing season the species is sensitive
to drought, which can lead to degradation of xylem shown as red rot and black spots (‘freckles’).
8
Site requirements:
Beech has relatively large soil amplitude. It develops well on deep soil with plenty of nutrients and
water. It tolerates also poor and dry soils, but this result in modest growth. Most problematic for
beech is poor drainage followed by periodically poor oxygen content in the soil. Circumstances like
that will not eliminate the species. Heavy cutting will make it more sensitive to drought and
stagnation, and subsequently crown degeneration and red rot. With pseudogley deeper than 80 cm
this is not necessarily a problem. Intermingling of more tolerant species such as ash, lime, oak,
hornbeam, silver fir, etc. might reduce these problems for beech. Peaty soil is inappropriate,
whereas chalky soil is suitable for beech. Compared to other species, dead leaves of beech are less
biodegradable and it is producing sour humus on poor soils. On optimal sites, however, it produces
an excellent humus layer (mull), especially in mixed stands with other broad-leaved tree species.
Beech Soils with free drainage Limited

4 moderate fresh
drainage
3 moderate dry
7 80 - 160 cm
8 40 - 80 cm
6 very fresh
1 very dry
9 < 40 cm
5 fresh
2 dry
1 very poor
Nutrition
2 poor
3 moderate poor
4 moderate rich
5 rich
6 very rich
Beech is the species most naturally distributed in Denmark. Consequently, it is dominating most
forest sites, except the poorest, most sandy, water logged and peaty ones. Similarly, beech is
playing a greater or lesser role in most forest development types. There are four beech dominated
types: Beech (11), Beech with ash and sycamore maple (12), Beech with Douglas fir and larch (13)
Beech with spruce (14). Furthermore, there are four other types where beech has an important role:
Oak with lime and beech (22), Spruce with beech and sycamore (51), Douglas fir with Norway
spruce and beech (61), and Fir and beech (71).
As a typical climax species beech is not very genetically differentiated, and beech from many
different parts of its range can grow in Denmark with a good result. From a gene conservation
viewpoint Danish (local) selected seed sources should be preferred.
2. Switzerland: Zurich-area (Sihlwald, Adlisberg)
3. Germany: Selected seed stands (Gebiet 81001, 02, 03, 07, 09)
4. Slovakia: Carpathian Mts. (Kakasovce eller Sigord)
5. Holland and Belgium: Selected seed stands
9
Birch (Betula pendula Roth. and Betula pubescens Ehrh.)
Silver birch (Betula Pendula) and downy birch (Betula pubescens) are both common almost
anywhere in Europe from N Scandinavia to S Italy, from Ireland far into Russia. Downy birch is not
as common in the south as silver birch. Up north, however, it is native in e.g. Greenland and
Iceland. Accordingly, you find it higher in the mountains than silver birch. In Denmark silver birch
is very common in most regions. Downy birch, however, grows mainly on peat and in swamp areas
and is very common in North- and Central Jutland as well as in North Zealand. In classic forestry
birch has been regarded as weed, something to control. Therefore, birch is rarely found in any
managed forest.
Birch is our most typical pioneer species and extremely light demanding. Birch depends on larger
disturbances e.g. windfalls, forest fire, etc. to be able to colonize effectively with its abundant,
easily wind dispersed seeds. It is playing an important role in the early successions after minor or
larger disturbances. Only on peat soil it is able to establish more or less stabile forests with other
pioneer species e.g. Scotch pine, mountain ash´, etc.
10
As a pioneer species birch is extremely climate tolerant and is rarely damaged by frost. Likewise, it
is very drought resistant. Silver birch is, however, more drought resistant than downy birch. With
sufficient water supply, evapotranspiration of birch stands is similarly very high. Both birch species
are considered very robust to climate changes.
Site requirements:
Birch has extremely wide soil amplitude. They can grow in every kind of soil from the most poor
and dry to the most nutrient rich and wet soils. The two species differ, however, from each other
regarding soil requirements. Silver birch prefers dry, sandy soils, while white birch tolerates
waterlogged peat sites. Birch is generally tolerant to the most harsh site conditions, including
pseudogley and groundwater stress.
Birch Soils with free drainage Limited

4 moderate fresh
drainage
3 moderate dry
7 80 - 160 cm
8 40 - 80 cm
6 very fresh
1 very dry
9 < 40 cm
5 fresh
2 dry
1 very poor
Nutrition
2 poor
3 moderate poor
4 moderate rich
5 rich
6 very rich
Birch is the most important pioneer species in the forest. Generally birch is important in the forest
as a ‘sore healer’ after windfalls or other disasters due to its ability to re-establish the forest climate
as an early successional species. Therefore, it is very important generally to leave single birch trees
in the forest, especially in the spruce dominated ones, as a safeguard against disasters. In nature-
based forest management it is associated with disasters and marginal sites (sand and peat). Hence, it
is playing a minor role in many forest development types as an “insurance” in case of major
disasters but frequently present in forest development types on marginal sites such as Birch with
Scotch pine and spruce (41) and Scotch pine, birch and spruce (81). As a pioneer species birch
plays a certain role in some Coppice forests (91) and in Grazing forests (92).
The following recommendations are for both species of birch.

2. Sweden: Selected stands and seed orchards south of the 57th latitude
3. Germany: Selected stands and seed orchards in northern Germany
11
Wild Cherry (Prunus avium L.)
Wild cherry is naturally distributed almost all over Europe. It is found in N England and in Mid-
Sweden. Its southern range goes through N Spain, Italy and S Balkan. The species is also growing
around the Black Sea, in Caucasus, Elburz Mountains and in N Africa. In the Black Forest you find
it up to 1000 m, in the Alps up to 1700 m and in the Caucasus up to 2000 m.
In Denmark wild cherry is a common species in the forests of Bornholm. In other parts of the
country it is mainly found in forest edges and on overgrown pasture land. You seldom see it in the
westernmost parts of Jutland.
Wild cherry is a typical pioneer light-demanding tree and only found as an intermixture, solitary or
in groups. In beech forests it is often found together with ash and sycamore or mixed with oak and
ash. In the Alps, it is primarily found in natural groups mixed with conifers, Scotch pine and larch.
It is a typical tree of the landscape and of the forest edges. Here its ecologic (insect pollinator) and
aesthetic (flowers and berries) advantages are displayed in full.
12
Generally Wild cherry prefers a warmer climate than we have at present in Denmark, which is close
to its northern range. It is a typical pioneer species with early succession, seldom frost-bitten and
drought resistant to some extent. Future climate changes will be a direct advantage for wild cherry.
Site requirements:
Wild cherry can grow on many different sites and tolerates different nutrients and water conditions.
Soils with limited or bad drainage are not acceptable (especially water supply type 9 and the wet
ones of type 8). The optimal site conditions are moderate to nutrient rich well drained soils.
However, it can also grow on dry, but not too poor soils. In that cases productivity is limited. The
species regenerates easily and its seeds are spread over far distances by birds. To ensure good
growth (for timber production) well drained soils with good water supply are necessary.
Cherry Soils with free drainage Limited

4 moderate fresh
drainage
3 moderate dry
7 80 - 160 cm
8 40 - 80 cm
6 very fresh
1 very dry
9 < 40 cm
5 fresh
2 dry
1 very poor
Nutrition
2 poor
3 moderate poor
4 moderate rich
5 rich
6 very rich
In Denmark, experiences with growing bird cherries are limited and the species is seldom found in
our forests. In future, it is entitled a more important position, due to its ecologic and aesthetic
advantages, but also because of its excellent wood quality. It seems, however, to be unfit for
growing in even aged stands. In the forest development types, like in nature, it has a position as an
intermixture. Likewise it appears in Beech (11), Beech with ash and sycamore (12), Oak with ash
and hornbeam (21) on not too clayey soils, and in Oak with lime and beech (22). Apart from that,
wild cherry is generally an important species in forest edges and in the landscape, but also in
Grazing forests (92) and on Woodland meadows (93).
When growing cherry for wood production the selection of provenance is of outmost importance. In
such cases only material from seed orchards or selected seed stands can be recommended.
1. Germany: Seed orchards in northern Germany
3. Germany: Seed stands in northern Germany
13
Hornbeam (Carpinus betulus L.)
Hornbeam is found throughout Central Europe: From S England to Denmark, from S Sweden to S
France, Italy and Greece. The eastern range goes along the Russian border. It is, however, also
found around the Black Sea, in Caucasus and in the Elburz mountains in Iran. In the northern
regions hornbeam is growing in the lowlands. In Central Europe, however, it is growing at 1000 m
and in the Caucasus even at 2000 m. It can be about 150-250 years old, but it seldom grows higher
than 25 m even at the best sites.
Even though hornbeam is shade tolerant, it is seldom dominating in its forest communities due to its
limited height growth. Ecologically it fills a niche as an intermixture in form of a sub-canopy in oak
forests and in other light-demanding forests communities. It is never found in age-class forests, but
often in mixtures with other broad leaved species. It easily regenerates by sprouts and is therefore
common in the Central European coppice- and coppice-with-standard forest systems.
14
Hornbeam is adapted to a sub-oceanic/sub continental climate with an optimum in the south Baltic
region (Lithuania and Poland). It prefers warm summers but tolerates cold winters. Compared with
beech it is less sensitive to late spring frost. In not too harsh places it can be cultivated on bare land
without surrounding forest or nurse crop. Near its northern range (for example Denmark) it prefers
warmer sites. The south-easternmost parts of Denmark are therefore the best place to grow
hornbeam. Expected warmer climate will be an advantage for hornbeam.
Site requirements:
Hornbeam is relatively tolerant to nutrient and water supply, but for developing large trees
sufficient nutrient supply is a prerequisite. It can grow on dry sites, as well as in very badly drained
soils. It also tolerates stiff clay with pseudogley, but does not grow well on peaty soil. Dead leaves
of hornbeam are easily decomposed and the species is a typical mull former.
Hornbeam Soils with free drainage Limited

4 moderate fresh
drainage
3 moderate dry
7 80 - 160 cm
8 40 - 80 cm
6 very fresh
1 very dry
9 < 40 cm
5 fresh
2 dry
1 very poor
Nutrition
2 poor
3 moderate poor
4 moderate rich
5 rich
6 very rich
Due to limited growth in height and good shade tolerance, hornbeam is often used as undergrowth
in oak stands fighting water sprouts. As such it is ideal for creating sup-canopy structures and
ground cover in light demanding forest development types and is especially important concerning
Oak with ash and hornbeam (21). Also in other forest development types on better soils hornbeam
has a certain position e.g. in Beech with ash and sycamore (12), Oak with lime and beech (22), and
Ash with alder (31). In the Coppice forest (91) you might also find hornbeam, especially on fertile
soils.
Experiences with provenances of hornbeam are absent, therefore, good local stands are
recommended for seed supply.
1. Denmark: Local seed stands
15
Lime (Tilia cordata Mill.)
Small-leaved lime is widespread almost everywhere in Europe. Its natural distribution from north to
south reaches from S Norway, Central Sweden and S Finland to S Italy and the Balkan mountains.
From east to west you find it in Ireland, England and from N Spain far into Russia. You also find it
in Caucasus and in the Elburz Mountains in Iran. Up north it is found in the lowland, in the Alps in
the mountains up to 1500 m.
About 6000 to 8000 years ago lime populated Denmark and was the dominant tree species when
temperatures were 2 - 3 degrees warmer than today. As the temperature dropped in the iron age and
man began forest clearings, beech gradually replaced lime. So today only few stands remain in
southern parts of Denmark. The best known places are Draved in southern Jutland and Vindeholme
on Lolland.
Lime is shade tolerant, but not as much as beech. On many forest sites it grows naturally in smaller
groups, but never in coherent stands. In nutrient rich forest stands you find it with oak, hornbeam,
ash, elm, poplar, etc. and on nutrient poor soils with beech, oak, birch, ash, rowan, etc. It has a
capability for vegetative regeneration and has therefore been an important species in the old days’
coppice- and coppice-with-standards forestry systems.
16
Lime has its optimum in a sub-continental climate with warm summers and cold winters. Our
Atlantic climate with cool summers is not ideal for lime; especially not in the seed ripening season.
Therefore, it is very unusual for lime to regenerate generatively in Denmark. In spite of rather early
bud-burst it tolerates late spring frost. It even tolerates winter frost and drought. Perhaps due to a
well-developed root system it grows well even in soils with alternating water table. Lime will
prosper by the expected higher temperatures.
Site requirements:
Lime has huge soil amplitude and is able to develop in moderate poor as well as in very rich soils. It
develops well in water- and nutrient rich soils. It tolerates heavy clay and poor drained soils with
pseudogley and gley in depth of 30-50 cm. Peaty soils, however, are not suitable.
Lime Soils with free drainage Limited

4 moderate fresh
drainage
3 moderate dry
7 80 - 160 cm
8 40 - 80 cm
6 very fresh
1 very dry
9 < 40 cm
5 fresh
2 dry
1 very poor
Nutrition
2 poor
3 moderate poor
4 moderate rich
5 rich
6 very rich
With its wide ecologic amplitude and tolerance to water logging lime is suitable in a number of
forest development types. Future climate changes will no doubt be vital for lime and improve its
regenerative ability. In the following forest development types lime is playing a more or less
important role: Oak with lime and beech (22), Beech with ash and sycamore (12), Beech and spruce
(14), Oak with ash and hornbeam (21), and Ash and alder (31). In addition lime is growing in the
historical forest development types Coppice forest (91) and Grazing forest (92).
Similar to beech lime is not very differentiated genetically, and lime from many different parts of its
range can be grown in Denmark with a good result. From a gene conservation viewpoint Danish
(locally) selected seed sources should be preferred.
2. Germany: Selected stands and seed orchards (Gebiet 82301, 02 og 03)
17
Mountain ash (Sorbus aucuparia L.)
Mountain ash is common most places in Europe – from the North Cape north of the polar circle to
Sicily and S Balkans. From Iceland, Ireland and N Spain to the Ural Mountains and Caucasus. And
even in N Africa in the mountains of Morocco. Similarly, it grows in the lowlands and up to the
Alpine timber line in the Central- and E European mountains at 2000 m. Mountain ash is found
most places in Denmark, most often in poor sites, with less competition from other more vigorously
growing species.
Mountain ash is a typical pioneer species with rapid, soon subsiding juvenile growth. It grows up to
20 m and will not reach more than 100 years of age. Primarily, it is attached to the early succession
stages right after larger or minor disasters, very often mixed with birch, alder, willow, oak, etc. In
the sub alpine forests it is growing in mixed stands with cembra- and mountain pine. In Denmark,
mountain ash is typical in forest edges often positioned in light open spruce stands. It often appears
after windfall, if not eaten by game. It plays, therefore, a very important role as gap filler. Due to its
tolerance and resistance to climatic extremes and not least due to its decorative autumn coloured
leaves and the rowanberries, mountain ash has become an important rural and urban tree species.
18
Mountain ash is extremely climate tolerant and in line with its pioneer qualities, it is frost hardy and
drought resistant. Similarly, it is growing well in a temperate and damp climate (e.g. on northern
slopes). The expected climatic changes will have little effect on mountain ash.
Site requirements:
The site requirements of mountain ash are low and its ecological amplitude is huge. It can grow
everywhere. On nutrient poor and dry sites, but also well on nutrient rich and ground water logged
sites. On the most nutrient poor sites, however, a certain amount of humus and high atmospheric
humidity is necessary. It grows well on peat, but raised bog is not a good place to grow mountain
ash. Dead leaves of mountain ash are easily degradable.
Mountain ash Soils with free drainage Limited

4 moderate fresh
drainage
3 moderate dry
7 80 - 160 cm
8 40 - 80 cm
6 very fresh
1 very dry
9 < 40 cm
5 fresh
2 dry
1 very poor
Nutrition
2 poor
3 moderate poor
4 moderate rich
5 rich
6 very rich

In classic forestry mountain ash has for a long time been regarded as weed. Consequently, the
species has been suppressed. Birds are interested in the berries and in return they help spreading the
species. In nature-based forest management the species has a number of qualities, which is making
it interesting. Converting moor land plantations for example, mountain ash is often appearing
spontaneously as an excellent intermixture. Together with birch it is an important element in
secondary successions after windfalls. In future forest management it is not going to play an
important role. In most forest development types on lighter soils, however, it will be playing a
minor role as an intermixture species.
Experiences with provenances of mountain ash are absent, therefore, good local stands are
19
Norway maple (Acer platanoides L.)
Norway maple is native to the sub oceanic and sub continental parts of Central and E Europe. It is
most common in lowlands, along river valleys, etc. It is also occurring in SE Scandinavia and in
Denmark, in S Finland, the Baltic States, and Poland and far into the Urals in Russia. Its natural
western border goes through Germany and E France. Apart from SE Europe, Norway maple is
growing in W Turkey and the Caucasus. It has often been planted outside its natural range and is
generally advancing up in N Scandinavia and W England, Ireland, and also into the cities. The
species was brought into N America, where it has been listed as an invasive species in some areas
because of an early and fruitful fructification with an extreme ability to regenerate.
Norway maple is like Sycamore maple intermediary light demanding. It grows up to 20 m when
mature and will therefore never be a very frequent or a dominant element in the forest. It is shadow
tolerant as juvenile, especially on rich soils. Later it becomes more light demanding. It is frequently
flowering and regenerating everywhere and is therefore ready immediately after there has been
made a gap in the crown canopy. It appears often in small groups or as a solitary tree in forests with
ash, lime and hornbeam, being part of a sub-canopy stratum. Earlier is has been used in coppice or
in coppice-with-standards systems because of its willingness to produce sprouts.
20
Norway maple prefers warm summers, but tolerates also cold winters and even some frost and
drought during autumn and spring. This general sturdiness, combined with a huge variety in forms
and colours, has made it popular in cities and parks. This sub continental climate profile and
drought adaptability indicates that it will prosper by future climate changes.
Site requirements:
Norway maple is like most other species growing well on rich, deep grounded soils with good water
supply. It is, however, also growing well on poor and dry soils. All considered Norway maple is
more tolerant than sycamore. Like sycamore, however, it is sensitive to stagnant water in the root
zone. Its dead leaves are easily decomposed.
Norway maple Soils with free drainage Limited

4 moderate fresh
drainage
3 moderate dry
7 80 - 160 cm
8 40 - 80 cm
6 very fresh
1 very dry
9 < 40 cm
5 fresh
2 dry
1 very poor
Nutrition
2 poor
3 moderate poor
4 moderate rich
5 rich
6 very rich
Norway maple has never been included in classic forestry and consequently always been reduced to
edges of wood and to open landscapes. In nature based forestry, it is will never be a dominating
species in any forest development site. It plays a certain part, however, as an intermixture species in
forest development types such as Beech with Douglas and larch (13), Oak with ash and hornbeam
(21), Ash and alder (31), Spruce with beech (51) and in the historical forest development types such
as Coppice forest (91), Grazing forest (92) and Woodland meadow (93).
Experiences with provenances of Norway maple are absent, therefore, good local stands are
2. Germany: Seed stands in northern Germany
21
Pedunculate oak (Quercus robur L.)
Pedunculate oak is one of the most common species in European lowlands. It is widely distributed
from S Scandinavia to Italy and the Balkan mountains, from Portugal and N Spain far into Russia.
Larger natural populations of this long living and light demanding tree species is found where the
competition of beech is weakened: on nutritious poor, dry sites and even more on rich humid,
groundwater influenced sites. It also competes well on typical pseudogley and along Central
European rivers, where it tolerates periodically flooding. Oak is a light-demanding tree species with
typical pioneer qualities. On nutrient rich sites it is growing in forest stands with hornbeam, ash,
elm, lime, poplar, etc. On nutrient poor soils it is mixed with sessile oak, birch, aspen, rowan, lime.
As a pioneer species pedunculate oak is tolerant to frost and drought (especially to spring frost due
to late leaf burst). Very hard winter frost, especially late winter frost can, however, damage the
cambium. Influenced by a variety of fungi, it can periodically lead to the degeneration phenomenon
‘Sudden Oak Death’. Likewise, early autumn frost can damage not yet hardened summer shoots. It
is drought tolerant. Due to better regeneration abilities pedunculate oak is considered to be
somewhat more tolerant to climatic and biotic damages than sessile oak. It has a pronounced
22
tendency for water sprouts, which is negative seen from a wood quality point of view. As a warmth
loving and drought tolerant species oak will prosper by the expected climatic changes.
Site requirements:
Pedunculate oak has huge soil amplitude. Best development is on nutrient rich, deep grounded sites
with ample water supply. It can also grow well on poor, sandy and dry sites and on clayey and poor
drained soils with gley and pseudogley. Even on very ground water influenced sites (type 9) it is
developing acceptably, conditioned there is no peat tendency. It is considered to be better suited for
fertile wet soils than Sessile oak, which is more suited to dry, poor soils. Pedunculate oak should
not be planted on peaty soils. The root system of oak is often considered to be very strong with a
tendency for deep growth.
Pedunculate oak Soils with free drainage Limited

4 moderate fresh
drainage
3 moderate dry
7 80 - 160 cm
8 40 - 80 cm
6 very fresh
1 very dry
9 < 40 cm
5 fresh
2 dry
1 very poor
Nutrition
2 poor
3 moderate poor
4 moderate rich
5 rich
6 very rich
Oak is a dominating tree species in three different forest development types: Oak with ash and
hornbeam (21), Oak with lime and beech (22) and Oak with Scotch pine and larch (23). Pedunculate
oak is more attached to the nutrient rich types. Pedunculate oak is also seen mixed in most forest
development types, especially in the light-demanding Ash and alder (31), Birch with Scots pine and
spruce (41), plus the historical forest development sites such as Coppice forest (91), Grazing forest
(92) and Pasture woodland (93). In the dune forests Pedunculate oak is more important than sessile
oak in forest development types with Sitka spruce with pine and broadleaves (52) and Mountain
pine (82).
The selection of pedunculate oak provenances depends to a great extend on the site and the purpose.
Protected fertile sites in eastern Denmark (mainly for wood production)
1. Denmark: Selected seed stands of Dutch origin
2. Holland: Special selected avenues
4. Holland: Selected stands (gebied-stands)
Frost and wind exposed sites in western Denmark (mainly for landscape and ecology)
1. Selected local seed stands
23
Sessile oak (Quercus petraea Liebl.)
Sessile oak is like pedunculate oak widespread in W Europe except from the eastern parts. Unlike
Pedunculate oak it is not doing well in continental climate. North/south distribution goes from S
Scandinavia to S Italy and the Balkan mountains; west/east from Portugal, N Spain and Ireland to
the Russian border. In most of its distribution sessile oak is growing together with pedunculate oak
and in southern direction even with pubescent oak. It is most common on lighter soils, but unlike
pedunculate oak sensitive to bad draining. Totally it has less ecologic amplitude than pedunculate
oak. In Denmark, sessile oak is especially dominant on lighter soils in Mid- and West Jutland (often
in oak scrubs) and on Djursland. Sessile oak is also common along the south coast of Zealand and
on Bornholm.
Sessile oak is a light-demanding tree species with typical pioneer qualities. On fertile soils it is
developing with pedunculate oak, hornbeam, ash, beech, lime, etc. On poor soils it is mixed with
beech, birch, ash, rowan, lime, etc.
Sessile oak prefers warm summers and due to its pioneer qualities it is frost tolerant, especially
spring frost due to late leaf burst. Very hard winter frost, particularly late winter frost, can damage
24
the cambium. With assistance of a variety of fungi, this can periodically lead to the degeneration
phenomenon ‘Sudden Oak Death’. In the juvenile phase it is fast growing and will develop a deep
root growth and is therefore both storm and drought hardy. Expected climate changes will not be a
problem and might even be an advantage for sessile oak.
Site requirements:
Sessile oak is like pedunculate oak very tolerant to different soil conditions. Ideal growth conditions
are fertile, well drained soils with ample water supply. It can also grow in poor, sandy and dry soils
and even in clay and compact soils. Sessile oak is usually considered to be better adapted to the dry
and poor soils than pedunculate oak. It should be avoided on the most water logged sites (type 9),
where Pedunculate oak is more suited. Peaty soils are to be avoided.
Sessile oak Soils with free drainage Limited

4 moderate fresh
drainage
3 moderate dry
7 80 - 160 cm
8 40 - 80 cm
6 very fresh
1 very dry
9 < 40 cm
5 fresh
2 dry
1 very poor
Nutrition
2 poor
3 moderate poor
4 moderate rich
5 rich
6 very rich
Sessile oak is a dominating tree species in three forest development sites: Oak with ash and
hornbeam (21), Oak with lime and beech (22) and Oak with Scotch pine and larch (23). Here sessile
oak is more attached to the nutrient poor types (22 and 23). Sessile oak is also mixed in a number of
forest development types, especially in the light-demanding Ash and alder (31) and Birch with
Scots pine and spruce (41) and (less than pedunculate oak) in the historical forest development
types such as Coppice forest (91), Grazing forest (92) and Pasture woodland (93).
The selection of oak provenances depends to a great extend on the site and the purpose.
Protected sites in eastern Denmark (mainly for wood production)
Sandy soils in western Denmark (mainly for landscape and ecology)
2. Norway: Selected stands (Agder)
25
Sycamore maple (Acer pseudoplatanus L.)
Sycamore maple is native to Central Europe: from N Spain, France and Belgium in west, through
Central and E Europe to the Caucasus and the Elburz mountains in Iran. Its northernmost habitat is
N Germany and the most southern is Calabria (S Italy). It has, however, naturalized many other
places and is even growing and regenerating vigorously in Tromsø in Arctic Norway. Sycamore
maple is nearly everywhere associated with beech (in the Caucasus oriental beech). It is a common
species in the low lands, but also typical in the mountain forests (German: Bergahorn) in M and S
Europe, where it is found near the timber line. It is debated whether sycamore is native or
introduced in Denmark, where it grows well and in some regions is expanding rapidly.
In the juvenile phase, sycamore maple is shade tolerant. Frequent disturbances (clearings) are,
however, necessary for continued growth. It regenerates as well after serious disasters/disturbances
such as windfalls, avalanche, landslides, etc. and is as such a typical sub-climax species. In the
lowlands it grows together with beech. As a typical gap specialist, it is developing well with ash for
example in beech dominated forest stands. Here sycamore maple seldom represents more than 5 per
cent. In the mountains, it often occurs more frequently and is typically seen with beech, Norway
spruce and silver fir. It hardly or never occurs in pure stands.
26
Sycamore maple is very climate tolerant. In the juvenile state it can be damaged by spring frost.
Due to rapid growth, however, these problems are not long lasting. Generally, it has huge
temperature amplitude and is growing well even in the cold temperate zone. It is relatively drought
tolerant. Smaller climate changes will not be a problem.
Site requirements:
Sycamore maple has a great amplitude regarding nutrient and water availability. It develops,
however, best on nutrient rich, chalky, deep grounded soils with good water supply. The roots do
not tolerate heavy soils (high volume weight) or ground water influence, but prefers good soil
structure and good air change. It is therefore sensitive to water logging in the root zone. Compared
to beech sycamore maple requires slightly better soils. Avoid peat soils and poorly drained, heavy
clay soils (water supply type 9, and the most wet of type 8). In Denmark, we have very little
experience growing sycamore maple on nutrient poor, sandy soil. Most properly, it can grow here,
but hardly with any impressive result. Dead leaves are easily decomposed.
Sycamore Soils with free drainage Limited

4 moderate fresh
drainage
3 moderate dry
7 80 - 160 cm
8 40 - 80 cm
6 very fresh
1 very dry
9 < 40 cm
5 fresh
2 dry
1 very poor
Nutrition
2 poor
3 moderate poor
4 moderate rich
5 rich
6 very rich
With its huge regeneration ability on a wide spectrum of sites, good ability to associate with other
species, and generally good ecologic skills, sycamore maple is playing an important role in nature-
based forest management. The role as an intermixture in a series of natural forest sites gives
sycamore maple a similar role in a number of forest development sites: Beech (11), Beech with ash
and sycamore (12) and Grand fir with beech and sycamore (51) up to 20 per cent, and in a number
of other forest development sites: 13, 14, 61, 61 as a ‘minor’ intermixture species.
Experiences with provenances of sycamore maple are rare, therefore, good local stands are
2. Germany: Selected stands and seed orchards (Gebiet 80101 og 02)
27
Douglas fir (Pseudotsuga menziesii (Mirb.) Franco)
Douglas fir is indigenous to NW America. There are two different types: a coastal and an inland
one. The coastal type ranges from British Columbia in Canada through the states Washington,
Oregon and down to N California. The inland-type ranges from Canada through Rocky Mountains
and with some spread populations all the way down to Mexico. The coastal type and especially the
provenances from Washington have been widespread all over Europe. In Denmark it has
successfully been used in forestry for more than 120 years.
Douglas fir is basically a pioneer species. Indigenously, it is regenerating after forest fire. Due to its
enormous growth (height up to 100 m) and a long life span (up to 1000 years), it can dominate the
forest community for ages and in this way get the role as a sub climax species. In the early
succession stages (after forest fires, etc.) it is associating with American Alder (Alnus rubra), and
later with other succession species such as Sitka Spruce (along the coast), grand fir, thuja, hemlock
and many others. Without any new catastrophes (forest fires) during the succession stages, the
development will after 600 to 800 years turn into a shade tolerant climax community dominated by
western hemlock.
In Denmark Douglas fir is mainly growing in even-aged stands. It has, however, also been
successful in mixed stands with other coniferous trees (grand fir, Sitka spruce and Norway spruce)
and it is also associating well with beech. It regenerates willingly, especially on lighter soils.
28
Douglas fir develops well in the mild, coastal climate in Denmark. In spite of its pioneer
characteristics, the use of it is limited due to climatic damages in the early succession. It is often
damaged by late spring frost and by frost desiccation in connection to black frost in late winter.
Consequently, it is regenerating best under shelter wood or in half-shade. The species is rather
drought resistant, especially in deep soils. It is easily wind crooked (sword-shaped) and the first 30-
40 years very storm sensitive. Later on it is more storm hardy. Expected climate changes will not be
a problem for Douglas fir.
Site requirements:
Douglas fir has very large soil amplitude and is growing fairly well even on rather dry and poor
soil. For optimal growth it requires deep soil conditions. Soils with hard pans and layers with high
density should be omitted. Generally, badly drained soils should be avoided, too.
Douglas fir Soils with free drainage Limited

4 moderate fresh
drainage
3 moderate dry
7 80 - 160 cm
8 40 - 80 cm
6 very fresh
1 very dry
9 < 40 cm
5 fresh
2 dry
1 very poor
Nutrition
2 poor
3 moderate poor
4 moderate rich
5 rich
6 very rich
Douglas fir is, due to its ecological profile and excellent growth the non indigenous tree species
with the greatest potential in nature-based forest management. Therefore, it is playing an important
role in a number of forest development types especially together with beech: Beech with Douglas
fir and larch (13), Beech and spruce (14), Spruce with beech and sycamore (51), Douglas fir, spruce
and beech (61), Silver fir/grand fir and beech (71).
The selection of the right provenance is of outmost importance for Douglas fir. Many provenance
trials have been established. They show that Douglas fir from the western slopes of the Cascades
and the Olympic Mountains in Washington perform the best under Danish conditions.
1. Denmark: Selected seed stands and seed orchards
2. France: Seed orchard FP 232 (selected in Danish stands of Darrington origin)
3. USA/Washington: Seed zones 030, 222, 231, 240
29
Grand Fir (Abies grandis Lindl.)
Grand fir is indigenous to NW America, where it ranges in two areas: 1) The western, marine parts
round Vancouver Island and the western areas of the states of Washington and Oregon, and 2) The
eastern, continental part through Rocky Mountains from SE of British Columbia, Idaho, Montana
and E Washington and Oregon.
Grand Fir is a shade tolerant tree, a typical climax species connected to late successional stages.
Together with thuja and western hemlock it will slowly replace other pioneer species such as
Douglas (west) and Ponderosa fir (east) unless a forest fire is starting a new succession order. Most
interesting from a Danish point of view is the western coastal area. Here, it is most common as a
solitaire intermixture or in small groups with Douglas fir, thuja and western hemlock. In the eastern
continental area it appears mixed with Ponderosa pine and Douglas fir.
In Europe grand fir has been cultivated for almost 100 years. In Danish forestry it’s become
important due to its productivity (highest volume yield of all tree species) and because it replaces
silver fir as a climax species on poorer and dryer soils. Grand fir is excellent in mixed groups with
other species and is regenerating well, especially on lighter soils.
30
Like a typical shade tolerant tree, grand fir is sensitive to early frost, but not quite as much as silver
fir. On the other hand, the species is very drought tolerant, partly due to its ability for deep rooting
even in heavy gley soils with pseudogley. Grand fir is expected to adapt well to future climate
changes.
Site requirements:
Grand fir is growing well on a wide water- and nutrient gradient, from very poor to very rich, and
from very dry to ground water influenced soils. Like silver fir, its root system is tolerating wet,
alternately ground water influenced rich clay soil (type 8). On dry soils it is less water demanding
than common silver fir and is generally growing well here. Therefore, in Denmark it’s most
common on light and dry soils in Djursland and East Jutland. Probably, it is unstable on peat soils
and in extreme ground water influenced, humus rich gley soils. Grand fir has very fast leaf
convertibility; the fastest of all coniferous trees.
Grand fir Soils with free drainage Limited

4 moderate fresh
drainage
3 moderate dry
7 80 - 160 cm
8 40 - 80 cm
6 very fresh
1 very dry
9 < 40 cm
5 fresh
2 dry
1 very poor
Nutrition
2 poor
3 moderate poor
4 moderate rich
5 rich
6 very rich
The ecologic profiles of grand fir and silver fir are very much alike. Grand fir has, however, a
somewhat higher growth and is also more frost tolerant. In various forest development types, grand
fir and silver fir are substituting each other. On poor soils with less rainfall grand fir is preferred,
however. In the following forest development types grand fir is an important stand element: Beech
with fir (14), Fir with beech and sycamore (51), Douglas fir, Norway spruce and beech (61), Silver
fir/grand fir and beech (71).
Danish seed sources are in general good. For direct import of seed the western coastal area in
Washington is most interesting from a Danish point of view.
2. Washington, seed zone 221, 222, 212, 231, 232
3. British Columbia, Vancouver Island (east coast)
31
Larch (Larix dicidua Mill., Larix kaempferi Carr. and Larix x eurolepis)
European larch is native to central European mountains, growing at 600-2300 m in mixed stands
with spruce, beech, silver fir, sycamore and rowan. Certain places it is bordering the timber line
with mountain pine.
Japanese larch is primarily distributed centrally in the temperate forest zone of main island Honshu.
Here it is growing in mixed groups with i.e. Abies veitchii, Tsuga diversifolia and Pinus densiflora.
Hybrid larch is an artificial hybrid between selected trees of European and Japanese larch. The
hybrid combines European larch’s excellent shape with Japanese larch’s resistance against larch
canker. Besides, hybrid larch displays a considerably higher growth.
Originally, the European larch (Larix decidua) was imported into Denmark, but due to a high
predisposition to larch canker, Japanese larch (Larix kaempferi) has taken over its place. After the
development and improvement of hybrid larch (Larix x eurolepis) it has become the most widely
used. There is, however, still some uncertainty about segregation (F2-generation), i.e. when hybrid
larch is grown with natural reproduction in view. Larch is a typical pioneer species and like Scotch
pine very light demanding, but more shady. In open fields it is a natural successor after fire,
landslide, and avalanches or after end grassing. Very fast it is establishing a forest climate.
Distribution of Japanese larch (left) and European larch

(above)
Larch in general and especially hybrid larch are frost hardy and drought tolerant. Unlike most other
species, larch has its shoot elongation mid August and therefore it is to a certain extent sensitive to
late summer drought. It is rather storm tolerant, but will often get a bad shape if there is too much
32
wind. Larch and especially hybrid larch are supposed to be very both robust to climate changes
except for extreme summer drought.
Site requirements:
The site amplitude of larch is huge. It develops well on poor and rich soils and is not especially
water demanding. The roots are not tolerating soil horizons with pseudogley or gley, and it is
unsuitable on compact, oxygen poor soils. Especially Japanese larch is sensitive to late summer
drought. For this reason hybrid larch is more suitable on dry sites than Japanese larch. Larch leaves
are not easily decomposed and the species often form thick layers of non decomposed needles.
Larch Soils with free drainage Limited

4 moderate fresh
drainage
3 moderate dry
7 80 - 160 cm
8 40 - 80 cm
6 very fresh
1 very dry
9 < 40 cm
5 fresh
2 dry
1 very poor
Nutrition
2 poor
3 moderate poor
4 moderate rich
5 rich
6 very rich
As a robust pioneer species larch has an important function as a nurse tree in afforestation and/or
after windfalls, etc. It has also a more permanent role in light-demanding tree development types
such as Oak with pine and larch (23), and in more climax ones, i.e. Beech with Douglas fir and
larch (13), Beech and spruce (14), Spruce with beech and sycamore maple (51), Sitka spruce with
fir and broadleaf (52), Silver fir/grand fir and beech (71).
European larch is hardy used anymore. Due to successful long-term breeding programs in larch a
large number of excellent seed sources – seed orchards and seed stands - are available. Primarily
hybrid larch should be used. However, due to uncertainties about negative out crossing effects in
the F2-generation Japanese Larch should be considered when used in permanent mixtures and
natural regeneration.
1. Denmark: Seed orchards (Hybrid larch and Japanese larch)
2. Denmark: Selected seed stands (Hybrid larch and Japanese larch)
3. Germany: Japanese larch from selected stands and seed orchards (Gebiet 83901)
33
Norway Spruce (Picea abies (L.) Karst.)
Norway spruce is growing throughout Europe. It is naturally distributed in the mountains of Central
and SE Europe (the Vosges, the Black Forest, the Alps, the Harz, the Erzgebirge, the Bavarian
Forest, the Bohemian Forest, the Fichtel Mountains, the Sudeten, the Tatra and the Carpathians) up
to 500-2000 m. Furthermore, Norway spruce is widely spread in the lowlands of the Baltic region
and in Russia and in most of the Scandinavian Peninsula. Norway spruce is not native to Denmark
and was introduced successfully 250 years ago. With approx. 27 per cent of the total forest area, it
is presently our most widespread forest tree.
Norway Spruce is between pioneer- and climax species. It regenerates well both after disasters
(often with birch and rowan) but also group wise in an established forest stand. Usually it is
growing in mixed stands with beech, silver fir, sycamore, larch and others (in Central Europe) and
with Scots pine, birch, ash (in N and E Europe). In natural monocultures you only find it near its
Alpine limit.
Norway spruce grows naturally in a cold temperate, sub-continental climate. Although it is very
robust and frost tolerant in the regeneration period, it is sensitive to summer drought, wind and salt.
34
It is also very sensitive to storms and exposed edges are unstable. Its climatic optimum is at a mean
temperature between 5 - 7.5 degrees centigrade and with rainfall in the vegetative period of more
than 550 millimetres. These climatic conditions for Norway spruce are not fulfilled in Denmark.
Consequently, it shows pronounced decline symptoms after mild winters and in periods with
summer drought. Since the expected climatic changes seem to increase these problems, the future
for Norway Spruce seems somewhat bleak.
Site requirements:
The nutrient requirements of Norway spruce are low, but it is sensitive to water logging and very
dry soils should be avoided. On heavy clay and alternate wet soils Norway spruce is declining at an
early age and exposed to windfall. The development of the root system is very sensitive and
dependent on the soil conditions. It develops a root system very close to the surface when growing
on pseudogley or gley soil horizons, making it sensitive to drought and storms. On deep grounded
soils it will similarly develop a deep root system. Its dead leaves are decomposing very slowly and
it is developing many fine roots in the typical humus layer.
Norway spruce Soils with free drainage Limited

4 moderate fresh
drainage
3 moderate dry
7 80 - 160 cm
8 40 - 80 cm
6 very fresh
1 very dry
9 < 40 cm
5 fresh
2 dry
1 very poor
Nutrition
2 poor
3 moderate poor
4 moderate rich
5 rich
6 very rich
Norway spruce is the less acclimatized of our non-indigenous forest trees. Expected climatic
changes might worsen its adaptability to Danish growing conditions. It affects the role of the tree
species and its application in the tree development types. It appears in many types, but always in
mixed stands and never as the dominant species. As such you find it in: Beech with Douglas fir and
larch (13), Beech with spruce (14), Birch with Scots pine and spruce (41), Spruce with beech and
sycamore (51), Douglas fir, spruce and beech (61), Silver fir/ grand fir and beech (71) and Scots
pine, birch and spruce (81).
The frequent stability and health problems in Norway spruce underline the importance of climatic
adaptation and adaptability when selecting provenances in Norway spruce.
1. Denmark: Selected seed orchards and seed stands
2. Germany: Selected seed orchards and seed stands (Gebiet 84001, 02, 06)
35
Scotch pine (Pinus silvestris L.)
Scotch pine is native to the Scandinavian Peninsula. In Central and E Europe it is ranging in a belt
between 45° and 65° N from Germany, through Poland, the Baltic States through Russia to the
Bering Strait. In westward direction you find it in the Alps and special places in Scotland, France,
and Spain, on the Balkans and in the mountains along the Black Sea.
About 10,000 – 12,000 years ago, shortly after the last glacial period ended, Scotch pine invaded
Denmark. Only a few thousand years later, it was reduced to marginal sites by a number of other
invading climax species. Due to intensive human utilization it finally disappeared in the Middle
Ages. It was reintroduced during the 1700th century as prevention against sand drift and by
recultivation heath lands. By this reintroduction, unsuitable provenances were brought in,
unfortunately, which especially in the heath region of Jutland were unsuccessful. Therefore, Scotch
pine never became central in establishing forestry on former heath lands.
Scotch pine is a typical pioneer species, regenerating in connection with catastrophes after forest
fires, windfalls, landslides, avalanches, etc. In Scandinavia and NE Europe it is found on rather dry
sites with fire risk and often mixed with birch, ash and sometimes with spruce. In Central and S
European mountains, where it is growing on the most steep, shallow and small drought sites, it
associates with Norway spruce, silver fir, beech, sycamore and larch.
36
Scotch pine is climate tolerant and adaptable to various weather conditions, partly due to its
pronounced ability to develop races adapted to the local growing conditions. It is very frost hardy
and extremely drought resistant. Its crown can be wind formed, but is otherwise not sensitive to
wind. Due to its deep root system it is also rather storm tolerant. Although easily burnt by salt, it is
rather hardy in dune forestry. As species it will tolerate some climatic changes, problems with the
local adaptation of specific races might occur, however.
Site requirements:
Scotch pine is the one of our tree species with less site requirements. It tolerates poor, dry and wet
soils. It occurs on chalky soil and is also developing well on peat. The foliage of Scotch pine is not
easily converted and it is humus accumulating.
Scots pine Soils with free drainage Limited

4 moderate fresh
drainage
3 moderate dry
7 80 - 160 cm
8 40 - 80 cm
6 very fresh
1 very dry
9 < 40 cm
5 fresh
2 dry
1 very poor
Nutrition
2 poor
3 moderate poor
4 moderate rich
5 rich
6 very rich
With its pioneer abilities Scotch pine plays an important role when establishing forest on poor sites,
in harsh climate, and also in connection to afforestation, clear-cutting or after windfalls. In this way
it can facilitate the reestablishment of the forest climate. Further it is included in a number of light-
demanding forest development types on dry sites: Oak with Scotch pine and larch (23), Birch with
Scotch pine and spruce (41), Sitka spruce and pine with deciduous forest (52), Scotch pine, birch
and Norway spruce (81), and also mixed in a number of others.
Protected sites
1. Denmark: Selected seed stands and seed orchards of Baltic origin
2. Germany: Seed orchards of east Prussian origin
3. Latvia, Lithuania and north-east Poland: Selected stands
4. Sweden: Selected stands from Bohus län
Exposed sites
1. Denmark: Selected seed stands and seed orchards of Norwegian and Scottish origin
2. Norway: Selected stand from southwestern Norway
3. Sweden: Selected stands from southwestern Sweden
37
Silver fir (Abies alba Mill.)
Silver fir is naturally distributed in the mountain regions of Central and S Europe, abundantly
mixed with beech, sycamore, spruce and occasionally with larch. It is rarely found as monoculture.
Outside its natural range it is used in the coastal regions of N Germany and Denmark. Experiments
with different provenances of silver fir have shown big differences. In Denmark’s oceanic climate
provenances from S Italy (Calabria over 1600 m) have shown superior health and growth.
Silver fir is the most typical climax species of our coniferous trees. It regenerates willingly in shade
and can for decades survive in deep shadow and resume growth again when being liberated. Climax
communities with beech and spruce are being established many places in Central Europe.
Due to early burst Silver fir is very sensitive to spring frost. Therefore, it regenerates almost solely
on already established forest sites (under canopy or in forest edges). After clear cutting or at bare
land afforestation, preculture of some kind is necessary as nurse trees alone seldom are sufficient.
Silver fir needs frequent rainfalls (more than 800 mm yearly) or alternatively high atmospheric
humidity to develop. Abundant rainfall is more important for silver fir than nutrients. The reason
why the species is so widespread in the dune forestry is due to its extreme salt tolerance and the
38
need of high atmospheric humidity. Silver firs expected reaction to the assumed climate changes is
to be seen in relation to choice of provenances. Central European silver fir provenances with limited
adaptability will be problematic. Calabrian provenances (S Italy) should be able to adapt.
Site requirements:
Silver fir tolerates both wet clay soils as well as poor sandy soils. The most dry and extremely
ground water influenced soils should be avoided. The species is sensitive to lack of rainfall and
should not be planted in such sites. It tolerates calcareous soil and is less sensitive to stagnating
water in the rooting zone than Norway spruce. It has a well developed root system, and silver fir is
the coniferous tree with the deepest root system on poorly drained soils.
Silver fir Soils with free drainage Limited

4 moderate fresh
drainage
3 moderate dry
7 80 - 160 cm
8 40 - 80 cm
6 very fresh
1 very dry
9 < 40 cm
5 fresh
2 dry
1 very poor
Nutrition
2 poor
3 moderate poor
4 moderate rich
5 rich
6 very rich
Silver fir is only a dominating tree species in one forest development type i.e. Silver fir/Grand fir
and beech (71). This forest development type with the typical climax species is the closest we get in
Denmark to the “selection forest system” or ‘Plenterwald’- structure) with the intensive mixture of
species and age classes. In forest development types: Spruce with beech and sycamore (51), Sitka
spruce and pine with broad-leaved trees (52) and Douglas fir, Norway spruce and beech (61), Silver
fir and Grand fir can be minor but ecological important elements. Especially on soils with good
water supply and/or in areas with high atmospheric humidity as in sand dunes for example this is
possible.
Trials with silver fir have shown that provenances from South Italy (Calabria) and from the East
Carpathians in Romania exhibit a far better growth, health and adaptability than rovenances from
Central Europe do.
1. Italy: Italien/Calabrien, Gariglione nr. 120, 1600-1750 m)
2. Romania, Maramures/ Lapos
39
Sitka spruce (Picea sitchensis (Bong.) Carr.)
Sitka spruce is spread along the Pacific coastline in a narrow zone from N Alaska to S California.
The total distance from north to south is almost 3000 kilometres or 1700 miles, while the distance
from east to west is merely 210 kilometres or 130 miles. The species is solely connected to the
Pacific coastline, where it is found in the lowland (below 300 metres). Danish forestry has used
Sitka spruce for about 100 years. The use of it has been rising for the last decades.
Like Norway spruce it has qualities both as a pioneer and as a climax species. Even-aged Sitka
stands are quite common, especially when the pioneer qualities can flourish after a fire or earth
erosion. Under stabile conditions the species is mixed with hemlock, thuja, Douglas fir, grand fir.
Sitka spruce is strikingly adapted to a coastal climate with mild winters and temperate summers,
almost like the climate we are having in Denmark. Further, it is acclimatized to damp and saline
40
surroundings, like in dune forests. Generally, Sitka spruce is not very stable in Denmark. One year
it is looking healthy and in good growth. A few years later it is in total decay, probably due to
improper water supply. Sometimes it is weakened by summer drought followed by attacks of giant
bark beetles (Dendroctonus micans). Because it is very sensitive to late spring and early autumn
frost especially during the cultivation stadium, it is often being damaged by frost on exposed sites.
Although the species is wind tolerant, it is easily overthrown by storm. As a typical oceanic species,
Sitka spruce will probably tolerate expected climate changes, except from very dry summers which
might be problematic.
Site requirements:
Sitka spruce is developing well on many different sites. It is growing much better than Norway
spruce on poor dry sites and can tolerate salt deposition. It is also doing much better than Norway
spruce on soils with limited drainage (type 8). On very groundwater influenced soils (water supply
type 9), especially on rich clay soils with bad air change and on peat soil, Sitka spruce is unsuitable
due to risk of storm felling. On drained peaty soil it is regenerating fast, but decays often at young
age. The dead leaves have a fast turnover and the species is a typical raw humus builder.
Sitka spruce Soils with free drainage Limited

4 moderate fresh
drainage
3 moderate dry
7 80 - 160 cm
8 40 - 80 cm
6 very fresh
1 very dry
9 < 40 cm
5 fresh
2 dry
1 very poor
Nutrition
2 poor
3 moderate poor
4 moderate rich
5 rich
6 very rich
Sitka spruce has and will have a certain role in future forestry. Especially in the dune region where
its ecological advantages (salt tolerance) can develop in full. On other sites it complements Norway
spruce due to different climate profile. As already mentioned, you find the species in typical dune
forest development types: Sitka with pine and broadleaf tree (52) and in a number of other types as
supplement to Norway spruce: Beech and spruce (14), Birch with Scots pine and spruce (41),
Spruce with beech and sycamore (51), Silver fir/grand fir and beech (71), and Scots pine, birch and
spruce (81).
Protected sites
1. Denmark: Selected seed orchards and seed stands (Washington type)
Frost and wind exposed sites
1. Denmark: Selected seed orchards and seed stands (Queen Charlotte type)
2. British Columbia: Selected stands (Queen Charlotte Isl.)
41
Literature:
Ellenberg, H., 1996: Vegetation Mitteleuropas mit den Alpen in ökologischer, dynamischer und
historischer sicht, 5 edn, pp. 4-1095. Ulmer, Stuttgart.
Köstler, J.N., Brückner, E., & Bibelriether, H., 1968: Die Wurzeln der Waldbäume, pp. 11-283.
Verlag Paul Parey, Hamburg und Berlin.
Larsen, J.B. (Red.), 1997: Træarts- og Proveniensvalget i et bæredygtigt skovbrug. Dansk

Skovbrugs Tidsskrift, 82, 1 - 252
Röhrig, E., Bartsch, N., 1990: Waldbau auf ökologischer Grundlage / begründet von Alfred Dengler
Bd. 2: Baumartenwahl, Bestandesbegründung und Bestandpflege, 314 s
Savill, P.S., 1991: The silviculture of trees used in British forestry. Impressum Wallingford : C.A.B.
International, cop. 1991 pages etc. ix, 143 s., ill.
Schütt, P., Schuck, H.J. Aas, G., Lang, U.M., 1994: Enzyklopaedie der Holzgewächse. Handbuch
und Atlas der Dendrologie. Ecomed Verlag, Landsberg.
42

Larsen Et Al. Ecology of Tree Species and Selection

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Larsen Et Al. Ecology of Tree Species and Selection

Uploaded by

Copyright:

Available Formats

Ecology of tree species and species selection

J. Bo Larsen, Karsten Raulund Rasmussen and Ingeborg Callesen

1. Native range and role in the forest community

Beech Soils with free drainage Limited

Ecological diagram for beech (Fagus silvatica)

4. Matching species to sites and to forest development types:

Alder (Alnus glutinosa (L.) Gaertn.) 4

Ash (Fraxinus excelsior L.) 6

Beech (Fagus silvatica L.) 8

Birch (Betula pendula Roth. and Betula pubescens Ehrh.) 10

Cherry (Prunus avium L.) 12

Hornbeam (Carpinus betulus L.) 14

Lime (Tilia cordata Mill.) 16

Mountain ash (Sorbus aucuparia L.) 18

Norway maple (Acer platanoides L.) 20

Pedunculate oak (Quercus robur L.) 22

Sessile oak (Quercus petraea Liebl.) 24

Sycamore maple (Acer pseudoplatanus L.) 26

Douglas fir (Pseudotsuga menziesii (Mirb.) Franco) 28

Grand Fir (Abies grandis Lindl.) 30

Norway Spruce (Picea abies (L.) Karst.) 34

Scotch pine (Pinus silvestris L.) 36

Silver fir (Abies alba Mill.) 38

Sitka spruce (Picea sitchensis (Bong.) Carr.) 40

Native range and role in the forest community:

Red alder Soils with free drainage 4 moderate fresh Limited

Matching species to sites:

1. Denmark: Selected seed stands

Native range and role in the forest community:

Ash Soils with free drainage Limited

Matching species to sites:

Native range and role in the forest community:

Beech Soils with free drainage Limited

Matching species to sites:

Native range and role in the forest community:

Birch Soils with free drainage Limited

Matching species to sites:

The following recommendations are for both species of birch.

Native range and role in the forest community:

Cherry Soils with free drainage Limited

Matching species to sites:

Native range and role in the forest community:

Hornbeam Soils with free drainage Limited

Matching species to sites:

Native range and role in the forest community:

Lime Soils with free drainage Limited

Matching species to sites:

Native range and role in the forest community:

Mountain ash Soils with free drainage Limited

Matching species to sites:

Native range and role in the forest community:

Norway maple Soils with free drainage Limited

Matching species to sites:

Native range and role in the forest community:

Pedunculate oak Soils with free drainage Limited

Matching species to sites:

Native range and role in the forest community:

Sessile oak Soils with free drainage Limited

Matching species to sites:

Native range and role in the forest community:

Sycamore Soils with free drainage Limited

Matching species to sites:

Native range and role in the forest community: