24 European forests: an ecological overview

European forests: an ecological overview
D. de Rigo, T. Houston Durrant, G. Caudullo, J.I. Barredo
Ecosystems may be classified into a variety of groups or
zones according to their homogeneity1-9 . An Ecological
Zone may be defined as an area with broad yet relatively
homogeneous natural vegetation formations that are
similar, although not necessarily identical.
Several key aspects of forest resources are influenced at the
global and continental scale by the ecological zone where
a given forest or woodland grows10-17. Even at the regional
and country scale, the distribution of environmental and
ecological zones may contribute to the understanding of
core differences in local forest ecosystems18-22 .
For its Global Forest Resources Assessment, the Food and
Agriculture Organization of the United Nations (FAO) produced
a global ecological zoning classification1, 2 . The FAO zoning is
exploited by a rich variety of applications11-13, 17, 23-25 .
In this chapter, the classification of FAO is summarised and
complemented by a qualitative analysis of the secondary
ecozone components which may coexist in a given
European area as subordinate constituents of the local
forest ecosystems. These components have been derived
by means of a robust fuzzy analysis of climatic similarity26 .
The FAO Ecological Zones
Boundaries of the Ecological Zones approximately coincide
with Köppen-Trewartha climatic types, which are based on
temperature and rainfall27, 28 .
At the first level a total of five domains are distinguished
based on temperature: Tropical, Subtropical, Temperate, Boreal
and Polar. Four of these domains can be found in Europe
(Tropical is absent).
At the second level, precipitation is also used to subdivide
the domains into a total of 20 subclasses, of which 12 can be
found across the European continent.
"Mountain systems" are classified as a separate Ecological
Zone in each domain and are characterized by a high variation
in both vegetation formations and climatic conditions1, 2, 27.
Temperate domain
The temperate domain lies in a region where average
temperatures above 10 °C can be found from 4 to 8 months of the
year. The temperate domain covers a large area across mainland
Europe and is subdivided into Temperate Oceanic Forest, Temperate
Continental Forest, Temperate Steppe and Temperate Mountain
Systems.
Temperate Oceanic Forest is typically found on the western or
windward side of the continent and has the mildest climate of the
four temperate zones. The average monthly temperature is always
above 0 °C and there is adequate rainfall in all seasons. The annual
rainfall may vary quite significantly from 400-800 mm in lowlands
up to 2 000-3 000 mm on windward lower coastal mountain slopes.
The main vegetation type in these areas is deciduous broadleaved
forest. In Western Europe a typical example is beech.
Temperate Continental Forest may be found in the interior
and eastern areas of the continent. Winters are colder, with
a shorter frost-free season and at least one month having
average temperatures below 0 °C. There is a larger annual range
of temperatures than in the Oceanic zone. Rainfall generally
decreases with distance from the ocean and also at the higher
latitudes. Similar to the Oceanic zone the main vegetation cover is
deciduous broadleaved forest or mixed forest: a typical example in
Europe is oak-hornbeam in Central Europe.
Temperate Steppe is found in the deep interior of the continent
and is characterised by cold winters and relatively low rainfall (200400 mm per year). Evaporation exceeds precipitation. The vegetation
is dominated by grass and low shrubs.
Temperate Mountain Systems are found in the Alps and the
Pyrenees in Europe. They share several characteristics of the boreal
zone and are snow covered for large parts of the year. Typical
vegetation is pine forest.
Fig. 1: A key factor in the local variability of ecosystems is the influence
of water bodies. Water resources, such as lakes or rivers, may exercise a
variety of effects on the surrounding microclima. Beside providing additional
humidity, they may contribute to mitigate the temperature range inducing a
stabler pattern, and influence the local geomorphology - being a factor e.g.
in shaping the slope of mountain and hill sides.
Top: Subtropical dry ecological zone. France, Provence-Alpes-Cote d'Azur.
(Copyright Jean Latour, CC-BY, http://archive.is/vqbxT)
Bottom: Temperate oceanic ecological zone. The Saar river influences part
of northeastern France and western Germany.
(Copyright Wolfgang Staudt, CC-BY, http://archive.is/2XBaO)
Introduction
Boundaries of the Ecological Zones approximately coincide
with Köppen-Trewartha climatic types, which are based on
temperature and rainfall27, 28 .
At the first level a total of five domains are distinguished
based on temperature: Tropical, Subtropical, Temperate, Boreal
and Polar. Four of these domains can be found in Europe
(Tropical is absent).
At the second level, precipitation is also used to subdivide
the domains into a total of 20 subclasses, of which 12 can be
found across the European continent.
“Mountain systems” are classified as a separate Ecological
Zone in each domain and are characterised by a high variation
in both vegetation formations and climatic conditions1, 2, 27.
Subtropical domain
The subtropical domain is generally characterised by having
on average at least 8 months above 10 °C.
It occurs throughout the southern regions of Europe and is
divided into Subtropical Humid Forest, Subtropical Dry Forest,
Subtropical Steppe and Subtropical Mountain Systems.
The Subtropical Humid Forest zone has high humidity every
month with annual rainfall usually over 1 000 mm distributed
throughout the year. Relatively few regions of Europe can be
classified as Subtropical Humid Forest, but some may be found on
the northern coast of Turkey, north-eastern Spain and parts of central
south-east Italy. Vegetation may comprise evergreen broadleaved
forest, evergreen coniferous forest and deciduous forest.
Subtropical Dry Forest is the typical Mediterranean climate with
dry, hot summers and humid, mild winters with an annual rainfall
of 400-900 mm. Found throughout the Mediterranean Basin, typical
vegetation is sclerophyllous evergreen forest, woodland and shrub,
for example maquis dominated by Quercus ilex.
In the Subtropical Steppe zone, evaporation generally
exceeds precipitation. These regions are found at the southern
parts of the Mediterranean Basin and vegetation is dominated
by shrubs adapted to arid environments.
Subtropical Mountain systems can be found in the southern
mountain regions of Europe and the Middle East. The vegetation may
be quite varied depending on the altitude, exposure and humidity.
Fig. 2: FAO ecological zones for forest reporting (2010).
Polar
Temperate mountain system
Subtropical mountain system
Boreal tundra woodland
Temperate steppe
Subtropical steppe
Boreal mountain system
Temperate continental forest
Subtropical dry forest
Boreal coniferous forest
Temperate oceanic forest
Subtropical humid forest
Note: Only the main ecological components are represented.
24
European Atlas of Forest Tree Species | Introduction
Boreal domain
The boreal, or subarctic, domain is characterised by a large
annual range of temperatures, with one to four months with average
temperatures above 10 °C and generally low levels of rainfall, usually
below 500 mm. The name taiga has been given to the subarctic
lands of Eurasia with their extensive coniferous forests. This domain
is found across the northern regions of Eurasia and is subdivided
into Boreal Tundra woodland, Boreal Coniferous Forest and Boreal
Mountain systems.
In Boreal Coniferous Forests the summers are short with
at most 3 months having temperatures above 10 °C. Winters are
long and cold. Geographically this zone covers the northern part of
Eurasia, and is typified by dense coniferous forests, dominated by
spruce and fir in northern Europe and western Siberia, and larch in
central and eastern Siberia.
Boreal Tundra occurs at the northern limit of the Boreal zone,
where it meets the Polar domain. Climatic conditions are similar to
the Boreal Coniferous Forest zone but are colder and more extreme
with very low winter temperatures and permafrost. Species are
similar to those found in the Boreal Coniferous Forest zone, but the
vegetation cover is more open.
Boreal mountain systems may be found in parts of Norway
and the eastern part of the Russian Federation. Temperatures are
extremely cold and there is continuous permafrost. Vegetation
comprises open woodlands and scrub.
Polar domain
In the Polar domain there are months with average
temperatures below 10 °C throughout the year. There are no
sub-divisions in this domain as it is generally only very sparsely
vegetated. In Europe there is a small Polar region in the very
northern tip of Scandinavia.
Fig. 3: Local-scale pattern of ecological components. Orography, prevailing wind direction and intensity, slope, aspect
and the presence of surrounding peaks influencing the local solar radiation and rain shadow effects are among the
factors able to alter vegetation at very local scales by affecting the local pattern of disturbances and the availability of
resources.
Top: canton of Bern, Switzerland. Forest areas and grassland alternating along the peaks of a mountain ridge. Human
influence is evident in the complex patchiness of pastures/grassland fragmenting the forests in the lower part of the
valley. Along the mountain ridge, the impact of differential solar radiation, dominant winds and peak-induced rain
shadow effects may contribute to the regularity of the pattern of forests (in the picture, shadow side of the peaks) and
grassland/sparse vegetation (sunny side and saddle landforms). See also Figure 5.
(Adapted from an image authored by Vasile Cotovanu, CC-BY, https://archive.is/8dGPq)
Middle: Northern Sweden subarctic landscape. In the foreground, a dominant ecological component of boreal tundra woodland
is visible. However, the water body on the right and the partial wind protection offered by the corresponding valley allow a
boreal coniferous forest component to survive.
(Adapted from an image authored by Alexander Cahlenstein, CC-BY, https://archive.is/AxnHo)
Bottom: Retezat Mountains, Romania. At very local scale, the transition between sparse trees (right of the picture),
pine shrubland (right side of the two small valleys, i.e. their left bank) and grassland (left side of the picture) is clearly
connected with the aspect of the banks. See also Figure 6. Wider scale ecosystem characteristics may be influenced by
these details (for example, average connectivity and fragmentation of forest/shrub patches, average availability of core
undisturbed patches40-43).
(Adapted from an image authored by Horia Varlan, CC-BY, https://archive.is/aIL3J)
Introduction | European Atlas of Forest Tree Species
25
Polar
Boreal Tundra
Boreal Mountain
26
European Atlas of Forest Tree Species | Introduction
Boreal Coniferous
Temperate Mountain
Temperate Steppe
Introduction | European Atlas of Forest Tree Species
27
Temperate Continental
Temperate Oceanic
Subtropical Mountain
28
European Atlas of Forest Tree Species | Introduction
Subtropical Steppe
Subtropical Dry Forest
Subtropical Humid Forest
Introduction | European Atlas of Forest Tree Species
29
Fig. 5: The transition between different bioclimatic conditions and ecosystems
(even within the same ecological zone) may occasionally happen along sharp
boundaries. Slovakia, Western Carpathians. The sides of this mountain ridge
show a sudden transition between a dense coniferous forest and a grassland
with sparse trees. Different solar radiation, dominant winds and induced rain
shadow effects may contribute to this kind of transition along mountain ridges
(see also Figure 3). Anthropogenic factors such as delimited pastures may also
induce or reinforce these sudden transitions. The different ecosystems may
even influence the local micro-climate: for example, coniferous forests may
have a lower albedo than areas with predominant grassland - with subsequent
differential radiation/warming feedbacks10, 29, 30 .
(Adapted from an image authored by Ján Sokoly, CC-BY, http://archive.is/Oxamq)
FIg. 4: The transition between ecological zones sometimes happens within
relatively low distances.
Top: Italy, geographic sudden transition between the Alps (temperate
mountain system) and the Po valley (whose sparse forests are
predominantly characterised by a temperate oceanic climate). Although this
transition may be smoother in other areas of the Alps, in the picture the
urban/agricultural landscape (foreground, Schio, Italy) lies at an elevation
of less than 300 m above sea level while the mountain peaks (background,
at a distance of few kilometres) are more than 1000 m higher. Ecological
zones classified according to the Global Forest Resources Assessment of
the Food and Agriculture Organization of the United Nations (FAO FRA) 1, 2.
(Adapted from an image authored by Doc Searls, CC-BY, https://archive.is/pGEPG)
Middle: France, Massif Central, Puy de Sancy (foreground) and the contiguous
plain with low elevation hills (background). Even in this case, the sudden
transition is between temperate mountain ecosystems to temperate oceanic
forests. The transition is smoother in other areas of the Massif Central.
(Adapted from an image authored by Patrice, CC-BY, https://archive.is/BcSK5)
Bottom: Wider scale view of the Alpine transition between the temperate
mountain system and the surrounding plains hosting fragmented temperate
oceanic forests.
(Adapted from an image authored by Francisco Antunes, CC-BY, http://archive.is/vy5Fa)
Fig. 6: Norway, Femundsmarka National Park at the transition between the ecological zone of the boreal
mountain system and that of the boreal coniferous forests.
Top: Compared to the sometimes sudden transitions highlighted in Fig. 4, in this region the component of
boreal coniferous forests may show a smoother shift towards the boreal mountain component.
(Adapted from an image authored by Mahlum, PD, http://archive.is/r23pC)
Bottom: Detail of a typical forest ecosystem in the region.
(Adapted from an image authored by Robert Anders, CC-BY, https://archive.is/1EIGx)
30
European Atlas of Forest Tree Species | Introduction
Pages 26-29:
Polar zone: polar shrub vegetation in Sør-Varanger (Finnmark, Norway).
Temperate continental zone: mixed broadleaved forest on Veľká homoľa mountain near
Modra (Bratislava, Slovakia).
(Copyright Soldatnytt, CC-BY, https://archive.is/yiNIK)
(Copyright Ján Sokoly, CC-BY, https://archive.is/WRHEj)
Boreal tundra zone: shrub tundra vegetation in northern Kola Peninsula (Murmansk,
Russia).
Temperate oceanic zone: mixed broadleaved forest along Ystwyth river (Ceredigion,
Wales).
(Copyright Ninara, CC-BY, http://archive.is/AXysf)
(Copyright Ruben Holthuijsen, CC-BY, https://archive.is/0JCCJ)
Boreal mountain zone: boreal forests on the side of the cliff over the Aurlandsfjorden
(Sogn, Norway).
Subtropical mountain zone: mediterranean mixed forest on mountain area of the
Pollino National Park (south Italy).
(Copyright Stan, CC-BY, https://archive.is/I1IvB)
(Copyright Brian Gratwicke, CC-BY, https://archive.is/47l9B)
Boreal coniferous zone: coniferous forest by Hundtjärnen lake near Floda (Dalarna,
central Sweden).
Subtropical steppe zone: steppe dry grassland on Djurdjura Massif of the Tell Atlas
chain (Kabylie, Algeria).
(Copyright Taxelson, CC0, https://archive.is/KEuAl)
(Copyright Atif Rafik, CC-BY, https://archive.is/Chq1H)
Temperate mountain zone: mixed broadleaved and coniferous stands in the Black
Forest (Freiburg, Germany).
Subtropical dry forest zone: sclerophyllous evergreen vegetation on the coasts of the
Fethiye Gulf (Muğla, south-western Turkey)
(Copyright ilovebutter, CC-BY, https://archive.is/mNK71)
(Copyright Jorge Franganillo, CC-BY, https://archive.is/ONm91)
Temperate steppe zone: steppe grasslands near Poltava (Poltava Oblast, Ukraine).
Subtropical humid forest zone: mixed broadleaved and coniferous forest on the side of
Pontic Mountains near Trabzon (Trebisonda, Turkey).
(Copyright Vlad Butsky, CC-BY, http://archive.is/qf8qf)
(Copyright Aleksasfi, PD, https://archive.is/W8Ffz)
Local ecology characterisation by means of
Ecological zones components
(Copyright Alexandru Badarau, AP, https://archive.is/mQfqE )
This implies the existence of forest areas where the main
ecozone component (e.g. a forest whose natural vegetation
formations are predominantly similar to those typical of
boreal coniferous forests) is complemented by other secondary
components (e.g. ancillary characteristics typical of temperate
continental forests).
In order to make a first attempt to account for these
phenomena and to model a pan-European ecological zoning with
a more gradual transition between the zones (where needed),
for each ecozone a qualitative fuzzy set map was computed26
by estimating the similarity of each grid-cell’s climatic and
geographic conditions to the distribution of values typically
observed within the corresponding FAO ecozone.
The modelling of robust fuzzy ecological zones has been
based on a geospatial application32 of the semantic array
programming paradigm33, 34 . The similarity analysis exploited
the relative distance similarity (RDS) approach which is designed
to support semantically enhanced, scalable data-transformation
models35-39 . The set of maps on pages 26-29 show the most
prevalent ecozones of Europe along with where the computed
qualitative fuzzy boundaries might lie.26 Further details on the
modelling aspects can be accessed in the full online version of
this chapter.
At local scale, the transition between different ecosystems
may be definite due to sudden changes of key bioclimatic factors
(e.g. see Figures 3, 4 and 5). The FAO ecological zones offer a
partition of the European continent in categories whose sharp
boundary may be associated with geographic sudden transitions
(e.g. the boundary between mountain systems and nonmountainous areas within a certain ecological domain, see Figure
4). However, the spatial distribution of biomes may frequently be
“interspersed rather than sharply delineated”31 . Some of the sharp
boundaries between FAO ecozones, such as those between the
boreal coniferous forest and the temperate continental forest, or
that between the temperate continental forest and the temperate
steppe, are located in areas which in reality show a smooth
transition of bioclimatic and geographic features. In these cases,
the necessarily sudden transition between two different classes
- with a relatively homogeneous set of average characteristics
for all the areas within the first class and an abrupt, relatively
homogeneous change of average characteristics when crossing
the boundary into the second class - may not find an ecological
correspondence in the actual smoother transition of ecosystems.
Box 1: Other classification systems
biotemperature (mean of temperatures between freezing and
30 °C) and aridity (potential evaporation ratio to mean total annual
precipitation) as its main axes for classification44-47. The system was
originally designed and is most appropriate for tropical and subtropical
areas, although it can also be used globally.
25
62
0.1
.5
The FAO classification of ecological zones is not the only one in use.
A number of other systems use bioclimatic variables to distinguish
between broad categories1-9 , with similar although not identical
results. Another classical, simple categorization family is based
on the Holdridge Life Zones (Figure 7). This classification was first
described in 1947 and uses annual precipitation (logarithmic scale),
0
5
12
5
0.2
25
0.5
altitudinal belts
0
50
ion
ira
t
desert
10
00
po
rain
tundra
00
rain
forest
subalpine
steppe
moist
forest
wet
forest
rain
forest
6°C
montane
superarid
perarid
arid
semiarid
subhumid
humid
perhumid
0
lower montane
00
32
tropical
thorn
moist
wet
rain
dry
desert
desert
steppe/
forest
forest
forest
forest
scrub
woodland
thorn
moist
wet
rain
very dry
dry
desert
desert
woodland
forest
forest
forest
forest
forest
scrub
premontane
16
subtropical
16
warm temperate
80
00
desert
desert
scrub
40
po
8
cool temperate
3°C
00
wet
forest
20
tia
ten
moist
forest
alpine
)
boreal
dry
scrub
desert
1.5°C
mm
4
le
va
wet
tundra
superhumid
humidity provinces
Fig. 7: Holdridge Life Zones
12°C
24°C
biotemperature
moist
tundra
n(
tio
dry
tundra
alvar
ita
subpolar
2
tra
nsp
desert
cip
pre
desert
al
1
polar
nu
an
rat
io
latitudinal regions
Fig. 8: Romania, Transsylvania, Fanatele Oroiului. A meadow-steppe
grasslands in the temperate continental ecological zone. This landscape
shows components of temperate steppe and the beginning of some
characteristics from the temperate mountain system.
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This is an extended summary of the chapter. The full version of
this chapter (revised and peer-reviewed) will be published online at
https://w3id.org/mtv/FISE-Comm/v01/e01e873. The purpose of this
summary is to provide an accessible dissemination of the related
main topics.
This QR code points to the full online version, where the most
updated content may be freely accessed.
Please, cite as:
de Rigo, D., Houston Durrant, T., Caudullo, G., Barredo, J. I., 2016.
European forests: an ecological overview. In: San-Miguel-Ayanz,
J., de Rigo, D., Caudullo, G., Houston Durrant, T., Mauri, A. (Eds.),
European Atlas of Forest Tree Species. Publ. Off. EU, Luxembourg,
pp. e01e873+
Introduction | European Atlas of Forest Tree Species
31