Professional Documents
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FOREST TREE
SPECIES
EUROPEAN ATLAS OF
FOREST TREE
SPECIES
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San-Miguel-Ayanz, J., de Rigo, D., Caudullo, G., Houston Durrant,
T., Mauri, A. (Eds.), 2016. European Atlas of Forest Tree calling 00 800 6 7 8 9 10 11
Species. Publication Office of the European Union, Luxembourg. (freephone number from anywhere in the EU) (*).
(*) The information given is free, as are most calls (though some operators, phone boxes or hotels
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Preambles 2
Publication details 2
Contributors 2
Introduction 5
Preface on the European Atlas of Forest Tree Species 6
The European Union Forest Strategy and the Forest Information System for Europe 7
Forest resources in Europe: an integrated perspective on ecosystem services, disturbances and threats 8
Forest bio-based economy in Europe 20
European forests: an ecological overview 24
European forest classifications 32
European Forest Types: tree species matrix 34
Past forests of Europe 36
Tree species 40
The European Atlas of Forest Tree Species: modelling, data and information on forest tree species 40
How to read the Atlas 46
Abies alba Silver fir 48 Picea omorika Serbian spruce 117
Abies spp. Circum-Mediterranean firs 50 Picea sitchensis Sitka Spruce 118
Acer campestre Field maple 52 Pinus cembra Arolla pine 120
Acer platanoides Norway maple 54 Pinus halepensis and Pinus brutia Aleppo pine and Turkish pine 122
Acer pseudoplatanus Sycamore or sycamore maple 56 Pinus mugo Dwarf mountain pine 124
Aesculus hippocastanum European horse-chestnut 60 Pinus nigra Black pine 126
Ailanthus altissima Tree of heaven 61 Pinus pinaster Maritime pine 128
Alnus cordata Italian alder 62 Pinus pinea Stone pine 130
Alnus glutinosa Common or black alder 64 Pinus sylvestris Scots pine 132
Alnus incana Grey alder 66 Populus alba White poplar 134
Alnus viridis Green alder 68 Populus nigra Black poplar 136
Betula sp. Birches 70 Populus tremula Eurasian aspen 138
Carpinus betulus Common hornbeam 74 Prunus avium Wild cherry 140
Carpinus orientalis Oriental hornbeam 76 Prunus cerasifera Cherry plum 142
Castanea sativa Sweet chestnut 78 Prunus mahaleb Mahaleb cherry 143
Celtis australis Nettle tree 80 Prunus padus Bird cherry 144
Chamaecyparis lawsoniana Lawson cypress 81 Prunus spinosa Blackthorn 145
Cornus mas Cornelian cherry 82 Pseudotsuga menziesii Douglas fir 146
Cornus sanguinea Common or red dogwood 84 Quercus cerris Turkey oak 148
Corylus avellana Common or European hazel 86 Quercus frainetto Hungarian oak 150
Cupressus sempervirens Mediterranean cypress 88 Quercus ilex Holm or evergreen oak 152
Eucalyptus globulus Tasmanian blue gum 90 Quercus palustris Pin oak 154
Euonymus europaeus Spindle tree 92 Quercus pubescens Downy or pubescent oak 156
Fagus sylvatica European beech 94 Quercus pyrenaica Pyrenean oak 158
Frangula alnus Alder buckthorn 96 Quercus robur and Quercus petraea Pedunculate oak and sessile oak 160
Fraxinus angustifolia Narrow-leaved ash 97 Quercus suber Cork oak 164
Fraxinus excelsior Common ash 98 Robinia pseudoacacia Black locust 166
Fraxinus ornus Manna ash 100 Salix alba White willow 168
Ilex aquifolium European holly 102 Salix caprea Goat willow 170
Juglans regia Common walnut 103 Sambucus nigra Black elderberry 172
Juniperus communis Common juniper 104 Sorbus aria Common whitebeam 174
Juniperus oxycedrus Prickly juniper 105 Sorbus aucuparia Rowan or Mountain ash 176
Juniperus phoenicea Phoenician juniper 106 Sorbus domestica Service tree 178
Juniperus thurifera Spanish juniper 107 Sorbus torminalis Wild service tree 180
Larix decidua European larch 108 Tamarix spp. Tamarisks 182
Olea europaea Olive 111 Taxus baccata European or English yew 183
Ostrya carpinifolia European hop-hornbeam 112 Tilia spp. Limes (Linden) 184
Picea abies Norway spruce 114 Ulmus spp. Elms 186
Appendices 194
The European Commission 194
Information on copyright, citation and disclaimer 196
Preface
Dear readers,
Happy reading!
Fig. 4. Top: The role of wood and other solid biofuel among the renewable sources for the primary production of energy. Data for the EU-28,
from 1990 to 2013. (Source: Eurostat2) e
Bottom: An indicator of biodiversity such as the common bird index is a composite multispecies statistic and is considered by Eurostat as
a headline indicator on the status of natural resources in the European Union31-33 . Overall, the index declined between 1990 and 2013.
However, while the common farmland bird index experiences a noticeable decrease (which has mainly been associated with agricultural
changes33), the index associated with common forest birds shows a recovery with a positive trend between 2000 and 2013. (Source: Eurostat33)
Fig. 8: The ability of forest tree species to tolerate periodic flooding and soil-water saturation varies from taxon to taxon. Different trees are
better suited to cope with different intensity of soil moisture, flooding and waterlogging. Among other factors, this also depends on the flooding
tolerance of the rooting system, while anthropic factors (e.g. agriculture, pasture, management of forests or water bodies, ...) may alter the
natural stratification of taxa with respect to the frequency of waterlogging154-156 . Within forested mountain valleys of hills, some areas may be
subject to more intense run-off (gullies, shallow secondary drainages or valleys). There, a sustainable, healthy forest cover requires the presence
of suitable tree species. Riparian forest buffers also provide remarkable services and require flooding-tolerant tree species38, 157, 158 .
Left: the side of a secondary valley (Italy, Trentino Alto Adige) clearly shows the stratification of coniferous trees (upper part) and more flooding-
tolerant broadleaved trees/shrubs (lower part).
Right: qualitative visualisation of the typical dendritic network along which flow accumulation due to run-off is more intense (gullies).
Correspondingly, erosion rates and thus forest soil carbon losses may be concentrated in relatively few critical areas. The presence of flooding-
tolerant vegetation may mitigate erosive phenomena and protect the carbon accumulated in the forest soil. The partial coverage by debris and
sediments with high permeability (due to periodic intense runoff and sediment transport) may sometimes require these strategic tree species
also to be drought tolerant20, 159, 160 .
(Adapted from an image authored by F. Delventhal, CC-BY, http://archive.is/TNMT8)
b c
Fig. 10: Source: Daniele de Rigo, et al., http://dx.doi.org/10.6084/m9.figshare.2247472 (doi: 10.6084/m9.figshare.2247472). In Europe, the larger
proportion of forest carbon pools is constituted by forest soils (54.1%)1 . Continent-wide, the boreal, temperate and subtropical
mountain systems host a variety of forest ecosystems44, 161, 162 . At the same time, European mountainous areas are among the
one more susceptible to potential soil erosion by water163 . Forests and other vegetation may provide a very effective protection
to mitigate soil and carbon loss and can support the provision of some key ecosystem services concerning mass stabilisation
and buffering/attenuation of mass flow such as sediments, rockfall, shallow landslides117, 131, 164, 165 . The extent of this protection
is also related to particular species composition of forests166 . In particular, some forested areas in hills and mountains may be
subject to a multiplicity of disturbances, with a limited number of tree species suitable to thrive under the combined effect of
these stressors154, 155, 159, 160, 167. This box qualitatively illustrates the network of valleys, hollows, gullies and riparian areas where
higher rates of potential erosion coexist with higher frequency of waterlogging or flooding. A healthy, sustainable forest cover in
these critical areas is able to display a multifunctional mitigation and protection.
a b
Fig. 12: Austria, a flood meadow. Areas subject to frequent intense run-off
and even occasional waterlogging tend to host flooding-tolerant tree
c d
species and other resistant vegetation.
(Adapted from an image authored by Stanislav Doronenko, CC-BY, http://archive.is/EwMjv)
e f
Fig. 14: Biodiversity of forest ecosystems may also be appreciated for the rich variety of animals living in European forests and woodlands, which include
birds (see Figures 4-bottom and 5) and mammals, along with other key vertebrates and invertebrates. Although it is impossible to summarise even a
simplistic overview of these animals in a picture, a few well known mammals are here illustrated in their forest environment.
A: Eurasian wolf (Canis lupus lupus), Bavarian Forest National Park, Germany. Copyright MrT HK, CC-BY, http://archive.is/4abQB
Fig. 13: Forest tree species may serve to mitigate and control desertification. B: Eurasian lynx (Lynx lynx), Bavarian Forest National Park, Germany. Copyright MrT HK, CC-BY, https://archive.is/SBD3v
Forested sand dunes in Fr, Gotland, Sweden. C: Red deer (Cervus elaphus), United Kingdom. Copyright Dave Hamster, CC-BY, https://archive.is/zrInn
(Adapted from an original image authored by Bernt Rostad, CC-BY, https://archive.is/oMGnD )
D: European beaver, (Castor fiber). Copyright NTNU, Faculty of Natural Sciences and Technology, CC-BY, https://archive.is/CIbc5
E: Red fox, (Vulpes vulpes), United Kingdom. Copyright Neil McIntosh, CC-BY, https://archive.is/GpQUM
F: Wolverine (Gulo gulo). Copyright NH53, CC-BY, https://archive.is/qUpx0
Nutrition (biomass and Provisioning Regulation and Mediation of waste, toxics and
potable water) Maintenance other nuisances
Forest
ecosystem
Simplified overview of the main
services forest ecosystem services.
The schema is based on the categori-
sations proposed by the Common
International Classification of
Cultural section: it covers all the Ecosystem Services (CICES) and the
non-material, and normally non-con- Mapping and Assessment of Ecosys-
sumptive, outputs of forest ecosys- tems and their Services (MAES). The
tems that affect physical and mental complete classification defines a
states of people. hierarchy of four levels. Here, a
simplified visualisation is proposed.
Non-material interactions with Some aspects of ecosystem services
ecosystems and land-/ may reverberate in multiple catego-
seascapes (coastal forests): ries with complex feedbacks (e.g.
surface / ground water; hydrological
cycle / water flow, filtration and
Physical and intellectual chemical conditions of freshwaters).
Fig. 20: Within forests, erosion may be unnoticed by non-experts due to the
leaf litter and the geological irregularities of the terrain in mountainous areas
(although litter dams may ease the identification of erosive phenomena).
Top: minor gullies converging in a beech forest (Fagus sylvatica), Italy. It may
be observed that beech trees are absent where the intensity of run-off causes
high soil moisture or even occasional waterlogging. In this particular example,
no flooding-tolerant species appears to occupy the niche left empty by beech.
In these cases, the erosion of forest soil might progress in the gullies with a
weakened protective effect by vegetation.
(Adapted from an image authored by Francesco Ciabatti, CC-BY, http://archive.is/rhon5)
Bottom: removal of trees within forest gullies may intensify erosion and
slope instability. Occasionally, intense precipitation and subsequent run-
off/waterlogging may remove as sediments the forest topsoil, negatively
affecting or even preventing the establishment of a new forest cover.
Fig. 19: Rills and incipient gullies of erosion within a forest of broadleaved trees, where the understorey has been removed close to a
(Adapted from an image authored by Nociveglia, CC-BY, http://archive.is/wdJQi)
cattle crossing. Weakened protection is offered during wintertime, with a subsequent unsustainable imbalance between soil formation
and soil erosion rates. Similar circumstances may reduce the provision of soil protection ecosystem services by a forested area.
(Adapted from an image authored by J. Kelley, CC-BY, http://archive.is/dPvzB)
Fig. 21: The European Forest Fire Information System (EFFIS) was established jointly by the European
Commission (EC) services (DG ENV and JRC) and the relevant fire services in the EU Member States and the
Forest and Civil Protection services other countries222 as the EC focal point of information on forest fires.
The Rapid Damage Assessment module of EFFIS was set up to provide reliable and harmonised estimates
of the areas affected by forest fires during the fire season. The methodology and the spatial resolution of
the satellite sensor data used for this purpose can map all fires of about 40 hectares or larger. Although
fires smaller this are not mapped, the analysis of historical fire data has determined that the area burned
by wildfires of at least 40ha accounts for about 75% of the total area burnt every year in the Southern EU.
The figure shows the total cumulated burnt areas mapped by EFFIS from 2000 to 2015 across the entire
region covered. As expected, the regions with the most fires are in a band across southern Europe, and
the five most affected countries (Portugal, Spain, Mediterranean part of France, Italy and Greece) account
for around 85% of the total burnt area each year222 . However, almost all countries have been affected,
at least in some years, by large fires of more than 40ha. The northern regions such as UK, Ireland and
Scandinavia are not usually as affected the southern regions, but in dry years, especially in the early parts
of the season before the new green vegetation has started to sprout, large wildfires can occur.
The northern half of Portugal and parts of northern Spain are historically the The other most affected parts of Europe are concentrated around the Mediterranean region.
most affected regions with a significant proportion of the annual total burnt Particularly affected are the large islands (Corsica, Sardinia and Sicily), Greece and Croatia. Like
area recorded here. In these regions, the main fire season occurs in summer. the other most affected countries, the main fire season for these countries is in summer.
Top right: Significant wind throw damage after a storm in Loch Bharcasaig, Scotland.
(Copyright Andrew, CC-BY, https://archive.is/gWjss)
Bottom right: Storm Gudrun struck Denmark and Sweden in January 2005. The
damage resulted in the creation of the worlds largest wood stockpile.
(Copyright Granssons keri AB i FRILA: AP)
In several European countries, the Nordic common rights (right of public Chestnuts are a very important non-wood forest product of forests in In 1993-2013, the European average production of chestnuts has been
access to the wilderness, or freedom to roam) grant access for picking Mediterranean countries. The high market price of chestnuts and the low 130000 tonnes per year, i.e. more than 10% of the global production18 .
non-wood forest products such as berries. Top: Chestnut bur.
Top: Blueberry picking. level of inputs required in the chestnut systems have led to the recent (Copyright William Warby, CC-BY, https://archive.is/0qaSn)
(Copyright Ragnar Jonsson, CC-BY) expansion of chestnut agro-forestry systems in some Mediterranean regions. Bottom: Chestnuts.
Bottom: Understorey of Vaccinium myrtillus in a coniferous forest. (Copyright Maja Dumat, CC-BY, https://archive.is/05xeX)
(Copyright Yuri Timofeyev, CC-BY, https://archive.is/D6fqw) This process is favoured by the movement of people from the countryside
Variety of mushrooms found in the forests of Priekui, Latvia. Raspberries. Truffle hunter with his dog, Tuscany, Italy. Italian white
(Copyright Inga Vitola, CC-BY, https://archive.is/5L6W2) (Copyright Maja Dumat, CC-BY, https://archive.is/wqkTZ ) truffles are highly sought after and have high value.
(Copyright Michela Simoncini, CC-BY, https://archive.is/P2xPu)
and Polar. Four of these domains can be found in Europe zone and are snow covered for large parts of the year. Typical Bottom: Temperate oceanic ecological zone. The Saar river influences part
(Tropical is absent). vegetation is pine forest. of northeastern France and western Germany.
(Copyright Wolfgang Staudt, CC-BY, http://archive.is/2XBaO)
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.
Introduction
Boundaries of the Ecological Zones approximately coincide
with Kppen-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 10C.
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 000mm 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 400-900mm. 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
Fig. 2: FAO ecological zones for forest reporting (2010).
parts of the Mediterranean Basin and vegetation is dominated
by shrubs adapted to arid environments.
Polar Temperate mountain system Subtropical mountain system
Subtropical Mountain systems can be found in the southern
Boreal tundra woodland Temperate steppe Subtropical steppe
mountain regions of Europe and the Middle East. The vegetation may
be quite varied depending on the altitude, exposure and humidity. 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.
Polar domain
In the Polar domain there are months with average
temperatures below 10C 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)
Boreal Tundra
Boreal Mountain
Temperate Mountain
Temperate Steppe
Temperate Oceanic
Subtropical Mountain
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)
Fig. 6: Norway, Femundsmarka National Park at the transition between the ecological zone of the boreal
Bottom: Wider scale view of the Alpine transition between the temperate mountain system and that of the boreal coniferous forests.
mountain system and the surrounding plains hosting fragmented temperate
oceanic forests. Top: Compared to the sometimes sudden transitions highlighted in Fig. 4, in this region the component of
(Adapted from an image authored by Francisco Antunes, CC-BY, http://archive.is/vy5Fa) 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)
Boreal tundra zone: shrub tundra vegetation in northern Kola Peninsula (Murmansk, Temperate oceanic zone: mixed broadleaved forest along Ystwyth river (Ceredigion,
Russia). 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 Subtropical mountain zone: mediterranean mixed forest on mountain area of the
(Sogn, Norway). 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 Hundtjrnen lake near Floda (Dalarna, Subtropical steppe zone: steppe dry grassland on Djurdjura Massif of the Tell Atlas
central Sweden). 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 Subtropical dry forest zone: sclerophyllous evergreen vegetation on the coasts of the
Forest (Freiburg, Germany). Fethiye Gulf (Mula, south-western Turkey)
(Copyright ilovebutter, CC-BY, https://archive.is/mNK71) (Copyright Jorge Franganillo, CC-BY, https://archive.is/ONm91)
Fig. 8: Romania, Transsylvania, Fanatele Oroiului. A meadow-steppe
grasslands in the temperate continental ecological zone. This landscape
Temperate steppe zone: steppe grasslands near Poltava (Poltava Oblast, Ukraine). Subtropical humid forest zone: mixed broadleaved and coniferous forest on the side of shows components of temperate steppe and the beginning of some
(Copyright Vlad Butsky, CC-BY, http://archive.is/qf8qf) Pontic Mountains near Trabzon (Trebisonda, Turkey). characteristics from the temperate mountain system.
(Copyright Aleksasfi, PD, https://archive.is/W8Ffz)
(Copyright Alexandru Badarau, AP, https://archive.is/mQfqE )
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[13] Y.Pan, R.A. Birdsey, O.L. Phillips, R.B.
results. Another classical, simple categorization family is based originally designed and is most appropriate for tropical and subtropical Jackson, Annual Review of Ecology, P.Panagos, Natural Hazards and Earth
Evolution, and Systematics 44, 593 System Science 15, 225 (2015).
on the Holdridge Life Zones (Figure 7). This classification was first areas, although it can also be used globally. (2013). [38] C.Bosco, D.deRigo, T.A. Dijkstra,
described in 1947 and uses annual precipitation (logarithmic scale), [14] M.Lindner, etal., Impacts of climate G.Sander, J.Wasowski, IFIP Advances
change on European forests and options in Information and Communication
for adaptation, European Commission Technology 413, 321 (2013). ISSN:1868-
Directorate-General for Agriculture and 4238. Special issue: Environmental
Rural Development, Brussels (2008). Software Systems. Fostering sharing
0.1
information.
.5
62
176 (2014).
5
25
an
[19] M.G. Barbour, etal., North American published in Environmental Modelling &
0
[20] X.Yue, L.J. Mickley, J.A. Logan, J.O. [42] V.Amici, etal., Ecological Complexity 21,
rec
nsp
1.5C
767 (2013).
10
biotemperature
Social and Behavioral Sciences 120, 788 Earthzine 5, 384031+ (2012). 2nd quarter
on
va
3C (2014).
le
20
(m
[22] F.Evrendilek, S.Berberoglu, O.Gulbeyaz, [44] L.R. Holdridge, J.A. Tosi, Life Zone Ecology
4
[23] P.S.A. Beck, S.J. Goetz, Environmental [45] W.R. Emanuel, H.H. Shugart, M.P.
00
40
[24] J.I. Barredo, J.San-Miguel-Ayanz, [46] A.E. Lugo, S.L. Brown, R.Dodson,
8
12C Commission. EUR - Scientific and Technical [47] T.Yue, etal., Ecological Modelling 144,
80
16
warm temperate thorn dry moist wet rain lower montane Research 25730, 16 pp (2012). 153 (2001).
desert desert steppe/ [25] A.Langner, F.Achard, G.Grassi,
forest forest forest forest
subtropical scrub woodland premontane Environmental Research Letters 9,
0
00
desert thorn very dry dry moist wet rain 24C 124008+ (2014).
32
16
super- perarid arid semi- sub- humid per- super- This is an extended summary of the chapter. The full version of
arid arid humid humid humid 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
humidity provinces main topics.
This QR code points to the full online version, where the most
Fig. 7: Holdridge Life Zones 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+
Phytosociology The sector of phytosociology which deals with vegetation system (EUNIS), available at http://eunis.eea.europa.eu.
dynamics and aims at detecting successional series is known This database provides information about European habitat
Phytosociology is a rather young discipline which started
as symphytosociology. The vegetation series (or sigmetum) classification, data sheets on species, habitats and designed
in Europe in the early decades of the last century with the
is a group of spatially and/or temporally interconnected protected sites compiled in the framework of Natura 20008 ,
Swiss botanist and ecologist Josias Braun-Blanquet1 . This
vegetation units that may co-occur in different succession and species mentioned in relevant international conventions
discipline is focused on describing plant communities through
stages or steps within the same place. Progressive succession and in the IUCN Red Lists. The EUNIS habitat classification
a multi-purpose approach, taking into consideration different
is the natural dynamic process from pioneer to mature and is a hierarchical classification of the terrestrial, freshwater
parameters such as species composition, frequency, cover,
stable communities (the so-called climax or head series), and marine habitats for the whole of Europe9 . Up to now this
structure (tree, shrub, herb, moss layers), spatial distribution
while regressive succession is a disruptive process from classification provides a pan-European reference set of units
(the so-called sociability; i.e. causal, clumped, etc.). The aim
more complex communities to open and less developed plant for meeting requirements in policy objectives and in supporting
of phytosociology is to provide effective synthetic information
assemblages; the latter mostly issues from intense and/or applications that relate to biodiversity monitoring and reporting.
about plant communities in order to assign them to different
frequent anthropogenic disturbance. A crosswalk from the EUNIS habitats at level 3 to the European
and recognisable units called syntaxa. Syntaxa are then
In the last century a very large body of phytosociological phytosociological syntaxa and vice-versa is also available10 .
grouped hierarchically within a classification system which
literature has been published, and a variety of schools with
is ruled by the ICPN (International Code of Phytosociological European Forest Types
different approaches formed, especially in southern and eastern
Nomenclature)2 . Phytosociologists usually collect data through
Europe, while this approach found no or little consensus in the The European Forest Types (EFTs) scheme has been
vegetation relevs recording the species occurrences within
United Kingdom and in north European countries. Recently the developed by an international consortium of experts with the aim
selected plot areas using a semi-quantitative cover-abundance
European Vegetation Survey, a working group established in to create a user-friendly classification system. It is, in fact, able
scale. Collected data are then analysed looking for similarities
1992, joined European phytosociologists in order to develop to facilitate understanding, interpretation and communication of
and dissimilarities in order to detect distinct vegetation types.
common standards, organize scientific meetings and survey data on indicators describing the status and trends of forests,
More recently several numerical models have been developed,
programmes, and to produce shared protocols and publications5, and forest management in Europe. The EFTs is a hierarchical
which help to identify dominant and diagnostic species, to 6
(http://euroveg.org). In the first overview of vegetation units, classification consisting of 14 categories, including 78 forest
evaluate species-richness and species-evenness, and which can
80 classes, 233 orders and 928 alliances have been detected types11-13 . The 14 categories represent groups of ecologically
lead to a more objective classification of vegetation units3 .
all over Europe7. distinct forest communities dominated by specific assemblages
The hierarchical classification foresees, as the botanical
of trees, including introduced tree species, while the types
one, different ranks. The association is the basic vegetation EUNIS Habitat Classification correspond to a finer level of division of the category in terms
unit: it represents a plant community defined by a particular
The European Topic Centre on Biological Diversity (ETC/BC), an of tree species composition. The EFTs is, therefore, a flexible
and uniform floristic composition and habitat structure, where
international consortium working with the European Environment system to compare forest information on ecologically similar
a relative constancy or abundance of characteristic species is
Agency (EEA), developed the European nature information forests, unlike other classification systems that present an
recognisable (called also diagnostic or dominant), which can
describe the community and its ecology. The upper units are
a group of lower ones, which share one or more diagnostic RANK SUFFIX EXAMPLE DESCRIPTION
and dominant species4 . Each unit is assigned a scientific name Class -etea Quercetea ilicis All the evergreen woody plant communities of the Mediterranean basin.
and is defined with compound names formed by one or two
Order -etalia Quercetalia ilicis All the Mediterranean forests dominated by evergreen broadleaved trees.
scientific names of the dominant and diagnostic plant species
with a different suffix for each rank2 . Alliance -ion Quercion ilicis All the Mediterranean forests dominated by holm oak (Quercus ilex).
Association -etum Aceri campestris- Mixed wood dominated by holm oak (Quercus ilex) and several deciduous
Quercetum ilicis broadleaved species typical of the North-Western Sicilian calcareous
mountains in the meso- and supra-Mediterranean bioclimatic belts.
Subalpine larch-arolla pine forest near Morgex (Valle dAosta, North-West Italy).
(Copyright Giovanni Caudullo: CC-BY)
Spruce-birch boreal forest in Norra Kvill National Park (Kalmar County, South Sweden).
(Copyright Tracy Houston Durrant: CC-BY)
References
[1] J.Braun-Blanquet, Pflanzensoziologie: habitat classification and related data
Grundzge der Vegetationskunde sets, Tech. rep., Alterra, Wageningen, NL
(Springer-Verlag, Berlin, 1964). (2014).
[2] H.E. Weber, J.Moravec, J.P. Theurillat, [11] A.Barbati, P.Corona, M.Marchetti, Plant
Journal of Vegetation Science 11, 739 Biosystems - An International Journal
(2000). Dealing with all Aspects of Plant Biology
[3] L. Mucina, E. van der Maarel, Vegetatio 141, 93 (2007).
81, 1 (1989). [12] A.Barbati, P.Corona, M.Marchetti,
[4] M.D. Jennings, D.Faber-Langendoen, O.L. European forest types: Categories and
Loucks, R.K. Peet, D.Roberts, Ecological types for sustainable forest management
Monographs 79, 173 (2009). reporting and policy (European
Environment Agency, Copenhagen, 2007),
[5] L.Mucina, etal., Lazaroa 30, 267 (2009). second edn.
[6] M.Chytr, etal., Applied Vegetation [13] A.Barbati, M.Marchetti, G.Chirici,
Science 19, 173 (2015). P.Corona, Forest Ecology and
[7] J.S. Rodwell, etal., The diversity of Management 321, 145 (2014).
European vegetation - An overview of [14] T.HoustonDurrant, J.San-Miguel-Ayanz,
phytosociological alliances and their E.Schulte, A.SuarezMeyer, Evaluation
relationships to EUNIS habitats (National of BioSoil Demonstration Project: Forest
Reference Centre for Agriculture, Nature biodiversity - Analysis of biodiversity
and Fisheries, Wageningen, 2002). module, vol. 24777 of EUR - Scientific and
[8] Council of the European Union, Official Technical Research (Publications Office of
Journal of the European Union 35, 7 the European Union, 2011).
(1992). [15] R.McRoberts, etal., National Forest
[9] C.E. Davies, D.Moss, M.O. Hill, EUNIS Inventories: Contributions to Forest
habitat classification - revised, Tech. rep., Biodiversity Assessments, G.Chirici,
European Environment Agency, European S.Winter, R.E. McRoberts, eds. (Springer
Topic Centre on Nature Protection and Netherlands, 2011), vol.20 of Managing
Biodiversity, Paris (2004). Forest Ecosystems, pp. 4197.
[10] J.H.J. Schamine, etal., Development of
vegetation syntaxa crosswalks to EUNIS
CONIFERS
CONIFERS BROADLEAVES
BROADLEAVES
Pinus peucesempervirens
sempervirens
CATEGORY
CATEGORY FOREST
FOREST
TYPE
TYPE
communis
communis
Abies nordmanniana
nordmanniana
Juniperus phoenicea
phoenicea
articulata
articulata
Juniperus oxycedrus
oxycedrus
Juniperus thurifera
thurifera
Populus canescens
canescens
Carpinus orientalis
orientalis
Ostrya carpinifolia
Ostrya carpinifolia
Castaneacoccifera
coccifera
Juglanspubescens
Betula pubescens
Pinus canariensis
canariensis
Cedrus brevifolia
brevifolia
atrocinerea
atrocinerea
Abies borisii-regi
borisii-regi
betulus
betulus
Pinus halepensis
halepensis
avellana
avellana
Populus tremula
Populus tremula
Pinus heldreichii
Pinus heldreichii
Fagus moesiaca
Fagus moesiaca
sativa
sativa
orientalis
Fagus orientalis
Pinus sylvestris
sylvestris
Fagus sylvatica
sylvatica
Alnus orientalis
Alnus orientalis
Alnus glutinosa
glutinosa
pendula
pendula
Salix eleagnos
Salix eleagnos
Quercus suber
Taxus baccata
baccata
pinaster
Pinus pinaster
Picea omorika
omorika
Alnus cordata
cordata
nigra
nigra
Pinus cembra
cembra
regia
regia
Larix decidua
Larix decidua
Juniperus sp.
Juniperus sp.
Cedrus libani
libani
alba
alba
Alnus incana
incana
Salix cinerea
Salix cinerea
Abies others
others
Taxusfragilis
Salix fragilis
Quercus ilex
ilex
Salix caprea
caprea
Betulaviridis
viridis
peuce
Populusfaya
Juglansfaya
Pinus mugo
mugo
Pinus pinea
pinea
Abies abies
abies
nigra
nigra
Abies alba
alba
Cupressus
Cupressus
Tetraclinis
Tetraclinis
alba
alba
Juniperus
Juniperus
Juniperus
Juniperus
Juniperus
Castanea
Carpinus
Carpinus
Carpinus
Quercus
Quercus
Quercus
Populus
Populus
Populus
Populus
Corylus
Corylus
Cedrus
Cedrus
Myrica
Myrica
Betula
Betula
Fagus
Fagus
Alnus
Alnus
Alnus
Alnus
Alnus
Abies
Abies
Abies
Pinus
Pinus
Pinus
Pinus
Pinus
Pinus
Pinus
Pinus
Pinus
Pinus
Picea
Picea
Picea
Salix
Salix
Salix
Salix
Salix
Salix
1. Boreal
1. Boreal
forestforest 1.1 Spruce
1.1 Spruce
and spruce-birch
and spruce-birch
borealboreal
forestforest
1.2 Pine
1.2 and
Pinepine-birch
and pine-birch
borealboreal
forestforest
2. Hemiboreal
2. Hemiboreal
forestforest
and nemoral
and nemoral2.1 Hemiboreal
2.1 Hemiboreal
forestforest
coniferous
coniferous
and mixed
and mixed
broadleaved-
broadleaved-
2.2 Nemoral
2.2 Nemoral
ScotsScots
pine forest
pine forest
coniferous
coniferous
forestforest
2.3 Nemoral
2.3 Nemoral
sprucespruce
forestforest
2.4 Nemoral
2.4 Nemoral
BlackBlack
pine forest
pine forest
2.5 Mixed
2.5 Mixed
ScotsScots
pine-birch
pine-birch
forestforest
2.6 Mixed
2.6 Mixed
ScotsScots
pine-pedunculate
pine-pedunculate
oak forest
oak forest
2.7 Atlantic
2.7 Atlantic
maritime
maritime
pine forest
pine forest
2.8 Nemoral
2.8 Nemoral
silversilver
fir fir
3. Alpine
3. Alpine
coniferous
coniferous
forestforest 3.1 Subalpine
3.1 Subalpine
larch-arolla
larch-arolla
pine and
pinedwarf
and dwarf
pine forest
pine forest
3.2 Subalpine
3.2 Subalpine
and mountainous
and mountainous
sprucespruce
and mountainous
and mountainous
mixedmixed
spruce-silver
spruce-silver
fir forest
fir forest
3.3 Alpine
3.3 Alpine
ScotsScots
pine and
pineBlack
and Black
pine forest
pine forest
3.4 Mountainous
3.4 Mountainous
birch birch
forestforest
4. Acidophilous
4. Acidophilous
oak and
oakoak-birch
and oak-birch4.1 Acidophilous
4.1 Acidophilous
oakwood
oakwood
forestforest 4.2 Oak-birch
4.2 Oak-birch
forestforest
5. Mesophytic
5. Mesophytic
deciduous
deciduous
forestforest 5.1 Pedunculate
5.1 Pedunculate
oakhornbeam
oakhornbeam
forestforest
5.2 Sessile
5.2 Sessile
oakhornbeam
oakhornbeam
forestforest
5.3.1 5.3.1
Ashwood
Ashwood
and oak-ash
and oak-ash
forestforest
lowlands
lowlands
5.3.2 5.3.2
Ashwood
Ashwood
and oak-ash
and oak-ash
forestforest
uplands
uplands
5.4 Maple-oak
5.4 Maple-oak
forestforest
5.5 Lime-oak
5.5 Lime-oak
forestforest
5.6 Maple-lime
5.6 Maple-lime
forestforest
5.7 Lime
5.7 Lime
forestforest
5.8 Ravine
5.8 Ravine
and slope
and slope
forestforest
5.9 Other
5.9 Other
mesophytic
mesophytic
deciduous
deciduous
forests
forests
6. Beech
6. Beech
forestforest 6.1 Lowland
6.1 Lowland
beechbeech
forestforest
of southern
of southern
Scandinavia
Scandinavia
and north
and north
central
central
Europe
Europe
6.2 Atlantic
6.2 Atlantic
and subatlantic
and subatlantic
lowland
lowland
beechbeech
forestforest
6.3 Subatlantic
6.3 Subatlantic
submountainous
submountainous
beechbeech
forestforest
6.4 Central
6.4 Central
European
European
submountainous
submountainous
beechbeech
forestforest
6.5 Carpathian
6.5 Carpathian
submountainous
submountainous
beechbeech
forestforest
6.6 Illyrian
6.6 Illyrian
submountainous
submountainous
beechbeech
forestforest
6.7 Moesian
6.7 Moesian
submountainous
submountainous
beechbeech
forestforest
7. Mountainous
7. Mountainous
beechbeech
forestforest 7.1 South
7.1 South
western
western
European
European
mountainous
mountainous
beechbeech
forestforest
7.2 Central
7.2 Central
European
European
mountainous
mountainous
beechbeech
forestforest
7.3 Apennine-Corsican
7.3 Apennine-Corsican
mountainous
mountainous
beechbeech
forestforest
7.4 Illyrian
7.4 Illyrian
mountainous
mountainous
beechbeech
forestforest
7.5 Carpathian
7.5 Carpathian
mountainous
mountainous
beechbeech
forestforest
7.6 Moesian
7.6 Moesian
mountainous
mountainous
beechbeech
forestforest
7.7 Crimean
7.7 Crimean
mountainous
mountainous
beechbeech
forestforest
7.8 Oriental
7.8 Oriental
beechbeech
and hornbeam-oriental
and hornbeam-oriental
beechbeech
forestforest
8. Thermophilous
8. Thermophilous
deciduous
deciduous
forestforest
8.1.1 8.1.1
DownyDowny
oak forest
oak forest
- western
- western
8.1.2 8.1.2
DownyDowny
oak forest
oak forest
- Italian
- Italian
8.1.3 8.1.3
DownyDowny
oak forest
oak forest
- Greek,
- Greek,
Anatolian
Anatolian
8.1.4 8.1.4
DownyDowny
oak forest
oak forest
- steppe
- steppe
8.2 Turkey
8.2 Turkey
oak, Hungarian
oak, Hungarian
oak and
oakSessile
and Sessile
oak forest
oak forest
8.3 Pyrenean
8.3 Pyrenean
oak forest
oak forest
8.4 Portuguese
8.4 Portuguese
oak and
oakMirbecks
and Mirbecks
oak Iberian
oak Iberian
forestforest
8.5 Macedonian
8.5 Macedonian
oak forest
oak forest
8.6 Valonia
8.6 Valonia
oak forest
oak forest
8.7 Chestnut
8.7 Chestnut
forestforest
8.8.1 8.8.1
Thermophilous
Thermophilous
ash forest
ash forest
8.8.2 8.8.2
Fraxinus
Fraxinus
ornusornus
and Ostrya
and Ostrya
carpinifolia
carpinifolia
forestforest
8.8.3 8.8.3
Hop-hornbeam
Hop-hornbeam
(Ostrya
(Ostrya
carpinifolia)
carpinifolia)
forestforest
8.8.4 8.8.4
Oriental
Oriental
hornbeam
hornbeam
(Carpinus
(Carpinus
orientalis)
orientalis)
forestforest
8.8.5 8.8.5
Thermophilous
Thermophilous
maplemaple
(Acer (Acer
spp.) forest
spp.) forest
8.8.6 8.8.6
Mediterranean
Mediterranean
lime (Tilia
lime (Tilia
spp.) forest
spp.) forest
8.8.7 8.8.7
CeltisCeltis
australis
australis
forestforest
8.8.8 8.8.8
HorseHorse
chestnut
chestnut
and walnut
and walnut
mixedmixed
woodswoods
9. Broadleaved
9. Broadleaved
evergreen
evergreen
forestforest 9.1 Mediterranean
9.1 Mediterranean
evergreen
evergreen
oak forest
oak forest
9.2 Olive-carob
9.2 Olive-carob
forestforest
9.3 Palm
9.3 Palm
groves
groves
9.4 Macaronesian
9.4 Macaronesian
laurisilva
laurisilva
9.5 Other
9.5 Other
sclerophlyllous
sclerophlyllous
forests
forests
10. Coniferous
10. Coniferous
forests
forests
of theof the 10.1.110.1.1
Mediterranean
Mediterranean
pine forest
pine forest
- Pinus
- Pinus
pinaster
pinaster
Mediterranean,
Mediterranean,
Anatolian
Anatolian
and and 10.1.210.1.2
Mediterranean
Mediterranean
pine forest
pine forest
- Pinus
- Pinus
halepensis
halepensis
Macaronesian
Macaronesian
regions
regions
10.1.310.1.3
Mediterranean
Mediterranean
pine forest
pine forest
- Pinus
- Pinus
pineapinea
10.2.110.2.1
Mediterranean
Mediterranean
BlackBlack
pine forest
pine forest
10.2.210.2.2
Anatolian
Anatolian
BlackBlack
pine forest
pine forest
10.3 10.3
Canarian
Canarian
pine forest
pine forest
10.4 10.4
Mediterranean
Mediterranean
and Anatolian
and Anatolian
ScotsScots
pine forest
pine forest
10.5 10.5
Alti-Mediterranean
Alti-Mediterranean
pine forest
pine forest
10.6 10.6
Mediterranean
Mediterranean
and Anatolian
and Anatolian
fir forest
fir forest
10.7 10.7
Juniper
Juniper
forestforest
10.8 10.8
Cypress
Cypress
forestforest
10.9 10.9
CedarCedar
forestforest
10.1010.10
Tetraclinis
Tetraclinis
articulata
articulata
standsstands
10.1110.11
Mediterranean
Mediterranean
yew stands
yew stands
11. Mire
11. and
Mireswamp
and swamp
forestforest 11.1 11.1
Spruce
Spruce
mire forest
mire forest
11.2 11.2
Pine mire
Pine forest
mire forest
11.3 11.3
Alder Alder
swamp swamp
forestforest
11.4 11.4
Birch Birch
swamp swamp
forestforest
11.5 11.5
Pedunculate
Pedunculate
oak swamp
oak swamp
forestforest
11.6 11.6
AspenAspen
swampswamp
forestforest
12. Floodplain
12. Floodplain
forestforest 12.1 12.1
Riparian
Riparian
forestforest
12.2 12.2
FluvialFluvial
forestforest
12.3 12.3
Mediterranean
Mediterranean
and Macaronesian
and Macaronesian
riparian
riparian
forestforest
13. Non
13. riverine
Non riverine
alder,alder,
birch,birch,
or or 13.1 13.1
Alder Alder
forestforest
aspenaspen
forestforest 13.2 13.2
ItalianItalian
alder alder
forestforest
13.3 13.3
Birch Birch
forestforest
13.4 13.4
AspenAspen
forestforest
14. Introduced
14. Introduced
tree species
tree species
forestforest
In this table the European Forest Types (EFTs) scheme is of the main tree and shrub species of forest interest in Europe in the EFT; the species presence in the EFT is either secondary
presented as proposed and revised by Barbati and colleagues1-3, has been evaluated, separated into three main groups: conifers, or predominant but in peculiar and not characteristic ecological
counting 14 broad categories which include 78 forest types, some broadleaved and alien trees. The species presence in the EFTs is conditions of the EFT; the presence in the EFT is both dominant
of which are divided into sub-types. For every EFT the presence categorized in three classes: the species is abundant and dominant and secondary in some cases. It has to be taken into account that
JRC_EFDAC_forest_atlas.indd 35
Quercus
Quercus pubescens
pyrenaica
Quercus
Quercus faginea
pubescens
Quercus
Quercus ithaburensis
faginea
Quercus
Quercus macrolepis
ithaburensis
Ulmus
Quercusminor
macrolepis
grabra
Ulmus minor
Ulmus
Ulmus
Ulmus laevis
grabra
Celtis australis
Ulmus laevis
Species is present but not dominant
Species is present in either category
Aesculus ippocastanus
Celtis australis
Species is dominant in that forest type
Laurus nobilis
Aesculus ippocastanus
Platanus orientalis
Laurus nobilis
Pyrus pyraster
Platanus orientalis
Pyrus
Pyrus communis
pyraster
Malus sylvestris
Pyrus communis
Malus
Malus domestica
sylvestris
domestica
Sorbusdomestica
Malus
Sorbus
Sorbus aucuparia
domestica
torminalis
Sorbus aucuparia
Sorbus
Sorbus
Rhamnus alaternus
References
Buxus sempervirens
frangula
Rhamnus alaternus
Rhamnus
Tilia tomentosa
Rhamnus frangula
Tilia
Tilia cordata
tomentosa
Tilia
Tilia platyphyllos
cordata
Tamarix sp.
Biosystems - An International Journal
Tilia platyphyllos
Cornus
Tamarixsanguinea
sp.
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Cornus sanguinea
Cornus
Erica
Cornusarborea
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scoparia
Erica arborea
Erica
Arbutus unedo
Erica scoparia
ornus
Fraxinusunedo
Arbutus
Fraxinus
Fraxinus excelsior
ornus
Fraxinus
Fraxinus angustifolia
excelsior
Olea europaea
Fraxinus angustifolia
Phillyrea latifolia
Olea europaea
Sambucus nigra
Chamaecyparis lawsoniana
Pseudotsuga menziesii
European forest types: Categories and
Tsuga heterophylla
Agency, Copenhagen, 2007), second edn.
Chamaecyparis lawsoniana
types for sustainable forest management
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reporting and policy (European Environment
Picea
Tsugasitchensis
heterophylla
ALIENS
Larix kaempferi
This QR code points to the full online version, where the most updated content may be freely
Pinus
Pinus radiata
strobus
This is an extended summary of the chapter. The full version of this chapter (revised and peer-
Populus hyb.
Pinus radiata
In: San-Miguel-Ayanz, J., de Rigo, D., Caudullo, G., Houston Durrant, T., Mauri, A. (Eds.), European Atlas
Quercus rubra
Populus hyb.
Pividori, M., Giannetti, F., Barbati, A., Chirici, G., 2016. European Forest Types: tree species matrix.
reviewed) will be published online at https://w3id.org/mtv/FISE-Comm/v01/e01e1b6. The purpose of
Quercus
Quercus palustris
rubra
Robinia
Quercuspseudoacacia
palustris
Ailanthus
Robinia altissima
pseudoacacia
Eucalyptus sp.
Ailanthus altissima
P.Corona, Forest Ecology and
Prunus serotina
Eucalyptus sp.
[3] A.Barbati, M.Marchetti, G.Chirici,
Acer negundo
Prunus serotina
35
Acer negundo
08/04/2016 11:08
Past forests of Europe
H. J. B. Birks, W. Tinner
European forests have varied in their composition, structure,
and extent over the last 5 million years or more in response
to global climate changes. European forests have also
undergone very major changes due to the alternating
glacial-interglacial cycles of the Quaternary (last 2.6
million years). European forests have greatly changed in
their extent and structure in the last 5000 years due to
human activities (the Homo sapiens phase) in the current
Holocene interglacial in which we live. Contemporary
ecologists and foresters can learn from lessons from the
past about forest responses and resilience to environmental
changes in the past.
Introduction
Were European forests 500, 5000, 15000, 150000, 1.5
million, 2.5 million, and 5 million years ago similar in species
composition, structure, and extent to the forests of Europe today?
As we cannot directly observe the forests of the past, to answer
these questions we need to reconstruct past forests indirectly
using the fossil record. This involves the study of seeds, fruits,
leaves, wood, and charcoal (macrofossils)1 and of microscopic
pollen grains, spores, cells (e.g. stomata), and charred particles
(microfossils) preserved in lake, bog, alluvial, and other sediments
where organic material can be preserved2 . Pollen analysis as
a tool for vegetation reconstruction - invented in 1916 by the
Swedish geologist Lennart von Post - was and still is the dominant
technique in the Quaternary period, especially the last 15000 years
of the late-Quaternary. Von Post had the idea of expressing fossil
pollen assemblages as percentages of the sum of pollen grains
counted, and of presenting these percentages as stratigraphical
pollen diagrams with pollen assemblages plotted against their
stratigraphical position through the sediment sequence (Fig. 1). He
showed strong similarities in pollen diagrams from a small area,
and striking differences between different areas. He was thus able
to provide the dimension of time (vegetations fourth dimension) to
the study of past vegetation and forests2, 3 .
Pollen analysis
There are ten basic principles of pollen analysis1 (see Box 1).
The results of a pollen analysis are most commonly presented
as a pollen diagram, showing how the percentages of different
pollen types vary with depth, and hence age, in the sedimentary
sequence (Fig. 1). When many sequences have been studied, their
pollen data can be mapped for a particular time interval (e.g. 5000
years ago) to produce so-called isopollen maps for particular
pollen types where the contours represent different pollen values
(e.g. 2.5%, 5%, 10%) (Fig. 2)4 . Alternatively when interest is
centred on the directions and rates of tree spreading, so-called
isochrone maps can be constructed where the contours represent
ages established by radiocarbon dating (e.g. 5000, 6000, 7000
years ago). When the value of a particular pollen type exceeds a
certain threshold value it can be interpreted as reflecting the first
expansion of that taxon at different sites (Fig. 3)5 . The first arrival
of a taxon is more difficult to assess, because the absence of
pollen or macrofossils may not mean a true absence of the taxon
in the landscape. Interpretation of pollen-stratigraphical data in a
qualitative manner in terms of major past vegetational changes is
relatively straightforward2 . Quantitative interpretation of such data
Fig. 2: Isopollen maps of Quercus (oak) pollen percentages across Europe for 12000, 10000, 8000,
in terms of quantitative estimates of past plant abundances is less 6000, 4000, and 2000 radiocarbon years before present (BP). Note the progressive northward spread into
straightforward because of the differential production, dispersal, southern Scandinavia by 6000 BP and the subsequent contraction at 2000 BP in Norway. The percentage
contours are percentages of total tree and shrub pollen.
and hence representation of different pollen types. Approaches for (Modified from Huntley and Birks4)
quantitative interpretation are currently an area of active research
within Europe and elsewhere (e.g.6, 7).
Fig. 1: Summary pollen diagram from Loch Cill an Aonghais (Argyll), a small
lake in south-west Scotland covering the last 12000 radiocarbon years. The
horizontal lines represent partitions of the pollen stratigraphy into pollen-
assemblage zones. The vertical axis is radiocarbon (14C) years before present
(BP) based on eight radiocarbon dates. The small arrows by the Betula (birch),
Quercus (oak), Alnus (alder), and Corylus/Myrica (hazel/bog myrtle) indicate
when these trees or shrubs are inferred to have first expanded near this site.
Cryocratic taxa are coloured red and stippled. These taxa become abundant
again in the open conditions of the Homo sapiens phase where they are
shown in plain red. Protocratic trees are coloured blue, mesocratic trees are
green, oligocratic and telocratic taxa are orange, and taxa associated with
human activity and the Homo sapiens phase of the Holocene are shown in
red. All the pollen and spore percentages are expressed as percentages of the
total number of terrestrial pollen and spores counted (generally 500-600 per
sample). Pollen analyses by Sylvia M. Peglar.
r
u
cre
cold-stress tolerant and adversity traits17.
Fe
rich assemblages of herbs, shrubs, and trees (birch, pine, willow) xim
as
Within these three broad groups of protocratic, mesocratic,
ing
Ma
unleached fertile soils
Incr
and oligocratic and telocratic plants, the actual floristic and
soil
M or
Decre
Dec
forest composition varies from interglacial to interglacial in north- Mesocratic:
Mesocratic Oligocratic
fertility, P limitation
e fe rtile s olis, in cr
e a sin g te m p e
reasin
western and central Europe17. Factors such as location of refugia mid interglacial stage
asing temp
temperate deciduous forests
In cre a sin g
in the cryocratic phase, rates of spreading, distances over which
g biomass
spread occurred, competition, predation, genotypic variation, and fertile brown-earth soils
Protocratic Telocratic
chance as it affects survival, dispersal, and establishment may Oligocratic & Telocratic:
all have contributed to the observed differences in interglacial late interglacial stage
bio
forest patterns17. Similar cycles occurred in southern Europe, yet open conifer (spruce, pine), ericaceous heaths, bogs Cryocratic
ea
e
m
r
rat
s
as
infertile, humus-rich podsols and peats
sin
a
with substantial differences in comparison to central and north-
tur
g
ss
ur
western Europe10, 11, 22 . Due to warmer conditions, European tree &
N
L ow bio m a
e
P
e
species persisted locally, although strongly reduced, in the steppe- Unique to the Holocene ls
like environments of the glacial stages. This corresponds to the Base- e s oi
Homo sapiens: ric h in fertil
cryocratic phase in central and northern Europe. At the onset of an mid-late Holocene (6000 years ago-present)
interglacial, corresponding to the protocratic phase in central and forest clearance, agriculture G la cial
north-western Europe, temperate taxa (e.g. deciduous Quercus, range of soil types, often fertilised (Modified from Birks and Birks18)
Ulmus, Ostrya hop-hornbeam, Carpinus) form open forests together
with evergreen broad-leaved trees (e.g. Quercus ilex holm oak, Olea
europaea olive) and mediterranean shrubs (e.g. Pistacia pistachio),
mixed Fagus-Ilex holly-Quercus forests developed whereas in other cleared or cultivated areas, relaxation in grazing pressure, or
while boreal and steppe vegetation declines (e.g. Betula, Juniperus,
areas there was a rapid change from Tilia- or Quercus-dominance reduction in fire frequency17.
Artemisia wormwood, Chenopodiaceae goosefoot)11, 22, 23 . In the
to Fagus-dominance17. These changes commonly occurred after an The westward, northward, and southward spread and expansion
following phase during the mid-interglacial, corresponding to the
extensive phase of human activity involving clearance and grazing of Picea abies through Finland, Sweden, and Norway over the last
mesocratic phase in central and north-western Europe, warm-
followed by the abandonment of cleared and cultivated areas. This 6000-7000 years4, 31 may be a contemporaneous response to subtle
temperate and Mediterranean conifers (e.g. Abies, Pinus) expand
abandonment may have occurred as a result of local population step-wise climate change, a delayed migration unrelated to simple
into the broad-leaved deciduous and broad-leaved evergreen
collapse following, for example, climate change, emigration, or climate change, a response to forest disturbance creating gaps
forests and arboreal cover increases, probably in response to
over-exploitation of environmental resources30 . for colonisation, or a combination of these factors32 . Whatever its
rising moisture availability. Towards the end of the interglacials,
Other types of secondary woodland developed in areas beyond causes, the invasion of Picea into northern and central Fennoscandia
corresponding to the oligocratic phase in north-western and central
the natural geographical range of Fagus, for example woods of over the last 6000-7000 years resulted in major changes in forest
Europe, moisture-loving taxa such as Fagus, Alnus, and Abies
pure Fraxinus excelsior European ash, Quercus spp., Taxus baccata composition and structure and in soil conditions, with widespread
gradually replace Mediterranean evergreen broad-leaved trees,
English yew, Betula spp., or Ilex aquifolium common holly became accumulation of mor humus, soil leaching, and podsolisation and
while broad-leaved deciduous trees remain important11, 22, 23 .
established on particular soil types following abandonment of changes in the natural fire regime within the boreal forest15, 33.
Finally, forest cover declines and steppe-like environments expand
during the climatic deterioration at the transition from the
interglacial to the next glacial (temperature decreases, reduced Box 3: Palaeo-model comparison: past, present and future Mediterranean vegetation
moisture), corresponding to the telocratic phase. There is an
apparent order within interglacial forest patterns when viewed at Simulation of future vegetation dynamics at Lago di Massaciuccoli, a a b c
the broad-scale of an entire interglacial cycle of 10000-15000 coastal lake in Tuscany (central Italy), with a dynamic vegetation model
years, whereas within each phase of an interglacial (ca. 5000 (LANDCLIM) for different climatic conditions (today vs. warming) and
years) there is often great variation between interglacials, hence levels of disturbance (low vs. moderate). The mid- to late-Holocene
the ability of pollen stratigraphy to differentiate between many of sedimentary pollen record of Lago di Massaciuccoli is used to validate
the different interglacials17. the model, in particular LANDCLIM is able to simulate extinct vegetation d e
types which were growing in the past at the site before anthropogenic
Europes forests in the Holocene (11700 disturbance became excessive.
years ago-today) a) Present-day (1950-2000 AD) mean monthly temperature (1
The mesocratic phase in the Holocene interglacial stage was standard deviation) and average total monthly precipitation at Lago di
greatly modified about 5000-6000 years ago by the onset of Massaciuccoli close to Pisa (Tuscany).
forest clearance and prehistoric shifting cultivation and livestock f
b) Map of Italy and Switzerland with Lago di Massaciuccoli denoted by a black
farming (Fig. 1). This new phase, unique to the Holocene is called
star, red star shows position of Gorgo Basso in southern Sicily (Fig 4).
the Homo sapiens phase (see Box 2)17. There was a steep fall in
Ulmus pollen values (Fig. 1), probably a result of an interaction c) Future (2071-2100 AD) mean monthly temperature and precipitation
between prehistoric human activities and a tree pathogen, with projected by a regional climate model (SMHI) for Lago di Massaciuccoli.
elm pollen values halving within 5 years at a site in southern
d) and e) Vegetation simulated at Lago di Massaciuccoli with LANDCLIM,
England24 . Similarly, 5000-6000 years ago Abies disappeared
a dynamic vegetation model with d) present climate and future climate Figure from Henne et al.52
from the Mediterranean and sub-Mediterranean lowlands of the
e). All vegetation models were initialised with the same present-day
Italian Peninsula, probably in response to excessive Neolithic
climate scenario and moderate disturbance before 2010.
disturbance by fire and by browsing25, 26 . As with Ulmus in England,
Abies collapses were rapid, with pollen values of Abies halving f) Holocene pollen percentages of upland trees and shrubs at Lago di
within 13 and 22 years at sites in Italy27 and Italian Switzerland28 , Massaciuccoli.
respectively.
Simulations of todays vegetation under low disturbance shows
In some areas of central and north-west Europe, forest clearance
Abies alba co-dominance with Quercus ilex (see right image) in the
and subsequent dereliction of clearings may have facilitated local
Mediterranean forest. This vegetation type disappeared during the late
colonisation and expansion of new immigrants such as Fagus
Holocene most likely in response to excessive anthropogenic burning
sylvatica European beech, Picea abies Norway spruce, and possibly
and land use25 . In agreement, simulations show the disappearance of
Carpinus betulus European hornbeam4 . While the establishment of
this vegetation type under current climate with moderate land use.
Fagus sylvatica during Mesolithic times followed climate change
Future climate and vegetation conditions at Lago di Massaciuccoli are
(cooling and a moisture increase) in southern and southern-central
comparable to present climate and vegetation conditions at Gorgo
Europe29, it is possible that the rapid spread of Fagus across
Basso, southern Sicily (Fig 5). With low land use, evergreen oak forest
central Europe in the last 4000-5000 years4 may have only been
will prevail16 , while under moderate land use forests will be reduced and
facilitated by the creation of abundant, large clearings within Tilia- Spontaneous regeneration of Abies alba and Quercus ilex
maquis (low biomass) will expand. in a cryptic Mediterranean stand in lowland Tuscany.
or Quercus-dominated forests on well-drained soils. In some areas (Copyright Willy Tinner: CC-BY)
tree species distribution and habitat suitability for the current Bottom left: France, Stratification over the years of forest and land
situation5-11 . However, even this apparently simpler objective management. Even the uniformity of tree species within managed forest
stands may be subject to border effects with increased diversity of species.
displays an impressive set of challenges for it to be addressed at (Adapted from an image authored by Hans Fransen, CC-BY, http://archive.is/fqFff)
the continental scale.
Top right: Romania. Grazing and managed forest stands generate patches
of locally uniform vegetation. However, at the landscape scale this results in
Heterogeneity of forests and forest data at a high heterogeneity.
(Adapted from an image authored by Sergey Norin, CC-BY, http://archive.is/JYzDI)
the European scale
Europe is densely populated, with about 3 % of world
land hosting almost 7% of the world population (estimates
for 2014)12 . As a consequence, the anthropogenic pressure on subtropical, temperate and boreal mountain systems, where
forest ecosystems is elevated, with intense landscape diversity highly variable bio-climatic conditions influence local forest
and relatively few undisturbed areas of high wilderness and ecosystems and their composition17, 20-22 (see also the chapters
low accessibility13, 14 (see Figure 2). Forest management may changes species. In both cases, it might be that the available field European forests: an ecological overview and Forest resources
significantly alter the local composition of forests compared observations are not dense enough to offer a statistical sampling in Europe: an integrated perspective on ecosystem services,
with the potential natural vegetation15, 16 , for example because without wide uncertainties on the real detailed composition and disturbances and threats).
management practices might prevent or mitigate the natural local distribution of tree species (see Figure 1). Europe spans These characteristics of the continent contribute to define
interspecific competition by other tree taxa. Therefore, while over the subtropical, temperate, boreal up to the polar climate its overall high heterogeneity. Continental-scale modelling of tree
undisturbed areas may display a rich variety of species adapted to domain, experiencing highly a wide range of climate patterns17-19 . species distribution and habitat suitability needs to adapt to this
coexist in the same mature ecosystem, areas with predominantly For example, mountains characterise more than one third of the exceptional challenge with a harmonisation effort to integrate
managed forests may exhibit very diverse patterns with sudden European land (see Figure 2) with peculiarities associated to the the different sources of forest-based field observations, also
considering how to better take advantage from the already
available land cover mapping. This Atlas is based on possibly
the richest set of information harmonised at the European scale
and focusing on recorded occurrences of forest tree species.
Several hundred thousand harmonised field observations have
been collected and integrated to cover several millions of
square kilometres. However, the local density of available field
observations (plot density, see Figure 6) is uneven with extensive
areas of the continent very poorly covered. In the mountainous
areas or where land use and landscape diversity is wide, a much
denser network of field observations would have been required
to reliably reconstruct at the km2 resolution the local distribution
of forest tree species (not only including the ones that occupy
Fig. 2: A qualitative visual overview for some of the dimensions of complexity and
heterogeneity in the European continent.
Top left: A view of the European continent at night shows clearly the large centres of population,
where anthropogenic influences might be greatest. From NASA, Earth Observatory40 .
Top right: Accessibility may be defined as the travel time to a location of interest using
land or water based travel, estimated using a cost-distance algorithm which computes
the cost of travelling between two locations, and usually measured in units of time. The
values in the map represent the cost required to travel across them (hence this is often
termed a friction-surface). This shows the varying levels of human influence, darker areas
- i.e. less accessible ones - more prevalent in the far north of the continent and along
mountain ranges. From European Commission and the World Bank14 . The similarity with
the top left view is evident.
Bottom left: The peculiar administrative heterogeneity of Europe (28 member states in the
European Union with 24 official languages and several states which are either federations,
federacies or in any case providing large autonomy to internal administrative units) further
increases the complexity and intrinsic uncertainty of continental-scale environmental
modelling. This is because the many regional datasets are often autonomously collected
and organised, with different spatial density of sampling, accuracy and uneven definitions
of measured/estimated quantities for different taxa. For example, information gaps between
multiple administrative units and institutions may potentially lead to classify regional field
observations on a given species (missing from the local inventory) as false negatives.
Bottom right: orographic complexity. About one third of the continent is covered by mountain
systems (according to a recent revised classification based on the UNEP-World Conservation
Monitoring Centre approach20) and in most massifs the forest cover is a key component up
to the timberline. Mountain forests are exposed to heterogeneous bio-climatic conditions:
temperature, solar irradiation, precipitation patterns may vary greatly depending on the local
elevation, slope, aspect, and resulting solar and rain shadow. Therefore, field observations
of forest tree species in these areas may be associated with information limited to very
local conditions.
categories - such as broadleaved and coniferous trees - instead the first time a systematic coverage of forest tree taxa distribution Right: Italy, Marche region. Another mixed land cover. Although fragmented,
here forests are more homogeneous with weaker evidences of border
of specific forest tree species36-39 . and habitat suitability at the continental scale, based on the most effects.
The available European-wide data and information have been comprehensively integrated set of available data and information. (Adapted from an image authored by Francesco Gasparetti, CC-BY, https://archive.is/PRbcg)
collected and harmonised within the Forest Information System for The modelling strategy here summarised is designed to be inclusive
Box 1: The review model and content-processing chain of the European Atlas of Forest Tree Species
The making of the European Atlas of Forest Tree Species required several Concerning the modelled maps, chorology areas may integrate several These few statistics refer to this printed version of the Atlas. As highlighted
years of work for the editorial board and intense exchange with more than sources within a single coarse-resolution overview. Each distribution (relative by Figure 4, the extended summaries of this book are associated with their
50 international experts who devoted their efforts in co-authoring, revising, probability of presence) and suitability (maximum habitat suitability) map corresponding updated online full version. The review model for the Online
reviewing, providing additional information, insights and comments for the is the result of an ensemble of hundreds of intermediate maps generated European Atlas of Forest Tree Species is similar to that applied to the printed
chapters of the Atlas. Here, a brief overview on the Atlas review model and via statistical resampling to ensure that the final estimate is more robust edition. In addition, each updated manuscript will undergo a more extensive
on its content-processing chain is summarised. and able to tolerate a larger amount of outlier data or data affected by peer review. If accepted, a chapter will be persistently published in the FISE
high uncertainties a feature which is essential at the considered spatial portal. Periodic updates might be possible for e.g. integrating recent or
The modelling, data and information on forest tree species, as harmonised
scale (see section Heterogeneity of forests and forest data at the European extended literature, improved data and modelling, extended statistics and
within FISE, has been complemented by open contributions from international
scale in this chapter). Overall, this required about 20000 core intermediate iconography, with the potential contribution of additional co-authors. All
authors, advisors and reviewers, under a clearly defined taxonomy of roles42
geospatial layers (at 1km2 spatial resolution) to be processed, without previous peer-reviewed versions of an updated chapter will remain accessible.
to ease the collaboration and with the systematic internal recording of all
considering other ancillary layers.
contributions and modifications by means of an internal version control
system. After the initial design and implementation of core materials and
methods (data, models, maps, diagrams, bibliography and text), a cycle of
internal review with the support of external reviewers and subsequent
content revision has been iterated up to finalise the extended summary for
each chapter (see Figure 4). To obtain the printed version of this Atlas, the
extended summaries with their cycles of review and revision required the
support of novel computational tools, with the design and implementation of
a dedicated chain of data-transformation modules (D-TM)43, 46 .
For a given chapter version, the content-processing chain of D-TMs starts
from text and references, annotated by the co-authors by using common
word-processing formats (DOC, DOCX). Although easy to edit with images,
tables and other supporting information, these formats are unsuitable for an
automated semantic enhancement of their content. The manual harmonisation
of the bibliography was based on the records stored in the Meta-information
Database on Integrated Natural Resources Modelling and Management
(INRMM-MiD, http://mfkp.org/INRMM ; about INRMM see also the chapter
Forest resources in Europe: an integrated perspective on ecosystem services,
disturbances and threats)43. The Atlas provides an overall bibliography with
more than 2400 cited references to scientific and technical publications, which
correspond to over 1600 unique references. For each of them, an INRMM-MiD
public record is available with integrated metadata and meta-information on
the cited publication. Furthermore, the INRMM-MiD catalogue covers about
5000 indexed publications, of which more than 2400 are on forest resources,
which may serve for further readings.
From the human-editable format, the D-TM chain for each chapter version
generates intermediate information with semantic enhancements, to derive
HTML, LaTeX, PDF and RTF documents with a harmonised, machine-readable
semantic structure. The content-processing chain is implemented on a GNU/
Linux computing environment44, 45 under the semantic array programming
paradigm43, 46 . Overall, the making of the printed version of the Atlas required
the processing and generation of more than 18000 files (considering only the
textual information content) organised in more than 2400 units of content.
Fig. 4: The review model of the European Atlas of Forest Tree Species.
Fig. 5: Examples of sparsely forested areas with mixed land cover. Although
the presence of tree species characterises these examples, some tree
formations may not fulfil the definition of forest and thus may be classified
differently in the available land cover maps. Furthermore, trees in non-
forest areas may be supported by scarcer field observations, since the main
effort of systematic data collections such as national forest inventories
is more focused on forest areas. However, remarkable biotic disturbances
(such as some forest pests) may spread also over landscapes with sparse
but susceptible trees. Therefore, even approximate information able to
systematically cover these European areas may be essential.
Bottom: France. Linear formations contribute to support connectivity among Fig. 6: Plot density, computed with a spatial grid of 50km2, (LAEA) for the datasets used: European Forest Inventories (EF),
forest patches. BioSoil (BS), Forest Focus (FF), EUFGIS (EG), Genetic Diversity (GD). Among other constraints, the array-based semantics of
(Adapted from an image authored by Fredi Bach, CC-BY, https://archive.is/9jRGz) each harmonised dataset expects the corresponding geospatial records to have nonnegative values (::nonnegative::), after the
removal of outliers, highly uncertain or missing data (::nanless::), all considered as not available information.
Fig. 9: Overly simplistic models may be affected by high modelling uncertainty because too many non-negligible
components of the real system are not taken into account by the model. However, with growing modelling
complexity, even the complexity of the underpinning software code increases. Software engineering techniques
and good computational science practices may help to mitigate the rise of software uncertainty. Since they
cannot completely prevent software uncertainty, a trade-off exists between increased complexity of models
(assuming that this increase is associated with a better approximation of the reality, which for some categories
of models may be structurally impossible77, 78) and the resulting increased software complexity.
Source: Daniele de Rigo, CC-BY, https://dx.doi.org/10.6084/m9.figshare
Fig. 10: Qualitative visualisation of the potential cumulated effect due to the combined uncertainty in modelling,
software and data. Integrated modelling frequently exploits multiple heterogeneous data sources by combining
specific data-processing and intermediate derived data as generated from specialised software modules. The
uncertainty of each data-transformation is propagated up to the final combined output, whose cumulated
uncertainty depends even on the initial uncertainty of the input data. More complex models might be associated
with a higher sensitivity of their estimates to outliers and input uncertainty, sometime with higher prediction
errors for new data77, 78 . This increase of the combined data uncertainty may be mitigated with robust modelling
techniques (e.g. statistical resampling and robust statistics) which may also contribute to mitigate software
uncertainty79-84 .
Source: Daniele de Rigo, CC-BY, https://dx.doi.org/10.6084/m9.figshare
Fig. 11: Qualitative visualisation of varying trade-offs for different available spatial resolution of input data and
information. A qualitative ranking is proposed to simplistically illustrate the different complexity associated to
the modelling approaches discussed in this chapter. Coarser spatial resolution of input data may be associated
with an intrinsic partial loss of information (e.g. finer-resolution details within highly heterogeneous areas such
as mountain systems and high-diversity landscapes). This may reverberate in higher final data uncertainty.
Among other aforementioned concepts and criteria, even this qualitative trade-off has been taken into account
for adapting the modelling complexity of the discussed maps and diagrams.
Source: Daniele de Rigo, CC-BY, https://dx.doi.org/10.6084/m9.figshare
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General information
This section provides a brief overview on how to understand
the information provided in the species chapters present in this
Atlas. In each chapter there is an extended summary of the
current state of knowledge about that species, which is aimed
to be written in an easily accessible style but at the same time Fagus sylvatica
scientifically grounded. Therefore, each chapter has been through
a revision by scientific experts and includes a comprehensive list
of scientific references. Although the chapters have been written
by a number of different authors, they have been harmonized to Fagus sylvatica in Europe: distribution, habitat, usage and threats
obtain species information that is as homogeneous as possible T. Houston Durrant, D. de Rigo, G. Caudullo Fag
throughout the Atlas. The full version of each chapter (expanded Fagus sylvatica L., or European beech, is one of the most important and widespread broadleaved trees in Europe. It is a
Fagus
and fully peer-reviewed) will be published in the online version of large deciduous tree that can maintain its high growth rate until late maturity. Its natural range extends from southern
Some
Scandinavia to Sicily, from Spain in the west to northwest Turkey in the east. Though not demanding of soil type,
the Atlas at http://w3id.org/mtv/FISE-Comm/v01/. beech requires a humid atmosphere with precipitation well distributed throughout the year and a well-drained soil. It
them
very s
tolerates rigorous winter cold, but is sensitive to spring frost. Owing to the capacity of its root system for assisting in
Each chapter starts with a summary and description of the circulation of air throughout the soil, and the amount of potash in its leaves, Beech trees conserve the productive
those
oppos
the species to continue with paragraphs concerning the species capacity of the soil better than many other species. Its wood is strong and wears well making it ideal for a wide range
the Ca
of uses, from furniture to musical instruments, as well as for pulp and firewood.
distribution, habitat and ecology, importance and usage and The European beech (Fagus sylvatica L.) is a large deciduous
of the
two18 .
finally threats and diseases. Most chapters deal with a single tree that commonly reaches 30-40 m and is capable of attaining Frequency
the va
heights up to 50 m in some locations1 . In contrast to many other becau
species, although in a few cases the information is presented at
< 25%
25% - 50%
tree species, it is able to maintain a high rate of growth until a 50% - 75%
taxon level (e.g. circum-Mediterranean firs). relatively mature age. The tree is usually single-stemmed with
silver-grey bark. The leaves are typically 10 7 cm, dark and shiny
> 75%
Chorology
Native
A key contribution of this Atlas is the inclusion of innovative green. They have an oval to elliptic shape, with wavy margins and
Large beech in a mountain pasture in Piani di Praglia (Genova, North Italy).
short teeth at the end of the parallel veins on each side2, 3 . Beech is
maps and diagrams concerning: 1) Frequency and Chorology; monoecious: the male and female flowers are borne on the same
(Copyright Ettore Balocchi, www.flickr.com: CC-BY)
2) Modelled Distribution; 3) Maximum Habitat Suitability; 4) branches. It has a typical life span of around 150-300 years, and
reproduces very late (40-50 years old). Fruiting normally occurs
Habitat and Ecology
Beech is a hardy species. It tolerates very shady situations
Autoecology, for all those species for which sufficient data exist. every 5 to 8 years. Its seed production is characterised by irregular
(it is the most shade-tolerant broadleaved tree in its range10),
mast years (when a very heavy crop is produced), usually following
High quality images are also included relative to forest habitat, hot summers of the previous year. The bitter edible nuts are sharply
so that natural regeneration is possible in silvicultural systems
with continuous crown coverage as the seedlings are able to
individual trees or more detailed images concerning the bark, tri-angled and are borne singly or in pairs in soft-spined husks. The
beech nuts are an important source of food for several animals
survive and grow below the canopy of established trees. The
predominance of beech means a reduction of light level in the
leaves, fruits and flowers. and birds including squirrels, woodpigeons, woodpeckers and jays;
understorey vegetation level and in that case beech seeds survive
they also play a major part in seed dispersal by hiding the seeds
better than those of other tree species. It is not particularly soil-
and failing to retrieve all of them1 .
Map 1: Plot distribution and simplified chorology map for Fagus sylvatica. sensitive11 and grows on a wide variety of soils with a pH range protec
Distribution Frequency of Fagus sylvatica occurrences within the field observations as from 3.5 to 8.5, although it cannot tolerate the most acidic and gn
reported by the National Forest Inventories. The chorology of the native spatial
conditions. Beech shows a moderate soil-acidifying ability12 . It deer is
Beech is widespread across Europe: it can be found from Sicily range for F. sylvatica is derived after Meusel and Jger, and EUFORGEN27, 28 .
prefers moderately fertile ground, calcified or lightly acidic and is frosts
in the south to Bergen in southern Norway4-6 . An analysis of pollen
also sensitive to late frosts13; therefore it is found more often on same
records indicate that the species has spread across Europe from orientalis). At the southern part of its range (Spain, Sicily) it is only
Frequency and Chorology small scattered populations left after the last glaciation, and is normally present at altitudes of more than 1 000 m, and can even be
the side of a hill than at the bottom of a clayey basin. It grows
well on soft soils in which the root system can easily penetrate
woolly
Ganod
currently probably at its maximum post-glacial spread7. It needs a found at elevations of up to 2 000 m1, 8 . High summer temperatures,
and its optimal growth is in humid soils situated on calcareous red he
growing season of at least 140 days, and for this reason cannot drought and moisture availability are limiting factors for the
This map summarises two basic pieces of information concerning survive too far north in Scandinavia7. Longitudinally its range is from distribution of beech in Europe, but continentality is also associated
or volcanic parent rocks. On the contrary, it does not thrive on among
sites that are regularly flooded or which have stagnant water,
the species: the Cantabrian Mountains in the west to the Carpathians and Balkan with limiting its spread in north-western regions4 . Climate change
since it needs good drainage and will not tolerate waterlogged
Mountains in the east, although there are some areas in Europe may have impacts on its future distribution, particularly at the
1. The species frequency over a 50km square grid (blue where it is not found as a native tree, such as the Po valley and extremes of its range where it is likely to become less competitive in
or compacted soils1, 14 . Beech furthers soil conservation due to
its production of a large quantity of litter (around 900 g/m2 per
dots): this shows the percentage of plots inside the grid the Hungarian plain. As the climate becomes more continental in the south and east (primarily because of drought), but could expand
year). The root system tends to be shallow, making it susceptible
the eastern parts of Europe it is replaced by oriental beech (Fagus its range into Scandinavia and the Baltic9.
that contain the species of interest. The sampling points to drought when compared to coniferous stands15 . However, there
appears to be some genetic variability across different climatic
are derived from the same datasets used to model the zones, since trees in southern Europe are able to cope better with
species distribution (Map 2) and the maximum habitat drought than those in the north1 .
ending
species selection and silviculture in
Ireland. (National Council for Forest
(Bioversity International, Rome, Italy, 2009).
[24] A. Oprea, C. Srbu, I. Goia, Contributii
to-date scientific literature, which has been revised by three
iceable
Research and Development (COFORD),
2003).
Botanice 46, 17 (2011).
[25] M. ijai Nikoli, J. Milovanovi, M. Noni,
scientific experts. All the references are sequentially included
ourable R. Kneevi, D. Stankovi, Genetika 45,
make
[9] K. Kramer, et al., Forest Ecology and
Management 259, 2213 (2010). The
ecology and silviculture of beech: from
369 (2013). through the text and listed at the end of the chapter. A revised
[26] T. Denk, G. Grimm, K. Stgerer, M. Langer,
es little Mature beech forest with autumn colour foliage in Delamere forest, Cheshire, UK.
gene to landscape.
[10] A. Praciak, et al., The CABI encyclopedia of
V. Hemleben, Plant Systematics and
Evolution 232, 213 (2002).
and extended set of references will be available in the full
(Forestry Commission, www.forestry.gov.uk: Crown Copyright) forest trees (CABI, Oxfordshire, UK, 2013).
[11] L. Walthert, E. Graf Pannatier, E. S. Meier,
[27] H. Meusel, E. J. Jger, Plant Systematics
and Evolution 162, 315 (1989).
online version of each chapter.
Autoecology diagrams based on harmonised Forest Ecology and Management 297, [28] EUFORGEN, Distribution map of beech
Field data in Europe (including absences) Observed presences in Europe field observations from forest plots. 94 (2013). (Fagus sylvatica) (2008).
[12] L. Augusto, J. Ranger, D. Binkley, A. Rothe, www.euforgen.org.
Annals of Forest Science 59, 233 (2002). [29] J. Jalas, J. Suominen, Atlas Florae Europaeae:
[13] L. Paule, Forest Genetics 2, 161 (1995). distribution of vascular plants in Europe Vol. 3
Average temperature of the coldest month (C)
Autoecology Diagrams
In most chapters, autoecology diagrams (also known possible combinations of variables is very large and for spot (one for every plot), while those plots containing
as climate-space diagrams) have been derived for this Atlas we have focussed on three: 1. Annual average the species of interest are coloured blue, thus illustrating
the described species, based on the datasets of field temperature vs Annual precipitation; 2. Potential spring- the specific climate niche of that species, and showing
observations as harmonised within the Forest Information summer solar irradiation vs. Average temperature of how a given species might be constrained by one or
System for Europe (FISE). These observations are the the coldest month; 3. Seasonal variation of monthly more climatic conditions. Grey patches on this page may
same as those used to estimate the coarse-resolution precipitation vs. Sum of precipitation of the driest month. be coloured blue in others where different species are
forest plot distribution presented in Map 1. The local In the online version of the Atlas other combinations of adapted to different conditions.
bioclimatic conditions where a given species is observed variables may also be found. For more details on the data and modelling aspects, see
are obtained by means of a number of high-resolution The overall climate space occupied by each of the field the Atlas chapter modelling, data and information on
bioclimatic and geographic variables. The number of observations on every species is represented by a grey forest tree species on page 40.
rounded dome as the tree grows older2 . The needles are dark green
and glossy on their upper side while the lower side has two silver-
green waxy bands of 6-8 rows of stomata, and can live for up to six
or eight years. Flowers only appear after 30 to 40 years, generally
in April or May, and the buds are red-brown and non-resinous.
The fully developed seeds are mainly dispersed by wind. With
particularly cool and moist habitats this tree can live up to 500-
600 years3, 4 and reach heights above 60m4-7 making it among
the tallest tree species of the genus Abies in Europe. This tree is
also the most heavily browsed of the commercially important tree
species in montane forests of central and southeastern Europe8 .
Distribution
Map 1: Plot distribution and simplified chorology map for Abies alba.
Silver fir is often distributed on relatively high elevated areas Frequency of Abies alba occurrences within the field observations as reported
(500-2000m a.s.l.). It requires relatively high moisture conditions by the National Forest Inventories. The chorology of the native spatial range
for A. alba is derived after several sources29-31 .
throughout the year, with mean yearly precipitation between
700 and 1800mm9 . Its main distribution is concentrated in
Central Europe, on the Suisse plateau and in South and Eastern Concerning its past distribution palaeo evidence suggests
Germany as well as in the Czech Republic and Austria. There are different ice-age refugia of silver fir in northern, central and
conspicuous numbers in the Pyrenees, Southern Alps of Northern southern Italy, the Balkans, the Pyrenees and potentially France,
Young tree near Zwardo village (South Poland).
Italy and Ticino and the Eastern Alps, the Carpathians and Albania. which is in agreement with results obtained using biochemical (Copyright Crusier, commons.wikimedia.org: CC-BY)
It is also found more sporadically in Eastern France, on the Massif and molecular markers10 . During the past decades silver fir
Central, and in the Apennines. Stands of silver fir are present was positively responding to climate warming in Central Europe Habitat and Ecology
in the Dinaric Alps and are continuously connected towards the and adjacent areas, as documented in many tree ring series11 . Silver fir tolerates a wide variety of soil types with different
Rodopi mountains in Bulgaria and Greece, where it naturally However, in Switzerland, silver fir is decreasing as a result of nutrient content and alkalinity conditions except compact and
hybridises with the Greek fir (Abies cephalonica) forming stable animal browsing and replacement by Norway spruce (Picea hydromorphic soils. Deep and moist but not too wet soils are
populations of intermediate forms described as Bulgarian firs abies), a more economically valuable species12 . The future preferred with a pH from acid to neutral. Silver fir shows a
(Abies x borisii-regis)2 . Plantations of silver fir are rare outside its distribution of Silver Fir is subject of a debate. Some studies noticeable soil-acidifying ability15 . Unlike the other European
natural range, possibly because of increased potential for insect suggest a reduction in response to future expected warming13, 14 , and Mediterranean Abies species, it prefers cooler and moister
damage in monocultures1 . while others suggest stable conditions or expansions15, 16 . conditions, favouring summer temperatures ranging from
approximately 14C to 19C9 . The main limiting factors are a
lack of summer heat and adequate moisture during the growing
season, while new seedlings are extremely sensitive to frost
damage. This tree is mostly found mixed with Norway spruce
Uncertain, no-data (Picea abies) or Scots pine (Pinus sylvestris) at the upper tree
Marginal/no presence < 5% limit1, 4, 18 . At lower altitudes it competes with beech (Fagus
Low presence 5% - 10% sylvatica), being the first conifer species to appear among them4 .
It is very shade tolerant and can remain as a seedling bank
Mid-low presence 10% - 30% under the canopy of older dominant trees for decades. It often
Medium presence 30% - 50% invades deciduous forests due to its easy natural regeneration.
Mid-high presence 50% - 70%
Importance and Usage
High presence 70% - 90% Silver fir is considered an important ecological and functional
Very-high presence > 90% balancer of European forests and can serve as a keystone species
for maintaining high biodiversity in forested ecosystems9 . The
wood is non-resinous, light and fine-grained and also easy to
work, which makes it a good material for carpentry and furniture.
During the seventeenth century, its wood was used to produce
ships masts. The essential oils obtained from the leaves were
also used in the past to heal bruises as well as for treating coughs
and colds1 . Along with Norway spruce (Picea abies), silver fir is
also used for paper production. During the 19th century it was
popular as a Christmas tree, although it has lately been replaced
by the cheaper Norway spruce1 .
Uncertain, no-data
Tundra, cold desert
Negligible survivability
Low survivability
Mid-low survivability
Medium survivability
Mid-high survivability
High survivability
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Map 2: High resolution distribution map estimating the relative probability of presence. Five-lobed yellow leaves with smooth margins, displaying autumn colours.
(Copyright Wendy Cutler, www.flickr.com: CC-BY)
References
[1] I. Chybicki, B. Waldon-Rudzionek, K. Meyza [9] A. Praciak, et al., The CABI encyclopedia of
10, 1739 (2014). forest trees (CABI, Oxfordshire, UK, 2013).
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Guidelines for genetic conservation and
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(Bioversity International, 2004). P. Panagos, Natural Hazards and Earth
System Science 15, 225 (2015).
Large field maples with large domed crown in a garden park (Weinsberg, South Germany). [5] P. S. Savill, The silviculture of trees used in
(Copyright Rosenzweig, commons.wikimedia.org: CC-BY) British forestry (CABI, 2013). [12] F. Florineth, H. P. Rauch, H. Staffler,
Proceedings of the International Congress
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Autoecology diagrams based on harmonised the main valuable broadleaved species (2002), vol. 2, pp. 827837.
Field data in Europe (including absences) Observed presences in Europe field observations from forest plots. in the Pyrenean area and neighbouring
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regions (Centre de la Propietat Forestal,
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[7] H. Meusel, E. Jager, S. Rauschert, E. mri10a15+ (2016).
Average temperature of the coldest month (C)
is dense with foliage. The leaves are opposite, simple, 10-15cm > 75%
Chorology
long, very variable in dimension depending on the age and the Native
vigour of the tree. They have five lobes with long and acuminate Introduced
appear from about 25-30 years and are insect pollinated. The
It tends to be located at the base of hills where it receives a
fruit is a double samara, 3-5cm each, greenish-yellow, dispersed
surface runoff and subsurface soil water flow. It also thrives at
by the wind. The two samaras are set at a wide angle approaching
higher elevations with sufficient precipitation8 . It germinates and
180. The bark of young trees is smooth and grey-brown; on older
grows quickly in shade, even under close canopy. When mature,
trees the bark becomes darker and shallowly furrowed with long
it becomes more light-demanding9, 10 . The height increment is
narrow ridges in a network. The wood is dark reddish-brown; the
about 1 m/year in the first 10 years. With its wide crown it tends
grain is straight, with a fine, uniform texture. Many cultivars have
Map 1: Plot distribution and simplified chorology map for Acer platanoides. to shade and suppress other slow-growing competitor species4 .
been selected for their distinctive leaf shape and coloration and Frequency of Acer platanoides occurrences within the field observations as Under optimal conditions Norway maple may live for more than
for crown shape1-5 . reported by the National Forest Inventories. The chorology of the native and
introduced spatial range for A. platanoides is derived after several sources6, 28-31 . 250 years1 . Over Europe it occurs in fresh and humid sites both
Distribution in coniferous and deciduous forests. In natural stands Norway
Norway maple is the most widespread native maple in Europe. maple occurs as a secondary species with low frequency, thus
and south-east Canada. It shares the ecological range of sugar
Its natural distribution ranging from Greece, Balkans, North Italy not forming pure stands but generally found in small groups or as
maple (Acer saccharum), which is taxonomically close. Norway
and Pyrenees to southern Fennoscandia, and toward the East it individual trees. In temperate continental mixed forests it can co-
maple may be found all over the world in towns and villages as
arrives as far as Russia but not over the Ural Mountains. It grows dominate with other broadleaves such as English oak (Quercus
an ornamental and shade tree6-8 .
from sea level up to 1400m in the Alps. In Europe it is not native robur) and small-leaved lime (Tilia cordata)3, 8, 10-13 .
of western France, British Isles, Netherlands and Denmark. The Habitat and Ecology
subspecies Acer platanoides turkestanicum occurs in mountain The Norway maple is a fast-growing tree species, able to
Importance and Usage
The Norway maple has been used extensively as an ornamental,
forests of Turkey, Caucasus and northern Iran on the southern grow well across a wide range of soil conditions, shade, drought
shade and street-side tree because of its attractiveness, colourful
coasts of the Black Sea and Caspian Sea, reaching 2400m and pollution. However, it thrives best in deep, fertile, moist soils,
foliage and large, spreading crown, in combination with its tolerance
in elevation. Norway maple is commonly found throughout which are adequately drained and with a sub-acid pH. Exposure
of urban conditions. Its ability to resprout vigorously after trimming
mainland Europe, even in countries where it is non-native. It and strong calcareous soils are well tolerated4, 5 . It is intolerant
makes it suitable to be used as a live fence4 .
was also introduced in the United States in the 18th century and of low soil nitrogen conditions, high evapo-transpiration or
The Norway maple distribution range overlaps with many areas
now it is naturalised in some areas of central-east United States prolonged drought and it is rare on acidic soils (pH near 4)8 .
The large five-lobed leaves appear after the flowers. This species is frequently planted as an ornamental for its attractive
(Copyright Free Photos, www.flickr.com: CC-BY) autumn colouration. (Copyright Nicholas A. Tonelli, www.flickr.com: CC-BY)
Uncertain, no-data
Tundra, cold desert
Negligible survivability
Low survivability
Mid-low survivability
Medium survivability
Mid-high survivability
High survivability
Map 2: High resolution distribution map estimating the relative probability of presence. Maturing double samars set in a V shape.
(Copyright Wendy Cutler, www.flickr.com: CC-BY)
Uncertain, no-data
Tundra, cold desert
Negligible survivability
Low survivability
Mid-low survivability
Medium survivability
Mid-high survivability
High survivability
Reddish leaves on young seedling with five deep lobes and long stalks.
(Copyright Stefano Zerauschek, www.flickr.com: AP)
Annual average temperature (C) Potential spring-summer solar irradiation (kWh m-2) Seasonal variation of monthly precipitation (dimensionless)
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Description Frequency
European horse-chestnut (Aesculus hippocastanum L.) is the < 25%
25% - 50%
only European native species belonging to the Aesculus genus, 50% - 75%
which counts 13 tree and shrub species living in temperate > 75%
Chorology
deciduous forests1 . It is a large and tall tree growing up to 39m Native
Description
Frequency
Tree of heaven (Ailanthus altissima (Mill.) Swingle) is a short- < 25%
lived, fast-growing tree, reaching a height of around 20m and 25% - 50%
50% - 75%
70cm in diameter. The bark is greyish and slightly rough1 . The > 75%
Map 1: Plot distribution and simplified chorology map for Ailanthus altissima.
Frequency of Ailanthus altissima occurrences within the field observations as
reported by the National Forest Inventories.
References
[1] B. Shah, Arnoldia pp. 2127 (1997). [9] P. L. Burch, S. M. Zedaker, Journal of
[2] F. Clinovschi, Dendrologie (Editura Arboriculture 29, 18 (2003).
Universitatii Suceava, 2005). [10] E. Pan, N. Bussak, Journal of
[3] P. P. Feret, Journal of Arboriculture 11, Environmental Horticulture 4, 1 (1986).
361 (1985). [11] R. E. Landenberger, N. L. Kota, J. B.
[4] I. Kowarik, I. Sumel, Perspectives in Plant McGraw, Plant Ecology 192, 55 (2007).
Ecology, Evolution and Systematics 8, [12] E. Gravano, M. Ferretti, F. Bussotti, P.
207 (2007). Grossoni, Forest Growth Responses
[5] A. W. Sheppard, R. H. Shaw, R. Sforza, to the Pollution Climate of the 21st
Weed Research 46, 93 (2006). Century, L. Sheppard, Cape, eds. (Springer
Netherlands, 1999), pp. 267272.
[6] S. Y. Hu, Arnoldia 39, 29 (1979).
[13] L. Udvardy, Acta Botanica Hungarica 41,
[7] P. Pyek, et al., Handbook of Alien Species 299 (1998).
in Europe (Springer Netherlands, 2009),
vol. 3 of Invading Nature - Springer Series [14] R. M. Heisey, American Journal of Botany
in Invasion Ecology, pp. 4361. 83, 192 (1996).
[8] C. M. Enescu, Journal of Horticulture, [15] M. Ballero, A. Ariu, P. Falagiani, Allergy 58,
Forestry and Biotechnology 18, 66 (2014). 532 (2003).
the crown is pyramidal, compact and dense. The leaves are dark 25% - 50%
50% - 75%
bright green, lighter underneath. They are 5-12cm long, heart- > 75%
Chorology
shaped, with long stalks and persist from April to December. Native
This species is monoecious with male and female catkins in the
same shoot, appearing when 10-12 years old. The male catkins
are in groups of 3-5 and pendulous, cylindrical, 7-10cm long, Dark green heart-shaped leaf with toothed margins.
pale purplish; they turn to gold from February to April when wind (Copyright Franco Caldararo, www.actaplantarum.org: AP)
pollination occurs. The female catkins are ovoid, above the male more frequent on plantations outside the natural area range,
ones, erect, on a 2-5cm stalk, and with red stigmas when mature. even if in general pathogens affecting the Italian alder are of
The catkins appear at the beginning of summer, are dormant limited importance11 . Root system damage is reported by fungi
during winter until the end of February, when flowers are fully Armillaria spp., Phytophtora alni and Cryphonectria parasitica11,
mature and functional. The fruit is ovoid, 3cm long, green then 15-17
. In northern Europe a new species of genus Phytophthora
woody and dark brown when ripe, similar to conifer cones and is affecting foliage and causing bark necrosis, recently also
called pseudo-strobili. The winged seed are small, leaving the spreading in the Mediterranean region11, 18 . Insect pests Cossus
pseudo-strobili from September-October up to a year after. The cossus, Zeuzera pyrina and Saperda scalaris affect the cortical
bark is brown-grey, smooth in young trees, and then blistered with zones of plants in precarious health conditions1, 11 . In natural
fissures. The wood is a light tan to reddish brown, homogeneous, Map 1: Plot distribution and simplified chorology map for Alnus cordata.
ranges actually the main threats endangering Italian alder are the
Frequency of Alnus cordata occurrences within the field observations as
with a fine, even grain and with relatively wide annual rings reported by the National Forest Inventories. The chorology of the native reduction of clear cutting in mixed forests and in protected areas,
resulting from the fast growth1-5 . spatial range for A. cordata is derived after EUFORGEN19 .
heavy and unauthorised grazing in forests, and the isotherm
shift in the Mediterranean regions. The possibly increase in
Distribution
Importance and Usage temperature due to climate change may force alder ecosystems
This species is endemic to the western side of the Apennines
to shift to higher elevations in restricted areas5 .
in southern Italy and mountains in south-central of Corsica and In mountainous areas, Italian alder has been widely used
north-west of Albania, from 800m to 1500m of elevation, for soil protection and wind breaks, as an ancillary species
frequently down to 300-400m with higher rainfall regimes1, 4-10 . associated with walnut (Juglans regia), wild cherry (Prunus
It is considered a relict species of the Tertiary period for its avium), English oak (Quercus robur) and other noble hardwoods
resemblance to eastern alders, in particular to the Caucasian for high-quality timber plantations. In fact it stimulates the
alder (Alnus subcordata)1 . In Italy it has been introduced in growth of associated species thanks to its nitrogen-fixing root
Sardinia, in the northern Apennines and up to the Southern Alps. capacity, and to its nitrogen-rich and easily degradable leaves
Plantations were also established in different European countries which improve the litter1, 5, 10, 11 . In France it has also been used
during the late 20th century (France, Spain, Portugal, England, for biomass production5 . Stands can be managed as coppice with
Netherlands, etc.) and it has been recently introduced in other rotations of 15-20 years, but also as high forest with cuts every
continents (Chile, New Zealand)1, 11, 12 . 70-80 years13 . Timber quality is similar to other alders and hybrid
poplars, used traditionally for firewood and also for turning and
Habitat and Ecology carving as well as for carpentry, furniture, panelling, plywood and
Italian alder occurs in the Mediterranean sub-mountain paper pulp1, 5, 11, 14 . Like other alders, the wood is degraded rapidly Maturing catkins during mid summer: they appear in early summer, are
and mountain belt. Unlike other alders, it is less dependent on dormant in winter and are fully functional at the end of winter.
when exposed to air or soil, but is more durable when immersed (Copyright Franco Caldararo, www.actaplantarum.org: AP)
riparian habitats and is more drought tolerant, although it still in water1, 5 .
tends to concentrate in water accumulation zones and needs References
climates with an annual precipitation of at least 1000mm per Threats and Diseases [1] V. M. Loewe, C. R. Delard, Monografa de [10] A. Bezzi, P. Brandini, G. Menguzzato, G.
year13 . It is a heliophilous species, but can be shade-tolerant Despite its limited natural range, Italian alder is not aliso italiano (Alnus cordata), vol. 204 Tabacchi, Annali dellIstituto Sperimentale
(Instituto Forestal, Santiago, Chile, 1998). per lAssestamento Forestale e per
under favourable rainfall regimes5, 11 . It grows in on most kinds considered an endangered species, because it grows over a wide [2] A. F. Mitchell, A field guide to the trees lAlpicoltura 12, 3 (1989).
of soils, including degraded, but preferring calcareous. The root range of elevations and can spread very rapidly5 . Diseases are of Britain and northern Europe (Collins, [11] A. Praciak, et al., The CABI encyclopedia of
1974). forest trees (CABI, Oxfordshire, UK, 2013).
system promotes a symbiosis with the nitrogen-fixing bacterium [3] O. Johnson, D. More, Collins tree guide [12] V. M. Loewe, A. C. lvarez, L. M. Barrales,
Actinomyces alni (Frankia alni) improving soil fertility1, 11 . In (Collins, 2006). Proceedings of 4th International Scientific
[4] M. Goldstein, G. Simonetti, M. Watschinger, Conference on Hardwood Processing
optimal habitats this alder is fast-growing and behaves as a Alberi dEuropa (A. Mondadori, 1995). 2013, S. Berti, et al., eds. (2013), vol. 1,
pp. 5061.
pioneer species, forming pure stands beside Turkey oak (Quercus [5] F. Ducci, A. Tani, EUFORGEN Technical
Guidelines for genetic conservation and [13] R. Del Favero, I boschi delle regioni
cerris) and beech (Fagus sylvatica) woods5, 13 . It also tends to use for Italian alder (Alnus cordata) meridionali e insulari dItalia (Cleup,
Padova, 2008).
colonise open woods such as black pine (Pinus nigra) plantations (Bioversity International, 2009).
[6] H. Meusel, E. Jager, S. Rauschert, E. [14] P. S. Savill, The silviculture of trees used in
in wetter conditions and abandoned chestnut (Castanea sativa) Weinert, Vergleichende Chorologie der British forestry (CABI, 2013).
orchards5, 10, 13 . It can be found as first-stage species in bare soils Zentraleuropischen Flora (Gustav Fischer [15] J. F. Webber, J. N. Gibbs, S. J. Hendry,
Verlag Jena, 1978). Phytophthora disease of Alder
after wildfires or landslides13 . [7] J. Jalas, J. Suominen, Atlas Florae (Information note) (Forestry Commission,
Europaeae: distribution of vascular Edinburgh, 2004).
plants in Europe Vol. 3 Salicaceae to [16] T. Jung, M. Blaschke, Plant Pathology 53,
Balanophoraceae (Committee for Mapping 197 (2004).
the Flora of Europe and Societas Biologica [17] E. Dallavalle, A. Zambonelli, Forest
Fennica Vanario, Helsinki, 1976). Pathology 29, 97 (1999).
[8] P. W. Ball, Flora Europea. Volume 1. [18] A. Santini, G. P. Barzanti, P. Capretti, Plant
Psilotaceae to Platanaceae, T. G. Tutin, Disease 85, 560 (2001).
et al., eds. (Cambridge University Press,
1993), pp. 6970, second edn. [19] EUFORGEN, Distribution map of italian
alder (Alnus cordata) (2009).
[9] J. Briquet, R. de Litardiere, Prodrome de www.euforgen.org.
la flore corse, comprenant les rsultats
botaniques de six voyages excuts en
Corse sous les auspices de M. mile
Burnat, Vol. 1 (Georg & Co., Genve, Bale,
Lyon, 1910).
Beech leaves.
(Copyright Jannik Selz, unsplash.com: CC0)
Distribution
Alder can be found over most of Europe, from Scandinavia
to the Mediterranean countries and parts of North Africa4 . It Map 1: Plot distribution and simplified chorology map for Alnus glutinosa.
normally grows below 1000m in elevation, although in the Frequency of Alnus glutinosa occurrences within the field observations as Alder trees on the banks of the river Wey, England.
reported by the National Forest Inventories. The chorology of the native
mountains of central Europe it can occasionally be found along spatial range for A. glutinosa is derived after EUFORGEN22 .
(Forestry Commission, www.forestry.gov.uk: Crown Copyright)
Map 2: High resolution distribution map estimating the relative probability of presence. Male inflorescence, Sierra Madrona, Spain.
(Copyright Javier Martin, commons.wikimedia.org: PD)
Uncertain, no-data
Tundra, cold desert
Negligible survivability
Low survivability
Mid-low survivability
Medium survivability
Mid-high survivability
High survivability
Foliage and mature cones, Ispra, Italy. (Copyright Daniele de Rigo: CC-BY)
References
[1] P. Schtt, H. J. Schuck, B. Stimm, Lexikon [11] D. T. Funk, Alnus glutinosa (L.) Gaertn.
der Baum- und Straucharten: Das European Alder, Agriculture Handbook
Standardwerk der Forstbotanik (Nikol, 654 (U.S. Department of Agriculture,
Hamburg, 2002). Forest Service, Washington, DC., 1990),
[2] H. H. Ellenberg, Vegetation Ecology of pp. 239256.
Central Europe (Cambridge University [12] P. S. Savill, The silviculture of trees used in
Press, 2009), fourth edn. British forestry (CABI, 2013).
[3] A. Krstinic, J. Gracan, D. Kajba, Noble [13] D. N. McVean, Journal of Ecology 44 (1956).
Hardwoods Network: Report of the Fourth [14] H. Claessens, A. Oosterbaan, P. Savill, J.
Meeting, 4-6 September 1999, Gmunden, Rondeux, Forestry 83, 163 (2010).
Austria and the Fifth Meeting, 17-19
May 2001, Blessington, Ireland, J. Turok, [15] A. Akkermans, et al., Nitrogen-fixing
G. Eriksson, K. Russel, S. Borelli, eds. actinorhizal symbioses (Springer, 2008).
(Bioversity International, 2002), pp. 4449. [16] R. K. W. M. Klaassen, J. G. M. Creemers,
[4] D. N. McVean, Journal of Ecology 41 Journal of Cultural Heritage 13, S123
(1953). (2012). Wood Science for Conservation.
[5] A. F. Mitchell, A field guide to the trees [17] R. A. Housley, A. J. Ammerman, C.
of Britain and northern Europe (Collins, E. McClennen, Journal of Wetland
1974). Archaeology 4, 139 (2004).
Alder forest in Spree, Germany. (Copyright Paul Schulze, www.flickr.com: CC-BY) [6] D. Kajba, J. Graan, EUFORGEN Technical [18] A. Praciak, et al., The CABI encyclopedia of
guidelines for genetic conservation and forest trees (CABI, Oxfordshire, UK, 2013).
use for Black alder (Alnus glutinosa) [19] J. F. Webber, J. N. Gibbs, S. J. Hendry,
(2003). Phytophthora disease of Alder
Autoecology diagrams based on harmonised [7] D. N. McVean, Journal of Ecology 43, 61 (Information note) (Forestry Commission,
Field data in Europe (including absences) Observed presences in Europe field observations from forest plots. (1955). Edinburgh, 2004).
[8] U. Pietzarka, A. Roloff, Enzyklopdie der [20] O. Johnson, D. More, Collins tree guide
Holzgewchse: Handbuch und Atlas der (Collins, 2006).
Dendrologie, A. Roloff, H. Weisgerber, U. M. [21] J. N. Gibbs, M. A. Lipscombe, A. J. Peace,
Lang, B. Stimm, P. Schtt, eds. (Wiley-Vch
Average temperature of the coldest month (C)
63517/0+. www.euforgen.org.
[10] G. E. Hemery, et al., Forestry 83, 65 (2010).
the common alder (Alnus glutinosa)1 . The bark is smooth and > 75%
Chorology
deep grey, developing fissures with age2, 4 . The leaves are oval to Native
Distribution
Grey alder is native to most of central Europe, extending
Map 1: Plot distribution and simplified chorology map for Alnus incana.
westwards towards France and east into Russia, the Caucasus
Frequency of Alnus incana occurrences within the field observations as
and western Siberia3, 6 . It is widespread in Scandinavia and has reported by the National Forest Inventories. The chorology of the native
been introduced in Britain6 . Two subspecies (subsp. rugosa and spatial range for A. incana is derived after Meusel and Jger21 .
communities3 . It prefers mesic and moist conditions and it is hybrids, although this is not common as the two species flower at
tolerant of acid soils, able to stand pH levels of 3.5-4.0 without slightly different times: the grey alder around a week earlier than
Importance and Usage
The timber of the grey alder has little commercial value except
problems, but it is able to grow on a wide range of soil types, the common alder14, 15 .
as fuel wood, although it is suitable for carpentry and turning and
is reported to make good charcoal for drawing11 . It has several
advantages as a short-rotation crop. It is relatively untroubled by
grazing animals and has few pests and diseases, it has modest
site requirements, coppices easily and combines fast growth with
Uncertain, no-data the ability to improve soil fertility5, 10, 13, 16, 17. It is now increasingly
Marginal/no presence < 5% being considered for biomass production in several countries18, 19,
Low presence 5% - 10%
as well as a potential suitable alternative species for reforestation
of former noble hardwood areas18 . It is also useful for restoration
Mid-low presence 10% - 30% of disturbed sites including old mines, for consolidating the ground
Medium presence 30% - 50% in wet woods, river-banks and on unstable slopes3, 9, and it is
Mid-high presence 50% - 70%
suitable for planting on polluted sites10. It has been historically
used for medicinal purposes for a range of ailments from sprains
High presence 70% - 90% and bruises to urinary problems and anaemia3, 20 .
Very-high presence > 90%
Map 2: High resolution distribution map estimating the relative probability of presence. Ovate leaves with toothed margins.
(Copyright Free Photos, www.flickr.com: CC-BY)
Uncertain, no-data
Tundra, cold desert
Negligible survivability
Low survivability
Mid-low survivability
Medium survivability
Mid-high survivability
High survivability
References
[1] H. Hytteborn, A. A. Maslov, D. I. Nazimova, [11] V. L. Komarov, et al., Flora of the USSR -
L. P. Rysin, Ecosystems of the World, Vol. Volume V (Keter Press, Jerusalem, 1970).
6: Coniferous Forests, F. A. Andersson, ed. [12] A. Praciak, et al., The CABI encyclopedia of
(Elsevier, 2005), pp. 2399. forest trees (CABI, Oxfordshire, UK, 2013).
[2] O. Johnson, D. More, Collins tree guide [13] L. Kullman, New Phytologist 120, 445
(Collins, 2006). (1992).
[3] K. Shaw, B. Wilson, S. Roy, The IUCN Red [14] E. V. Banaev, V. Bazant, Journal of Forest
List of Threatened Species (2014), pp. Science 53, 66 (2007).
194472/0+.
[15] P. A. Tallantire, New Phytologist 73, 529
[4] A. F. Mitchell, A field guide to the trees (1974).
of Britain and northern Europe (Collins,
1974). [16] J. Aosaar, M. Varik, V. Uri, Biomass and
Bioenergy 45, 11 (2012).
[5] J. L. Fryer, Alnus incana. Fire Effects
Information System (2011). [17] V. Uri, et al., Ecological Engineering 37,
http://www.feis-crs.org/feis 920 (2011).
[6] P. S. Savill, The silviculture of trees used in [18] N. Arhipova, T. Gaitnieks, J. Donis, J.
British forestry (CABI, 2013). Stenlid, R. Vasaitis, Forestry 84, 337
(2011).
Isolated grey alders in swamp areas in Rheinhessen-Pfalz (Germany). [7] V. Bojnansk, A. Fargaov, Atlas of Seeds
and Fruits of Central and East-European [19] T. Krki, M. Maltamo, K. Eerikinen, New
(Copyright AnRo0002, commons.wikimedia.org: CC0)
Flora: The Carpathian Mountains Region Forests 20, 65 (2000).
(Springer Netherlands, 2007). [20] U. Quattrocchi, CRC World Dictionary of
Autoecology diagrams based on harmonised [8] E. Hultn, M. Fries, Atlas of North European Medicinal and Poisonous Plants: Common
Field data in Europe (including absences) Observed presences in Europe field observations from forest plots. vascular plants (North of the Tropic of Cancer), Names, Scientific Names, Eponyms,
Vols. I-III. (Koeltz scientific books, 1986). Synonyms, and Etymology (5 Volume Set)
(Taylor & Francis, 2012).
[9] K. Huss-Danell, J. E. Lundmark, Studia
forestalia Suecica 181, 20 (1988). [21] H. Meusel, E. Jger, eds., Vergleichende
Chorologie der Zentraleuropischen Flora
Average temperature of the coldest month (C)
[10] L. Rytter, Short Rotation Willow Coppice - Band I, II, III (Gustav Fischer Verlag,
for Renewable Energy and Improved Jena, 1998).
Environment: Proceedings of a joint
Swedish - Estonian seminar on Energy
Annual precipitation (mm)
from grey to blue-grey colour. The leaves are alternate, sticky > 75%
Chorology
when young, from 7 to 14cm long and from 3 to 10cm wide. The Native
Betula pendula, Betula pubescens and other birches in Europe: distribution, habitat, usage and threats
P. Beck, G. Caudullo, D. de Rigo, W. Tinner
Silver birch (Betula pendula Roth) and downy birch (Betula pubescens Ehrh.) are short-lived, relatively small broadleaved
trees that occur throughout most of Europe, particularly in northern regions. In southern Europe, birch trees are confined
to mountainous areas, as they do not tolerate prolonged summer drought. Birch has a light canopy of small serrated
leaves, and characteristic smooth, white to grey bark. In northern regions, birch trees can dominate the landscape up
to the treeline, whereas in the centre of their range they often occur early in secondary succession because of their
abundant seed production, low demands on soil quality, and intolerance of shade. Birch trees provide the predominant
hard wood source in northern Europe, and some varieties of Betula pendula produce highly priced veneers, while Betula
pubescens is mostly used for pulp and fire wood. Other rarer species of birch are endemic to Europe contributing to the
continental biodiversity even at high elevations and latitudes.
Betula pendula Roth is a medium-sized tree, growing up to
30m, while Betula pubescens Ehrh. is relatively shorter, rarely
growing beyond 20m and also less towards its northern range
limits, up to dwarf trees in extreme habitats in the northern
tundra and on mountains1, 2 . The bark of young trees is brown
in colour; when mature it turn to silvery-white, with horizontal
dark grey lenticels, that with age darkens and develops fissures.
The bark of the Betula pendula is a brighter white and shinier
than that of Betula pubescens, and its branches characteristically
droop, whereas those of Betula pubescens grow upwards or
horizontally. In addition, Betula pubescens shoots are covered
with a smooth fine down, as opposed to the hairless shoots of
Betula pendula. Betula pendula leaves are coarsely and unequally
double-serrated, larger than those of Betula pubescens (3-7cm
vs. 2-5cm), and end in a fine point. Betula pubescens leaves are
egg shaped, with a finely serrated margin and end in a shorter
point3 . Both species are monoecious with male and female
Silvery-white bark on a young tree with dark scars from dropped shoots and
small horizontal dark grey lenticels.
(Copyright Tracy Houston Durrant: CC-BY)
Distribution
Betula pendula and Betula pubescens occur naturally
throughout most of Europe up to central Siberia. Betula pubescens
Uncertain, no-data
has a more northerly and easterly distribution, growing further north
Marginal/no presence < 5% in Europe than any other tree species, whereas Betula pendula
Low presence 5% - 10% can reach southern regions such as Iberian Peninsula, South Italy
and Greece5 . Given their wide distribution, these two birches show
Mid-low presence 10% - 30%
a high morphological variability and different subspecies and
Medium presence 30% - 50% varietals have been described6 . Moreover, in most parts of Europe
Mid-high presence 50% - 70% they are sympatric and can naturally hybridise, generating plants
High presence 70% - 90%
Very-high presence > 90%
Annual average temperature (C) Potential spring-summer solar irradiation (kWh m-2) Seasonal variation of monthly precipitation (dimensionless)
In Europe other two main species of genus Betula are described: Betula
nana (artic dwarf birch) and Betula humilis (dwarf birch). Some authors
identify other birch species, often rare, endemic and at the limits of
the geographical ranges, which have not explicitly a defined systematic
status and are treated in some case as hybrids, varieties or subspecies7.
Betula nana is a shrubby birch occurring in a broad geographic range
of Northern Europe, which spans from Iceland, Scotland and northern
England up to Scandinaviaand the Baltic area. In Central Europe it
occurs at high elevations (northern Alps from Austria west to France;
Carpathian mountains)34 . This birch lives in Arctic of high-mountain
exposed environments. It is found in immature or peaty soils within
alpine tundra, rocky barrens and moorlands, subalpine damp moorlands
and open raised bogs34 . Betula humilis is another shrubby birch which
has a very wide but scattered distribution, ranging from Western Europe Triangular smooth leaves of silver birch (Betula pendula) with toothed margins.
(Copyright Tracy Houston Durrant: CC-BY)
with few locations in Germany, Austria, Poland, Romania and Switzerland,
through Siberia up to Korea35, 36 . It is a relict birch distributed from the
hill to the montane zone, preferring wet soils in forests and the edges
of lakes. It may grow in shrubby pastures and alder thickets, transitional
mires, on open raised or acid valley bogs and in natural/drained fens35, 37.
Both these dwarf birches are diploid. They can naturally and frequently
hybridise with Betula pubescens and Betula pendula in the overlapping
Annual precipitation (mm)
Uncertain, no-data
Tundra, cold desert
Negligible survivability
Low survivability
Mid-low survivability
Medium survivability
Mid-high survivability
High survivability
References
[1] K.Shaw, S.Roy, B.Wilson, The IUCN Red [22] I.Svanberg, etal., Acta Societatis
List of Threatened Species (2014), pp. Botanicorum Poloniae 81, 343 (2012).
194521/0+. [23] K.Shaw, etal., The Red List of Betulaceae
Map 3: High resolution map estimating the maximum habitat suitability for the whole genus Betula. [2] M.Walters, Flora Europaea, Volume 1: (Botanic Gardens Conservation
Psilotaceae to Platanaceae, T.G. Tutin, International, Richmond, United Kingdom,
etal., eds. (Cambridge University Press, 2014).
1993), pp. 6970, second edn. [24] C.Bosco, D.deRigo, O.Dewitte, J.Poesen,
associated with more general die-back with affects crown health Europe, it is also harmful for Betula pendula which partly coexists [3] J.Lid, Norsk flora (Norske samlaget, P.Panagos, Natural Hazards and Earth
(i.e. Anisogramma virgultorum and Discula betulina)28, 29 . Although with the natural niche of the large pine weevil30-32 . Herbivory 1994), 6th edn. System Science 15, 225 (2015).
the large pine weevil (Hylobius abietis) is mostly known as one by short-snouted weevils (Strophosoma melanogrammum and [4] A.Praciak, etal., The CABI encyclopedia of [25] J.E. Norris, A.DiIorio, A.Stokes, B.C.
forest trees (CABI, Oxfordshire, UK, 2013). Nicoll, A.Achim, Slope Stability and
of the most serious pests affecting young coniferous forests in Otiorhynchus scaber) is another threat to birch33 . [5] H.Meusel, E.Jger, eds., Vergleichende Erosion Control: Ecotechnological
Chorologie der Zentraleuropischen Flora Solutions, J.E. Norris, etal., eds. (Springer
- Band I, II, III (Gustav Fischer Verlag, Netherlands, 2008), pp. 167210.
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[6] .T. Thrsson, E.Salmela, Evolutionary Biology 20, 369 (2007).
Frequency K.Anamthawat-Jnsson, Journal of [27] J.U. Jepsen, S.B. Hagen, R.A. Ims, N.G.
< 25% Heredity 92, 404 (2001). Yoccoz, Journal of Animal Ecology 77,
25% - 50% [7] K.Jadwiszczak, Silva Fennica 46 (2012). 257 (2008).
50% - 75% [8] M.F. Schenk, C.-N. Thienpont, W.J.M. [28] S.Green, G.A. MacAskill, Plant Pathology
> 75% Koopman, L.J. W.J. Gilissen, M.J.M. 56, 242 (2007).
Chorology Smulders, Tree Genetics & Genomes 4, [29] S.-O. Holm, Ecography 17, 60 (1994).
Native 911 (2008). [30] J.I. Barredo, etal., EPPO Bulletin 45, 273
[9] M.D. Atkinson, A.P. Jervis, R.S. Sangha, (2015).
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1896 (1997). (2015). Invasive Species Compendium.
[10] M.D. Atkinson, Journal of Ecology 80, http://www.cabi.org
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[11] J.Webber, H.Evans, Annual Report and Entomology 8, 121 (2006).
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Edinburgh, 2003), pp. 1627. T.N. Welander, Forest Ecology and
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[13] A.F. Mitchell, P.Dahlstrom, E.Sunesen, Species (2014), pp. 194495/0+.
C.Darter, A field guide to the trees of [35] K.Shaw, S.Roy, B.Wilson, The IUCN Red
Britain and northern Europe (Collins, List of Threatened Species (2014), pp.
1974). 194645/0+.
[14] L.Augusto, J.Ranger, D.Binkley, A.Rothe, [36] K.A. Jadwiszczak, E.Jabonska,,
Annals of Forest Science 59, 233 (2002). S.Kosowski, A.Banaszek, Acta Societatis
[15] A.Moen, National atlas of Norway: Botanicorum Poloniae 80, 233 (2011).
vegetation (Norwegian Mapping Authority, [37] H.McAllister, K.Ashburner, Curtiss
Honefoss, 1999). Botanical Magazine 24, 174 (2007).
Map 1-B: Plot distribution and simplified chorology map for Betula pubescens. [16] L.Stritch, K.Shaw, S.Roy, B.Wilson, The [38] K. Anamthawat-Jnsson, A. Thr Thrsson,
Frequency of Betula pubescens occurrences within the field observations IUCN Red List of Threatened Species Plant Cell, Tissue and Organ Culture 75,
Young foliage of downy birch (Betula pubescens), which is covered by a as reported by the National Forest Inventories. The chorology of the native (2014), pp. 62535/0+. 99 (2003).
smooth down, unlike the silver birch (Betula pendula). spatial range for B. pubescens is derived after Meusel andJger5 . [17] P.Bekhta, S.Hiziroglu, O.Shepelyuk,
(Copyright S. Rae, www.flickr.com: CC-BY) [39] A.E. Palme, Q.Su, S.Palsson, M.Lascoux,
Materials & Design 30, 947 (2009). Molecular Ecology 13, 167 (2004).
[18] A.Felton, E.Andersson, D.Ventorp, [40] N.Wang, etal., Molecular Ecology 22,
Autoecology diagrams based on harmonised field M.Lindbladh, Silva Fennica 45, 1143
Field data in Europe (including absences) Observed presences in Europe observations from forest plots for Betula pubescens. (2011).
3098 (2013).
[41] O.Maliouchenko, A.E. Palm,
[19] K.Wannebo-Nilsen, J.W. Bjerke, P.S.A. A.Buonamici, G.G. Vendramin, M.Lascoux,
Beck, H.Tmmervik, Boreal Environment Journal of Biogeography 34, 1601 (2007).
Research 15, 43 (2010).
[42] EUFORGEN, Distribution map of silver
Average temperature of the coldest month (C)
domed in old trees. The bark is smooth and steel grey, having a > 75%
Chorology
muscled character to its appearance6 . The leaves are alternate, Native
to April and are wind-pollinated. The fruits are clustered in about native trees in Europe, though slightly less than beech7. For this
8 pairs of nutlets (achene), 6-8mm, each pair at the base of a reason this species can play roles both as a secondary species
green leathery tri-lobate bract, 3.5cm long1, 6, 7. The hornbeam and as an understorey tree and also as a coloniser on bare and
Map 1: Plot distribution and simplified chorology map for Carpinus betulus.
is an abundant seeding tree and is marked by vigorous natural Frequency of Carpinus betulus occurrences within the field observations disturbed soils whose fertility is improved by its growth3 . In mixed
regeneration. Seeds often do not germinate until the spring of the as reported by the National Forest Inventories. The chorology of the native
forests it can be a dangerous invader, regenerating better and
spatial range for C. betulus is derived after several sources8, 23-25 .
second year after sowing7. faster than valuable timber species, such as oaks, ash (Fraxinus
excelsior) or Scots pine (Pinus sylvestris)3, 10 .
Distribution
Habitat and Ecology The common hornbeam grows mostly in mixed stands
The hornbeam has a wide range which covers southern Europe
dominated by deciduous oaks (Quercus robur, Quercus petraea),
(excluding the Iberian Peninsula), Central Europe, up to southern The hornbeam is a typical mesophilous species of temperate
forming oak-hornbeam forest communities. This vegetation
England and the south of Sweden. Eastwards it occurs across the climates, occurring on lowlands, hills and the low mountain belt.
represents the classic European temperate forest on fertile
Black Sea reaching the Caucasus and northern Iran8 . Its altitudinal High summer temperatures limit its distribution in the south,
soils, typically with ash (Fraxinus excelsior), small-leaved lime
distribution ranges from sea level to 700m in Central Europe, while it is a very hardy species and even found in frost hollows.
(Tilia cordata), wild cherry (Prunus avium), field maple (Acer
1000m in the Western Alps and 1800m in Iran8, 9 . It favours deep moist and well-drained soils from sub-acid to
campestre), common hazel (Corylus avellana) and spindle
calcareous, although it can tolerate wet heavy clay to light dry
(Euonymus europaeus). The hornbeam can also be found in beech
sandy soils, but never acid3 . It grows in full to partial sunny
forests (Fagus sylvatica), while pure stands are more rare11-14 .
conditions and it is also one of the few strongly shade-tolerant
Uncertain, no-data
Marginal/no presence < 5%
Low presence 5% - 10%
Mid-low presence 10% - 30%
Medium presence 30% - 50%
Mid-high presence 50% - 70%
High presence 70% - 90%
Very-high presence > 90%
It is not flexible and shrinks greatly during the drying process. For
these reasons hornbeam has low commercial interest and has
never been industrially cultivated16 .
In the past when metals were scarce and costly it was used Uncertain, no-data
more, for making small items which require resistance to wear, Tundra, cold desert
such as tool handles, mill wheels, agricultural tools, wooden rivets,
Negligible survivability
etc.1 . More recently it is used for flooring, billiard cues, drumsticks
and piano mechanisms, and sometimes as an alternative to Low survivability
maple3, 7. The wood has a high calorific value as it burns slowly, Mid-low survivability
making it excellent fuel wood and charcoal3, 7.
Medium survivability
Because of its ability to regenerate from root suckers, it can
be cultivated in mixed coppices alongside oaks, limiting them in Mid-high survivability
producing epicormic branches. It responds well even when pollarded, High survivability
making it a good hedgerow and fodder tree. The hornbeam is also
planted with oak in afforestation plantations on bare sloping areas
for soil protection from erosion and landslides and maintained as a
bush when needed3. Different varieties are available for ornamental
purposes; one of the more frequently used is the Fastigicata
with a regular balloon-shape crown, more rare is the Columnaris
with densely teardrop shape, or the Incisa with small and deeply
lobed leaves. They can be found in urban parks, gardens and along
roadsides, locally abundant on richer soils2, 6 .
References
[1] V. L. Komarov, et al. , Flora of the USSR - [14] European Environment Agency, EUNIS,
Volume V (Keter Press, Jerusalem, 1970). the European Nature Information System
[2] O.Johnson, D.More, Collins tree guide (2015). http://eunis.eea.europa.eu
(Collins, 2006). [15] H.J. Elwes, A.Henry, The Trees of Great
[3] A.Praciak, etal., The CABI encyclopedia of Britain and Ireland Vol. 6 (Privately
forest trees (CABI, Oxfordshire, UK, 2013). printed, Edinburgh, 1912).
[4] S.Orze, Electronic Journal of Polish [16] M.Christy, The Journal of Ecology 12, 39
Agricultural Universities 10 (2007). (1924).
[5] MonumentalTrees.com, Monumental trees [17] A.Turb, etal., Disturbances of EU forests
(2015). caused by biotic agents - final report,
Tech. Rep. KH-32-13-151-EN-N (2011).
[6] A.F. Mitchell, P.Dahlstrom, E.Sunesen, Final Report prepared for European
C.Darter, A field guide to the trees of Commission (DG ENV).
Britain and northern Europe (Collins, 1974).
[18] T.Wesoowski, P.Rowiski, Forest Ecology
[7] P.S. Savill, The silviculture of trees used in and Management 221, 299 (2006).
British forestry (CABI, 2013).
[19] G.E. King, British Journal of Entomology
[8] H.Meusel, E.Jager, eds., Vergleichende and Natural History 11, 153 (1999).
Chorologie der Zentraleuropischen Flora
- Band I, II, III (Gustav Fischer Verlag, [20] D.deRigo, etal., Scientific Topics Focus 2,
Jena, 1998). mri10a15+ (2016).
[9] I. C. Paridari, S. G. Jalali, A. Sonboli, M. [21] CABI, Lymantria dispar (gypsy moth)
Zarafshar, P. Bruschi, Journal of Forestry (2015). Invasive Species Compendium.
Research 24, 301 (2013). http://www.cabi.org
[10] A. J. Kwiatkowska, K. Spalik, E. Michalak, A. [22] CABI, Thaumetopoea processionea (oak
Paliska, D. Panufnik, Plant Ecology 129, processionary moth) (2015). Invasive
1 (1997). Species Compendium. http://www.cabi.org
[11] H.H. Ellenberg, Vegetation Ecology of [23] J.Jalas, J.Suominen, Atlas Florae
Central Europe (Cambridge University Europaeae: distribution of vascular
Press, 2009), fourth edn. plants in Europe Vol. 3 Salicaceae to
Balanophoraceae (Committee for Mapping
[12] P.Koir, etal., Plant Biosystems - An the Flora of Europe and Societas Biologica
International Journal Dealing with all Fennica Vanario, Helsinki, 1976).
Aspects of Plant Biology 147, 84 (2012).
[24] Botanical Society of Britain & Ireland, BSBI
[13] U.Bohn, etal., Karte der natrlichen Big Database (2015). http://bsbidb.org.uk.
Vegetation Europas; Map of the
Natural Vegetation of Europe [25] Tela Botanica, eFlore (2015).
(Landwirtschaftsverlag, 2000). http://www.tela-botanica.org
Obovate leaves with serrated margins: they are quite similar to those
of the beech, but less shiny.
(Copyright AnRo0002, commons.wikimedia.org: CC0)
Distribution regions coppiced stands are also used as a food resource for
The oriental hornbeam is a tree species native to south- livestock in drought summers, when grasslands are completely
east Europe, the Pontic region and western Asia. It is found in dry11, 18 . This frugal hornbeam is also suitable for the reforestation
southern parts of Italy, Balkan Peninsula, Turkey, Syria, Caucasus and restoration of degraded dry lands5 and is highly resistant to
and northern Iran, usually occurring at lower altitudes or on wildfire19 . It may be used as an ornamental plant, appreciated for
southern slopes up to 1300m in Europe, but growing at over its dense foliage and pollution resistance, and also as a hedge
2500m in the Caucasus mountains5-8 . because of its re-sprouting capability20 .
References
[1] V. L. Komarov, et al. , Flora of the USSR - [13] A.arni, etal., Plant Biosystems 143, 1
Volume V (Keter Press, Jerusalem, 1970). (2009).
[2] H.J. Elwes, A.Henry, The Trees of Great [14] R.Popovi, M.Koji, B.Karadi, Bocconea
The oriental hornbeam is a shrub or small tree and rarely reaches 15m. Britain and Ireland Vol. 6 (Privately 5, 431 (1997).
(Copyright Stefano Zerauschek, www.flickr.com: AP) printed, Edinburgh, 1912). [15] V.Matevski, etal., Forest vegetation of
[3] O.Johnson, D.More, Collins tree guide the Galiica mountain range in Macedonia
(Collins, 2006). (Zaloba ZRC, Ljubljana, 2011).
Habitat and Ecology [4] A.F. Mitchell, P.Dahlstrom, E.Sunesen,
C.Darter, A field guide to the trees of
[16] U.Bohn, etal., Karte der natrlichen
Vegetation Europas; Map of the
This hornbeam is a thermophilous and xerophilous species, Britain and northern Europe (Collins, Natural Vegetation of Europe
1974). (Landwirtschaftsverlag, 2000).
drought-resistant, thriving principally on slopes in shallow The bark is smooth and grey. [5] M.Goldstein, G.Simonetti, M.Watschinger, [17] P.M. Pijut, The Woody Plant Seed Manual,
humus-poor or even rocky soils, and preferring calcareous (Copyright Stefano Zerauschek, www.flickr.com: AP) Alberi dEuropa (A. Mondadori, 1995). F.T. Bonner, R.P. Karrfalt, eds., Agriculture
[6] H.Meusel, E.Jger, eds., Vergleichende Handbook 727 (U.S. Department of
substrates (rendzina)9 . Over its wide distribution range, this Chorologie der Zentraleuropischen Flora Agriculture, Forest Service, 2008), pp.
wood. It was used more in the past for making tool handles and - Band I, II, III (Gustav Fischer Verlag, 328332.
species exhibits different ecological habits. In eastwards regions Jena, 1998). [18] V.P. Papanastasis, P.D. Platis, O.Dini-
it occurs at higher elevations tolerating lower temperatures in other small household items1, 17. Thanks to its high aptitude for Papanastasi 37, 187 (1997).
[7] S.Gcel, K.zkan, S.Celik, E.Ycel,
more temperate climates10 . In south-east Europe it is a typical regeneration from root suckers, it can be managed in coppice M.ztrk, Pakistan Journal of Botany 40, [19] S.S. Radanova, Ecologia Balkanica 5, 55
1497 (2008). (2014).
element of the sub-Mediterranean vegetation, and it can also stands for fuel production as firewood or charcoal5, 17. In southern
[8] F.Assadolahi, M.Barbero, P.Quezel, [20] T.Tsitsoni, M.Tsakaldimi, C.Tsouri, African
Ecologia Mediterranea 8, 365 (1982). Journal of Agricultural Research 8, 4501
be found in the inner regions with some continental influences (2013).
[9] A.Chiarucci, D.Dominicis, V.A. Gabellini,
(colder winters)11-13 . It is very frugal, easily able to colonise open Atti della Societ Toscana di Scienze [21] D.deRigo, etal., Scientific Topics Focus 2,
Naturali - Memorie serie B 103, 107 mri10a15+ (2016).
and degraded areas and to regenerate vigorously, promoting its (1996). [22] CABI, Lymantria dispar (gypsy moth)
presence in disturbed habitats, such as after the exploitation of [10] H. Akhani, H. Ziegler, Phytocoenologia 32, (2015). Invasive Species Compendium.
455 (2002). http://www.cabi.org
primary oak forests11, 14 . This species is found both as a dominating
[11] C.Blasi, R.DiPietro, L.Filesi, P.Fortini, [23] CABI, Thaumetopoea processionea (oak
and secondary species in wood and shrub lands11 . The principal Phytocoenologia 31, 33 (2001). processionary moth) (2015). Invasive
Species Compendium. http://www.cabi.org
tree communities in which it is found are the mixed deciduous [12] A.Kavgaci, A.arni, B.Tecimeni, G.zalp,
Archives of Biological Sciences 62, 705 [24] H. Meusel, E. J. Jger, Plant Systematics
forests with oaks, such as downy oak (Quercus pubescens), (2010). and Evolution 162, 315 (1989).
Turkey oak (Quercus cerris), Hungarian oak (Quercus frainetto),
and with hop hornbeam (Ostrya carpinifolia) and South European
This is an extended summary of the chapter. The full version of
ash (Fraxinus ornus)11, 15, 16 . this chapter (revised and peer-reviewed) will be published online at
https://w3id.org/mtv/FISE-Comm/v01/e01bf18. The purpose of this
Importance and Usage summary is to provide an accessible dissemination of the related
main topics.
This QR code points to the full online version, where the most
Like other hornbeams (sometimes called ironwoods), the updated content may be freely accessed.
wood of the oriental hornbeam is very hard17. Because of its Please, cite as:
Sikkema, R., Caudullo, G., 2016. Carpinus orientalis in Europe:
small size and bushy habit, this tree does not produce high value distribution, habitat, usage and threats. In: San-Miguel-Ayanz,
Male catkins are 2-3cm long.
J., de Rigo, D., Caudullo, G., Houston Durrant, T., Mauri, A. (Eds.),
(Copyright Silvano Radivo, www.actaplantarum.org: AP)
European Atlas of Forest Tree Species. Publ. Off. EU, Luxembourg,
pp. e01bf18+
Broadleaved and coniferous mixed forest over the coast cliffs of the
Forggensee lake (Schwangau, Germany).
(Copyright Thomas Richter, unsplash.com: CC0)
Edible nuts of the sweet chestnut: they are traditionally roasted but can
also be candied, boiled, dried, or used as flour.
(Copyright Patrik Krebs: CC-BY)
The trunk is vertical, robust and enlarging at the bottom with > 75%
Chorology
age, with girth of 3 m, exceptionally up to 6m3 . The bark is Native
lived (more than 600 years) and can reach heights of up to 50m
(exceptionally up to 70m in its native range) and a diameter
exceeding 2m1, 2 . The tree is narrowly columnar with slender,
down-curving branches; frequently with forked stems. The bark is
silvery-brown, becoming furrowed and very thick with age giving
mature trees good fire resistance2, 3 . The wood is highly aromatic
with a distinctive ginger-like odour, as is the foliage which has
a parsley-like scent when crushed3, 4 . The evergreen scale-like
leaves are around 2-3mm long5 . Abundant, round pea-sized
cones ripen in autumn with seed dispersal occurring immediately
after and continuing until the following spring6 .
Distribution
The native range of Lawson cypress is a narrow strip
between Oregon and north-west California, mainly near the Map 1: Plot distribution and simplified chorology map for Chamaecyparis lawsoniana.
Frequency of Chamaecyparis lawsoniana occurrences within the field observations as
coast. It was introduced into Europe in 1854 and named after reported by the National Forest Inventories.
the Scottish nursery (Lawson & Son) where it was first sent7. It
is now established, though not common, in Germany, France, the
Netherlands, Denmark and United Kingdom, and also outside
Importance and Usage Ornamental specimen in a park in Varese (North Italy).
Europe in Australia, South Africa, Kenya, New Zealand and Sri The main use for Lawson cypress outside its natural range (Copyright Achille Mauri: CC-BY)
Lanka1 . Other species of Chamaecyparis are present in Europe. is as an ornamental tree, and there are over 200 cultivars with
In particular, Hinoki cypress (Chamaecyparis obtusa) and Sawara different coloured foliage and forms1, 12 . The timber is also
cypress (Chamaecyparis pisifera) are the most frequent ones valuable as it has many good qualities: fine texture, straight
after Lawson cypress8-10 . grain, easy to work and resistant to decay, it is suitable for a
wide range of applications including general construction, railway
Habitat and Ecology sleepers, doors, toys, and in the past, arrow shafts and venetian
Lawson cypress prefers medium-textured soils with bind slats12 . Only lack of availability has prevented it being used
consistent summer moisture, but it can also grow in drier more widely commercially12 . Chamaecyparis mature stands may
conditions. It is relatively shade-tolerant and can cope with a offer a good protection from soil erosion and their root systems
wide range of conditions and soil types. It is able to grow either may mitigate shallow-landslide susceptibility13-15 .
under a forest canopy or as a pioneer in the open. Growth rate is
relatively slow for young trees, but older trees retain their ability
Threats and Diseases
to respond to more light and space and can become dominant Lawson cypress is highly susceptible to the oomycete
in old-growth forests. It is usually found in mixed coniferous Phytophthora lateralis that has spread throughout much of its
forests (fir, spruce, pine), or with broadleaved species such as range, causing heavy losses since first being described in 19231 .
oak11 . It is an interesting species ecologically as its natural range The pathogen causes root rot and can quickly kill trees of all
is extremely small, yet it is able to survive in a wide variety of ages. This has resulted in Lawson cypress now being classed
conditions4, 12 . as near threatened in the United States. The pathogen has
more recently been observed in Europe where it now poses an
increasing threat16, 17.
References
[1] A. Praciak, et al., The CABI encyclopedia of [9] GBIF Secretariat, GBIF Backbone
forest trees (CABI, Oxfordshire, UK, 2013). Taxonomy (Global Biodiversity Information
[2] R. J. Uchytil, Chamaecyparis lawsoniana. Facility (GBIF), 2014), chap. GBIF-
Fire Effects Information System (1990). ID:2683880.
http://www.feis-crs.org/feis [10] GBIF Secretariat, GBIF Backbone
[3] A.F. Mitchell, P.Dahlstrom, E.Sunesen, Taxonomy (Global Biodiversity Information
C.Darter, A field guide to the trees of Facility (GBIF), 2014), chap. GBIF-
Britain and northern Europe (Collins, ID:2683866.
1974). [11] A.Farjon, A handbook of the worlds
[4] D.B. Zobel, L.F. Roth, G.M. Hawk, Ecology, conifers (Brill, 2010).
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Orford-cedar (Chamaecyparis lawsoniana), (Chamaecyparis lawsoniana (A. Murr.)
Tech. rep., United States Department of Parl.) An American Wood, Tech. rep., Forest
Agriculture, Pacific Northwest Forest and Service (1984).
Range Experiment Station (1985). [13] H.Kitahara, Y.Okura, T.Sammori,
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(Collins, 2006). Landscape Research 82, 79 (2009).
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Taxonomy (Global Biodiversity Information (2013).
Facility (GBIF), 2014), chap. GBIF- [17] C.Robin, etal., Forest Pathology 41, 417
ID:2683849. (2011).
The fleshy fruits of Cornelian cherry are edible and used for a
variey of culinary purposes.
Annual average temperature (C) Potential spring-summer solar irradiation (kWh m-2) Seasonal variation of monthly precipitation (dimensionless) (Copyright Marion Schneider & Christoph Aistleitner, commons.wikimedia.org: PD)
Cluster of hermaphrodite flowers composed of 4 yellow stamens and petals and 4 greenish sepals.
(Copyright AnRo0002, commons.wikimedia.org: CC0)
For this purpose several cultivars have been selected, bearing blossoming in late winter, its plasticity in growing in different
fruits with different sizes, taste (acidity and sweet) and colours kinds of soils and its trimming tolerance, make this species
(from creamy white, yellow, orange, red, violet to black); e.g. well adapted for use in gardens and parks as an ornamental,
the Macrocarpa variety bears large and pear-shaped fruits, nectariferous, hedge and shade plant4, 5, 16 . As for fruits, many
Alba white fruits and Flava yellow and sweeter fruits4 . In Asia, cultivars are available in trade all over the world with several
Cornelian cherries are also made into an alcoholic beverage, habits, colours of leaves and flowering abundance; e.g. Nana
Floral bud blossoming in early spring.
similar to the Drenja beverage produced in Serbia22 . Fruits plump and growing in dwarf stature, Elegantissima with leaves (Copyright Franco Rossi, www.actaplantarum.org: AP)
are rich in tannins and sugars as well as phenols, ascorbic acid, with yellow or pink margins, Variegata with white margins26 .
flavonoids and anthocyanins23, 24 . Traditionally the fruits were
used as a diuretic, analgesic and tonic. In Middle Age cornelian Threats and Diseases References
[1] S.Pignatti, Flora dItalia (Edagricole, [14] M.M. Markle, American Journal of
cherry shrubs were commonly planted in monastery gardens4 . The cornelian cherry has been apparently free of disease and Bologna., 1982). Archaeology 81, 323 (1977).
Recently its therapeutic properties have been well documented, pest problems, a highly appreciated characteristic which gained [2] M.J. Saint-Hilare, Treatise on Trees [15] I.Brmaud, etal., Annals of Forest Science
and Shrubs Grown in France and in the 69, 373 (2012).
finding high antioxidant and anti-inflammatory properties with its use as an ornamental and orchard species. However, recently Countryside - Version translated by J [16] J.H. Wiersema, B.Len, World Economic
B Fleishman (Firmin Didot Press, Paris,
beneficial effects on cardiovascular, endocrine, gastrointestinal the first diseases have been reported, principally in nurseries and 1825).
Plants: A Standard Reference (CRC Press,
2013).
and immune systems22, 24, 25 . The abundance of flowers plantations, such as the leaf spot and fruit rot disease caused by [3] V.Drken, Jahrbuch des Bochumer [17] S.Ercsl, Journal of Fruit and Ornamental
Botanischen Vereins 1, 213 (2010). Plant Research 12, 87 (2004).
[4] L.Reich, Arnoldia 56, 2 (1996). [18] N.Mamedov, L.E. Craker, Acta
[5] B.K. Shishkin, etal., Flora of the USSR - Horticulturae 629, 83 (2004).
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Jerusalem, 1970). 818, 85 (2009).
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Chorologie der Zentraleuropischen Flora African Journal of Biotechnology 9, 1205
- Band I, II, III (Gustav Fischer Verlag, (2010).
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[21] H.Hassanpour, Y.Hamidoghli,
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Norwegens. Ein Beitrag zur Natur- and Ecology 48, 257 (2013).
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(Christiania, A.W. Brgger, 1975). [22] B.J. West, S.Deng, C.J. Jensen, A.K. Palu,
L.F. Berrio, International Journal of Food
[8] C.S. Sargent, Manual of the Trees of Science & Technology 47, 1392 (2012).
North America (exclusive of Mexico), vol.2
(Dover Publications, New York, 1961), [23] E.Mratini, M.F. Aki, V.Rakonjac,
second edn. R.Mileti, M.iki, Plant Systematics and
Evolution 301, 365 (2015).
[9] M.T. Mmbaga, E.C. Nnodu, HortScience
41, 721 (2006). [24] P.Mikaili, etal., Journal of pharmaceutical
and biomedical sciences 35, 1732 (2013).
[10] J.Q. Xiang, D.E. Boufford, Flora of China,
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Ericaceae, F.ofChina EditorialCommittee, Research and Essays 6, 98 (2011).
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760, 433 (2007). Phytopathology And Plant Protection 46,
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[29] S.Klimenko, Journal of Fruit and
[13] R.E. Weaver, Arnoldia 36, 50 (1976). Ornamental Plant Research 12, 93 (2004).
Description
Cornus sanguinea L., known as common or red dogwood, is Frequency
a deciduous shrub, which usually grows 3-4m in height, but it < 25%
25% - 50%
can develop into a small tree reaching 6m especially in southern 50% - 75%
> 75%
ranges1 . Young slender twigs, especially those exposed to sun, Chorology
are dark red and when crushed the bark has a characteristic Native
Flowers have 4 creamy white petals and 4 stamens and are arranged in
Annual precipitation (mm)
flat-topped inflorescences.
(Copyright Stefano Zerauschek, www.flickr.com: AP)
Bog in mire pine forest of the emeri National Park (Jrmala, Latvia).
(Copyright Jevgenij Voronov, unsplash.com: CC0)
The leaves are deciduous, rounded, 6-12cm long, with a double 25% - 50%
50% - 75%
serrate margin and hairy on both sides. The flowers appear in early > 75%
Chorology
spring, before the leaves, and are monoecious with single-sex Native
wind-pollinated catkins. Male catkins are usually grouped together
(2-4 flowers) and are yellowish-brown and up to 10cm long, while
female catkins are very small. During flowering, the inflorescence
becomes slender and doubles in length. The fruit is a nut, grouped
in clusters of one to four together. Each nut is held in a short leafy
involucre (husk) which encloses about half of the nut. The nut is
roughly spherical, up to 2cm long, yellow-brown with a pale scar
at the base. Hairy leaves with double-serrated margins.
The average life span of this species is about 80-90 years1 . (Copyright Daniele de Rigo: CC-BY)
Uncertain, no-data
Threats and Diseases
In Europe, the pathogens Pseudomonas avellana and P.
Marginal/no presence < 5%
syringae pv. coryli are responsible for the bacterial canker
Low presence 5% - 10% and decline of hazelnut25, 26 . The disease is characterised by a
Mid-low presence 10% - 30% sudden wilting of the twigs and branches, especially at the end
of spring and during the summer27. Moreover, in some European
Medium presence 30% - 50%
countries such as Spain or Poland, hazelnut is affected by the
Mid-high presence 50% - 70% Apple mosaic virus28, 29 . In North America, it was reported that
High presence 70% - 90% the main diseases are caused by Anisogramma anomala30, 31 .
The nuts could be affected by Fusarium lateritium, which is the
Very-high presence > 90%
causal agent of nut grey necrosis32 .
Map 2: High resolution distribution map estimating the relative probability of presence. Maturing nuts can be up to 2 cm long when ripe.
(Copyright Ettore Balocchi, www.flickr.com: CC-BY)
Uncertain, no-data
Tundra, cold desert
Negligible survivability
Low survivability
Mid-low survivability
Medium survivability
Mid-high survivability
High survivability
Ancient hazelnut coppice in Galloway, UK. Map 3: High resolution map estimating the maximum habitat suitability.
(Forestry Commission, www.forestry.gov.uk: Crown Copyright)
References
[1] F. Clinovschi, Dendrologie (Editura [20] S. B. Vander Wall, The Botanical Review
Universitatii Suceava, 2005). 67, 74 (2001).
[2] C. Contesa, N. Valentini, M. Caviglione, R. [21] G. Rodolfi, Acta Theriologica 39, 215
Botta, European Journal of Horticultural (1994).
Science 76, 170 (2011). [22] Y. Wang, et al., Journal of Biochemistry
[3] W. Tinner, P. Hubschmid, M. Wehrli, B. and Molecular Biology 40, 861 (2007).
Ammann, M. Conedera, Journal of Ecology [23] F. Bestoso, et al., BMC Biotechnology 6,
87, 273 (1999). 45+ (2006).
[4] V. Erdogan, S. A. Mehlenbacher, Journal [24] S. A. Mehlenbacher, D. C. Smith,
of the American Society for Horticultural HortScience 41, 482 (2006).
Science 125, 489 (2000).
[25] M. Scortichini, et al., Phytopathology 95,
[5] S. Mehlenbacher, Acta Horticulturae 290, 1316 (2005).
791 (1991).
[26] M. Scortichini, Plant Disease 86, 704
[6] K. Foetisch, et al., Biochemical Journal (2002).
383, 327+ (2004).
[27] M. Scortichini, U. Marchesi, M. P. Rossi, P.
[7] N. Nikolaieva, J. Brindza, K. Garkava, R. Di Prospero, Applied and Environmental
Ostrovsky, Modern Phytomorphology 6, Microbiology 68, 476 (2002).
53 (2014).
[28] J. Aramburu, M. Rovira, Plant Pathology
[8] A. E. Palm, G. G. Vendramin, Molecular 49, 423 (2000).
Ecology 11, 1769 (2002).
[29] T. Kobylko, B. Nowak, A. Urban, Folia
[9] J. Deacon, New Phytologist 73, 1055 Horticulturae 17, 153 (2005).
(1974).
[30] K. B. Johnson, S. A. Mehlenbacher, J. K.
[10] P. Boccacci, et al., Tree Genetics & Stone, J. W. Pscheidt, J. N. Pinkerton, Plant
Genomes 9, 1465 (2013). Disease 80, 1308 (1996).
[11] K. E. Hummer, Acta Horticulturae 556, [31] S. A. Mehlenbacher, J. N. Pinkerton, K. B.
25 (2001). Johnson, J. W. Pscheidt, Acta Horticulturae
[12] L. Braun, J. Gillman, E. Hoover, M. Russelle, 351, 551 (1994).
Canadian Journal of Plant Science 91, [32] S. Vitale, et al., Phytopathology 101, 679
773 (2011). (2011).
[13] O. Kull, U. Niinemets, Tree Physiology 12, [33] H. Meusel, E. Jger, eds., Vergleichende
311 (1993). Chorologie der Zentraleuropischen Flora
[14] R. Catoni, M. U. Granata, F. Sartori, L. - Band I, II, III (Gustav Fischer Verlag,
Varone, L. Gratani, Photosynthetica 53, Jena, 1998).
Typical shrub-form plant in a garden park (Lake Maggiore, North Italy). 35 (2015). [34] J. Jalas, J. Suominen, Atlas Florae
(Copyright Daniele de Rigo: CC-BY) [15] P. S. Savill, The silviculture of trees used in Europaeae: distribution of vascular
British forestry (CABI, 2013). plants in Europe Vol. 3 Salicaceae to
[16] B. Ustaoglu, M. Karaca, Applied Ecology Balanophoraceae (Committee for Mapping
Autoecology diagrams based on harmonised the Flora of Europe and Societas Biologica
Field data in Europe (including absences) Observed presences in Europe field observations from forest plots. and Environmental Research 12, 309
Fennica Vanario, Helsinki, 1976).
(2014).
[17] L. Kubiak-Martens, Vegetation History and [35] Sociedade Portuguesa de Botnica, Flora-
Archaeobotany 8, 117 (1999). On: Flora de portugal interactiva (2014).
http://www.flora-on.pt.
[18] A. lhami Kksal, N. Artik, A. imek, N.
Average temperature of the coldest month (C)
Gne, Food Chemistry 99, 509 (2006). [36] O. de Bols, J. Vigo, Flora dels pasos
catalans, vol I-IV (Barcino, Barcelona,
[19] Food and Agriculture Organization of 1984-2001).
the United Nations, FAOSTAT (Food and
Annual precipitation (mm)
Map 1: Plot distribution and simplified chorology map for Cupressus sempervirens.
Frequency of Cupressus sempervirens occurrences within the field
observations as reported by the National Forest Inventories. The chorology
of the native and introduced spatial range for C. sempervirens is derived
after Faini andDellaRocca16 .
Distribution
The natural distribution of this cypress is unclear, due to its
long horticultural history in the Mediterranean region2, 8 . Natural
stands occur in the south-western Mediterranean basin over
several geographically non-adjacent areas reaching eastwards
the Caucasus and south-western Iran, from sea level (Crete)
up to 2000m (Turkey)7. Various authorities attribute its native One-year old ovoid-shaped seed cones: they take two years to reach maturity.
distribution to the Aegean islands (Crete, Samos, Rhodes, Kos (Copyright Alan Gregg, www.flickr.com: CC-BY)
and Symi), Cyprus, Turkey, Middle East (Syria, Jordan, Lebanon Importance and Usage
and Iran), and in North-East Africa (Libya, Tunisia)3, 7, 9 , although
The Mediterranean cypress has a long history of exploitation of
recent studies on genetic and paleobotanic records presume
its natural forests, leading to their strong decline. On the other
the presence of central Mediterranean natural populations10 .
hand it has been widespread planted throughout the
Today the distribution of this species, principally the columnar
Mediterranean area, and other regions, for ornamental and
form, comprises most of the Mediterranean basin and Middle
religious purposes7, 9. Since ancient times its wood has been
East, favoured by human cultivations since the time of ancient
particularly appreciated for its resistance to fungi and insects,
civilisations, and more recently all over the world as an
especially if immersed in water7. The wood is suitable for small
ornamental tree3, 7. Some populations are recognised as separate
carpentry, exterior woodworks (doors, windows, garden furniture,
varietals and for some authors are treated as different species:
etc.) and ship building. The insect repellent odour makes the wood
Cupressus sempervirens var. numidica in Tunisia and Cupressus
suitable for chests and wardrobes to store linens and foods. It is
sempervirens var. indica in northern Iran10 . The endemic
also used for coffins6. The columnar or conical forms have been
Moroccan cypresses in the High Atlas Mountains are considered
used since Greek and Roman times as an ornamental tree for
as a separate species (Cupressuss altantica)8, 11 , but some
shading gardens, cemeteries, as a windbreak along roads, and it
authors classify it as varietal of the common cypress (Cupressus
has become a characteristic feature of the Mediterranean coastal
sempervirens var. atlantica)2 or of the Tassili cypress (Cupressuss
and urban landscapes7, 9, 10. More recently, thanks to its ecological
dupreziana var. atlantica)1, 10 .
qualities, this cypress has been used in forest protection against
Habitat and Ecology desertification and soil conservation in hot areas, where the soil is
shallow and degraded and no other forest tree species could
In natural habitats this cypress occurs in Mediterranean
grow16, 17. Its deep and dense litter and the crown are difficult to
Tall ornamental cipress with columnar habit in an urban garden climates with dry and hot summers and rainy winters, or in semi-
(Udine, North-East Italy). ignite, so it can be used as firebreak18, even if regeneration is
(Copyright Graziano Propetto, www.actaplantarum.org: AP) arid climates in the eastern and interior areas of its range1 . It is
scarce after wildfires12. Mediterranean cypress also tolerates salty
winds, so it is used as coastal windbreak. It has a good resistance
Autoecology diagrams based on harmonised
Field data in Europe (including absences) Observed presences in Europe field observations from forest plots.
to frequent frost damage, trimming and grazing, as it is able to re-
sprout quickly, thus is also suitable as evergreen hedge6, 7, 12, 17.
References
[1] A. Farjon, A handbook of the worlds [12] R. Del Favero, I boschi delle regioni
conifers (Brill, Leiden, 2010). dellItalia centrale (Cleup, Padova, 2010).
[2] J. E. Eckenwalder, Conifers of the World: [13] A. Santini, V. Donardo, Forest Pathology
The Complete Reference (Timber Press, 30, 87 (2000).
2009). [14] M. C. Vermigli, Genetic variability in italian
[3] A. Farjon, D. Filer, An Atlas of the Worlds populations of Cupressus sempervirens L.
Conifers: An Analysis of their Distribution, assessed by SSR and RAPD markers, Ph.D.
Biogeography, Diversity and Conservation thesis (2005).
Status (Brill, 2013). [15] MonumentalTrees.com, Monumental trees
[4] A. F. Mitchell, A field guide to the trees (2015).
of Britain and northern Europe (Collins, [16] A. Faini, G. Della Rocca, Gestione
1974). selvicolturale dei boschi di cipresso in
[5] O. Johnson, D. More, Collins tree guide aree naturali e naturalizzate (Centro
(Collins, 2006). Promozione Pubblicit, Firenze, 2004),
[6] A. Praciak, et al., The CABI encyclopedia of pp. 2328.
forest trees (CABI, Oxfordshire, UK, 2013). [17] G. Della Rocca, Il valore economico del
[7] M. Intini, G. Della Rocca, Il cipresso cipresso: paesaggio e ambiente (Centro
comune (Cupressus sempervirens L.): Promozione Pubblicit, Firenze, 2004),
caratteristiche botaniche, distribuzione, pp. 8890.
ecologia (Centro Promozione Pubblicit, [18] G. Della Rocca, et al., Journal of
Firenze, 2004), pp. 1322. Environmental Management 159, 68
[8] C. J. Earle, The gymnosperm database (2015).
(2015). http://www.conifers.org [19] G. Della Rocca, C. A. Eyre, R. Danti,
[9] A. Farjon, The IUCN Red List of Threatened M. Garbelotto, Phytopathology 101,
Species (2013), pp. 32518/0+. 1408 (2011).
[10] F. Bagnoli, et al., Molecular Ecology 18, [20] Z. Solel, Z. Madar, M. Kimchi, Y. Golan,
2276 (2009). Canadian Journal of Plant Pathology 9,
115 (1987).
[11] K. Rushforth, R. P. Adams, M. Zhong, Ma, R.
N. Pandey, Biochemical Systematics and
Ecology 31, 17 (2003).
Eucalyptus globulus and other eucalypts in Europe: distribution, habitat, usage and threats
S. Cerasoli, M. C. Caldeira, J. S. Pereira, G. Caudullo, D. de Rigo
The Eucalyptus globulus Labill., commonly named as Tasmanian blue gum, is an evergreen broadleaf tree native to
south-eastern Australia. In Europe it is mainly cultivated in the Iberian Peninsula for paper pulp production, managed
as short rotation coppice stands. It is appreciated for its adaptation and fast-growing aptitude. Rapid environmental
changes in the Mediterranean region menace Tasmanian blue gum trees increasing the risk of fire, drought and the
outbreaks of pests and diseases.
The Tasmanian blue gum (Eucalyptus globulus Labill.) is a
medium to large evergreen broadleaf tree, growing up to 70 m, and Frequency
is the tallest tree species recorded in Europe1 . The bark is smooth < 25%
25% - 50%
and shed yearly in long ribbons when a new layer of the outermost 50% - 75%
tissue is formed. Juvenile leaves are ovate, thin, sub-horizontal and > 75%
covered with a blue grey wax bloom that gives rise to the common
name of the species. The adult leaves are lanceolate and shift to
vertical hanging. The leaves have roughly circular glands containing
aromatic oils. Flowers are solitary and are formed in the axils of
leaves. The sepals and petals are united to form a cap (operculum)
that covers stamens and that drops off at anthesis2 . The name
Eucalyptus originates from this trait common to the entire genus,
the words eu and kalyptos, meaning well and covered respectively2 .
The fruit is a woody capsule 0.6 to 2.5cm in diameter3.
Distribution
The Tasmanian blue gum, a native tree of south-eastern
Australia (Tasmania and southern Victoria4), belongs to the
angiosperm family of Myrtaceae, which includes more than 800
Map 1: Plot distribution for Eucalyptus spp.
species2 . It was introduced in south-western Europe (Portugal,
Frequency of Eucalyptus spp. occurrences within the field observations as Bark sheds in long ribbons.
Spain) and Northern Africa in the mid 19th century and was planted reported by the National Forest Inventories. (Copyright J.R. Pinho: CC-BY)
for industrial purposes, mainly timber and paper pulp3 . Nowadays
it is the major pulpwood species planted in temperate regions of
the world5 . In Europe, it covers 1.3 million hectares of forested
area, mainly in the Iberian Peninsula (more than 80%), France and
Italy6 . In Portugal, the Tasmanian blue gum plantations are at their
best in the northern half of the littoral, where winters are mild
and precipitation higher (700-2000mm) than in the Southern and
inland regions of the country7. In Spain, eucalypt plantations are
mainly in the North Western region (Galicia)8 .
Adult leaves are lanceolate and contain glands with aromatic oils.
(Copyright Forest & Kim Starr, commons.wikimedia.org: CC-BY)
young twigs are green, 4-angled, without brown protuberances. 25% - 50%
50% - 75%
New shoots grow vigorously, also 4-angled and winged. The > 75%
Chorology
buds are 2-4mm long, ovoid, sharply pointed. The leaves are Native
simple, opposite, lanceolate or ovate-elliptical, 3-10 cm long
and 2-3.5cm wide, narrowing at tip and base. The leaf margins
are crenate and finely saw-toothed and the leaves have a rough
surface and are bluish-green beneath. The petioles are 0.5-1cm
long1-5 . This species is gynodioecious, having female flowers on
some individual plants and hermaphrodite flowers on others1 . Fleshy orange fruit ripening inside the four-valved red capsule.
(Copyright Aldo De Bastiani, www.actaplantarum.org: AP)
The flowers are small, delicate and about 1cm in diameter. They
are arranged in inflorescences of 3-10 flowers in leaf axils, on the current populations appear stable1, 17-19 . This species can be a
1-3.5cm long pedicels, having 4 elements of each of the floral host of other diseases, such as a strain of cucumber mosaic virus,
whorls (sepals, petals, stamens and carpels). They blossom in April- strawberry latent ringspot and in some countries, a strain of cherry
July4, 6 . The fruits are capsules, 1-1.5cm wide, with 4 angled lobes, leaf roll virus1 . Despite its toxicity, scales and aphids can adapt to
green then dark pink or red when mature. Ripe fruits open through survive its otherwise insecticidal chemistry1, 10, 17-19 .
4 valves, containing 4 whitish seeds covered by a fleshy red-orange
layer (pseudo-aril). Fruits ripen in September-October2-5 . Map 1: Plot distribution and simplified chorology map for Euonymus europaeus.
Frequency of Euonymus europaeus occurrences within the field observations
as reported by the National Forest Inventories. The chorology of the native
Distribution spatial range for E. europaeus is derived after several sources7, 20-25 .
The spindle tree is present in temperate climates, from
Central to Eastern Europe (except the extreme north and most of effects to foxglove (Digitalis spp.). Dried, powdered fruits and seeds
the Mediterranean area), and reaches eastwards the Urals and mixed in butter were used to deter lice4 . Antifungal chitin-binding
Caucasus2-5, 7. It grows generally at low to middle elevations, from proteins have been isolated from bark and leaves14-16 . The whole
sea level to 1300m of altitude in Sicily6, 8 . It has been planted plant contains compounds of medicinal and veterinary value1, 10, 11 .
outside its natural range, now naturalised in some areas (e.g.
Scandinavia). Exported also to other continents, in northeast United
Threats and Diseases
There are no serious threats for this species1 . The spindle tree
States and in New Zealand it is considered an invasive species1 .
is the primary overwintering host of the black bean aphid (Aphis
Habitat and Ecology fabae) which feeds on field beans (Vicia faba) and sugar beet (Beta
It inhabits mainly forest margins, clearings and open vulgaris) and the peach potato aphid/green peach aphid (Myzus
woodlands, preferring medium moisture levels, and well-drained, persicae), a widespread pest of a large number of crops1, 17-19 . As
preferably alkaline soils in full to partial shade1 . It is frequently a measure against black bean aphid, in the past spindle trees have
found as an understorey shrub, principally in mixed mesophilous been removed from hedges and woodlands (e.g. England), although
forests dominated by deciduous oaks (Quercus robur, Quercus
petraea) and hornbeam (Carpinus betulus), forming oak-
hornbeam forest communities, along with ash (Fraxinus excelsior),
field maple (Acer campestre), etc.9 . The pseudo-aril is eaten by
a range of animals and its seeds are widely dispersed mainly by Blossoming hermaphrodite flower with 4 green petals,
sepals and 4 yellow stamens.
birds (robins, blackbirds, blackcap, and song thrush) and rodents, (Copyright Franco Rossi, www.actaplantarum.org: AP)
but the seeds are poisonous1, 10, 11 .
References
Importance and Usage [1] P. A. Thomas, M. El-Barghathi, A. Polwart, [14] K. P. Bergh, et al., Planta 219, 221 (2004).
Journal of Ecology 99, 345 (2011). [15] N. van der Weerden, M. Bleackley, M.
[2] T. G. Tutin, Flora Europea. Volume 2. Anderson 70, 3545 (2013).
The spindle tree is used mainly as an ornamental shrub for Rosaceae to Umbelliferae, T. G. Tutin, [16] A. J. De Lucca, T. E. Cleveland, D. E. Wedge,
its impressive autumn display of orange, red and purple leaves, et al., eds. (Cambridge University Press, Canadian Journal of Microbiology 51,
1968), p. 242. 1001 (2005).
accented by magenta pink to red fruits with orange to red pseudo- [3] O. Polunin, Flowers of Europe: A Field [17] M. J. Way, M. E. Cammell, Journal of
arils. For example, the cultivar Red Cascade in autumn has red Guide (Oxford University Press, 1969). Applied Ecology 19, 929 (1982).
[4] T. Svulescu, ed., Flora Republicii [18] M. E. Cammell, G. M. Tatchell, I. P. Woiwod,
leaves and abundant red fruits with rose pseudo-arils12 . The wood Populare Romne, vol 6 (Editura Journal of Applied Ecology 26, 463
Academiei Romne, Bucureti, 1958).
of the spindle is homogenous, white or yellowish, and easy to work. (1989).
[5] O. Johnson, D. More, Collins tree guide [19] G. Powell, C. R. Tosh, J. Hardie, Annual
It is used, more in the past, for plywood and toothpicks4, knitting (Collins, 2006). Review of Entomology 51, 309 (2006).
needles, combs, and for making spindles, from which the common [6] S. Pignatti, Flora dItalia (Edagricole, [20] Tela Botanica, eFlore (2015).
Bologna., 1982). http://www.tela-botanica.org
name derives13; the wood is also heat resistant and it was used in [7] E. Hultn, M. Fries, Atlas of North [21] Anthos, Information System of the plants
making tobacco-pipes13. Charcoal from its wood is used for drawing, European vascular plants (North of of Spain (Real Jardn Botnico, CSIC -
the Tropic of Cancer), Vols. I-III. (Koeltz Fundacin Biodiversidad, 2015).
and to make charcoal powder. The red pseudo-aril was also used scientific books, 1986). http://www.anthos.es.
in the past to make dyes4 . The bark, the leaves and the seeds were Shrub form of spindle in late autumn in a rural area along the Upper Rhine [8] H. Meusel, E. Jger, eds., Vergleichende [22] S. I. Sokolov, O. A. Svjaseva, V. A. Kubli,
valley (Hockenheim, South-West Germany). Chorologie der Zentraleuropischen Flora Distribution ranges of trees and shrubs
used as a purgative, but they are toxic, with similar cardio-stimulant (Copyright AnRo0002, commons.wikimedia.org: CC0) - Band I, II, III (Gustav Fischer Verlag, in the USSR, vol 1-3 (Nauka, Leningrad,
Jena, 1998). 1977-1986). (In Russian).
[9] H. H. Ellenberg, Vegetation Ecology of [23] Sociedade Portuguesa de Botnica, Flora-
Autoecology diagrams based on harmonised Central Europe (Cambridge University On: Flora de portugal interactiva (2015).
Field data in Europe (including absences) Observed presences in Europe Press, 2009), fourth edn. http://www.flora-on.pt
field observations from forest plots.
[10] R. Nishida, Annual Review of Entomology [24] Bundesamtes fr Naturschutz, ed.,
47, 57 (2002). FloraWeb (2015). http://www.floraweb.de.
[11] P. Vichova, L. Jahodar, Human & [25] P. H. Davis, Flora of Turkey and the
Experimental Toxicology 22, 467 (2003).
Average temperature of the coldest month (C)
East and north faces of the Matterhorn from the Domhtte (Valais, Switzerland).
(Copyright Sven Scheuermeier, unsplash.com: CC0)
Meadows and spruce stands in summer in Badia Valley (Bolzano, North-East Italy).
(Copyright Giuseppe Milo, www.flickr.com: CC-BY)
Shiny dark green leaves with red galls caused by the fly Mikiola fagi
Map 2: High resolution distribution map estimating the relative probability of presence. (Diptera Cecidomyiidae).
(Copyright AnRo0002, commons.wikimedia.org: CC0)
Fagus orientalis
Fagus orientalis, or oriental beech, is closely related to Fagus sylvatica.
Some authorities consider them to be sub-species; others consider Uncertain, no-data
them to be two separate species1 . In appearance they are generally Tundra, cold desert
very similar. The leaves are slightly longer, darker and less glossy than
Negligible survivability
those of European beech, and tend to have more vein-pairs (9-14 as
opposed to 5-9)3 . Oriental beech can be found in the Balkans, Anatolia, Low survivability
the Caucasus, northern Iran and Crimea18 . Its range overlaps with that Mid-low survivability
of the European beech and there is frequently hybridisation between the
Medium survivability
two18 . Where both species are present, oriental beech tends to favour
the valleys while European beech is found further up the slopes; this is Mid-high survivability
because the European beech is more susceptible to late frosts12 . High survivability
References
[1] J . R. Packham, P. A. Thomas, M. D. [17] J. E. Norris, A. Di Iorio, A. Stokes, B. C.
Atkinson, T. Degen, Journal of Ecology Nicoll, A. Achim, Slope Stability and
100, 1557 (2012). Erosion Control: Ecotechnological
[2] A. F. Mitchell, A field guide to the trees Solutions, J. E. Norris, et al., eds. (Springer
of Britain and northern Europe (Collins, Netherlands, 2008), pp. 167210.
1974). [18] L. K. A. Dorren, F. Berger, C. le Hir, E.
[3] O. Johnson, D. More, Collins tree guide Mermin, P. Tardif, Forest Ecology and
(Collins, 2006). Management 215, 183 (2005).
[4] J. Fang, M. J. Lechowicz, Journal of [19] J. I. Barredo, et al., EPPO Bulletin 45, 273
Biogeography 33, 1804 (2006). (2015).
[5] R. Baker, et al., EFSA Journal 9, 2186+ [20] CABI, Hylobius abietis (large pine weevil)
(2011). 108 pp. (2015). Invasive Species Compendium.
http://www.cabi.org
[6] E. Hultn, M. Fries, Atlas of North European
vascular plants (North of the Tropic of [21] M. Lf, Forest Ecology and Management
Cancer), Vols. I-III. (Koeltz scientific books, 134, 111 (2000).
1986). [22] M. Lf, G. Isacsson, D. Rydberg, T.
[7] D. Magri, Journal of Biogeography 35, N. Welander, Forest Ecology and
450 (2008). Management 190, 281 (2004).
[8] T. Horgan, et al., A guide to forest tree [23] G. Kandemir, Z. Kaya, EUFORGEN Technical
species selection and silviculture in Ireland. guidelines for genetic conservation and
(National Council for Forest Research and use for Oriental beech (Fagus orientalis)
Development (COFORD), 2003). (Bioversity International, Rome, Italy, 2009).
[9] K. Kramer, et al., Forest Ecology and [24] A. Oprea, C. Srbu, I. Goia, Contributii
Management 259, 2213 (2010). The Botanice 46, 17 (2011).
ecology and silviculture of beech: from [25] M. ijai Nikoli, J. Milovanovi, M. Noni,
gene to landscape. R. Kneevi, D. Stankovi, Genetika 45,
[10] A. Praciak, et al., The CABI encyclopedia of 369 (2013).
forest trees (CABI, Oxfordshire, UK, 2013). [26] T. Denk, G. Grimm, K. Stgerer, M. Langer,
[11] L. Walthert, E. Graf Pannatier, E. S. Meier, V. Hemleben, Plant Systematics and
Mature beech forest with autumn colour foliage in Delamere forest, Cheshire, UK. Evolution 232, 213 (2002).
(Forestry Commission, www.forestry.gov.uk: Crown Copyright) Forest Ecology and Management 297,
94 (2013). [27] H. Meusel, E. J. Jger, Plant Systematics
[12] L. Augusto, J. Ranger, D. Binkley, A. Rothe, and Evolution 162, 315 (1989).
Autoecology diagrams based on harmonised Annals of Forest Science 59, 233 (2002). [28] EUFORGEN, Distribution map of beech
Field data in Europe (including absences) Observed presences in Europe field observations from forest plots. (Fagus sylvatica) (2008).
[13] L. Paule, Forest Genetics 2, 161 (1995).
www.euforgen.org.
[14] A. Geler, et al., Trees - Structure and
Function 21, 1 (2007). [29] J. Jalas, J. Suominen, Atlas Florae Europaeae:
distribution of vascular plants in Europe Vol. 3
[15] A. Granier, et al., Agricultural and Forest Salicaceae to Balanophoraceae (Committee
Average temperature of the coldest month (C)
Meteorology 143, 123 (2007). for Mapping the Flora of Europe and Societas
[16] M. Goldstein, G. Simonetti, M. Watschinger, Biologica Fennica Vanario, Helsinki, 1976).
Alberi dEuropa (A. Mondadori, 1995). [30] K. Browicks, J. Zieliski, Chorology of
Annual precipitation (mm)
Distribution
Its distribution covers the central-southern Europe and
northwest Africa, up to the Caucasus5, 6 . Due to its large
morphological variations, this ash species includes a complex of
taxa and its taxonomic status is still not clear. However, prevailing Map 1: Plot distribution and simplified chorology map for Fraxinus angustifolia.
Frequency of Fraxinus angustifolia occurrences within the field observations
opinion recognises three geographical subspecies on the basis as reported by the National Forest Inventories. The chorology of the native
of molecular and morphological data: the narrow-leaved ash spatial range for F. angustifolia is derived after Meusel andJger5 .
(Fraxinus angustifolia subsp. angustifolia) in south-western
Europe and north-western Africa, the Caucasian ash (Fraxinus large rivers, where it formed vast and continuous populations,
angustifolia subsp. oxycarpa) in central Europe, Balkans and now with more limited extent. In the Mediterranean regions its
the Black Sea region, and the Syrian ash (Fraxinus angustifolia distribution is more patchy and reduced to smaller and more
subsp. syriaca) in south-east Anatolia, Middle East to Iran7. The isolated populations along rivers or on wetland sites, but also on
northern part of the distribution overlaps with that of common drier sites at higher altitudes. Rarely, it creates pure stands, found
ash (Fraxinus excelsior), with which it can naturally hybridise, only in optimum conditions as a pioneer species. More often this
developing plants with intermediate trait forms8, 9 . Exported as an ash is a dominant or a secondary species in mixed broadleaved
Mature dry samaras at the end of summer.
ornamental tree, this ash is naturalised in southern Australia10 . forests with oak (Quercus spp.), elm (Ulmus spp.), maple (Acer (Copyright Franco Rossi, www.actaplantarum.org: AP)
spp.), willow (Salix spp.), poplar (Populus spp.) and lime (Tilia
spp.), forming stable and species-rich ecosystems3, 4, 13 .
References
Importance and Usage [1] A. F. Mitchell, A field guide to the trees
of Britain and northern Europe (Collins,
[13] H. G. Kutbay, M. Kilin, A. Kandemir, Turkish
Journal of Botany 22, 157 (1998).
1974).
The wood of narrow-leaved ash has similar properties to [14] J. Coello, et al., Ecology and silviculture of
[2] O. Johnson, D. More, Collins tree guide the main valuable broadleaved species
the common ash, although the quality is inferior in terms of (Collins, 2006). in the Pyrenean area and neighbouring
regions (Centre de la Propietat Forestal,
strength and elasticity3 . Timber plantations are not very common [3] D. Boshier, et al., Ash species in Europe:
Generalitat de Catalunya, Santa Perptua
biological characteristics and practical
over Europe, as it is mainly planted in combination with other guidelines for sustainable use (Oxford de Mogoda (Spain), 2013).
Forestry Institute, University of Oxford, [15] E. iek, M. Yilmaz, Proceeding of
species14 . Higher wood quality, comparable to common ash, can United Kingdom, 2005). 128 pp. International IUFRO Meeting: Management
be obtained on drier sites where tree growth is slower3 . In the [4] M. Bobinac, S. Andraev, M. ijai Nikoli, of Fast Growing Plantations, 11-13
Periodicum Biologorum 112, 341 (2010). September 2002. Izmit, Turkey (2002),
north-west of Turkey, where narrow-leaved ash is more used in pp. 192202.
[5] H. Meusel, E. Jager, S. Rauschert, E.
fast-growing plantations on swampy lowlands, the wood quality Weinert, Vergleichende Chorologie der [16] U. Byksar, T. Akbulut, C. Guler, N. As,
Zentraleuropischen Flora (Gustav Fischer BioResources 6, 4721 (2011).
is more similar to the poplars and is suitable for pulpwood and Verlag Jena, 1978). [17] G. M. Elmas, African Journal of
bonded wood products, such as plywood, laminated veneer [6] E. Hultn, M. Fries, Atlas of North European Biotechnology 10, 9812 (2011).
vascular plants (North of the Tropic of [18] R. Schicchi, L. Camarda, V. Spadaro, R.
lumber and glued laminated timber15-17. Leaves are palatable Cancer), Vols. I-III. (Koeltz scientific books, Pitonzo, Quaderni di Botanica ambientale
to livestock and in southern Europe this ash was traditionally 1986). e applicata 17, 151 (2006).
[7] E. Wallander, Belgische Dendrologie Belge [19] A. Galati, G. Migliore, C. Scaffidi Saggio,
used as a fodder tree. It is also widely used as an ornamental 2012, 38 (2013). Colture artificiali di piante medicinali -
[8] C. Raquin, S. Brachet, S. Jeandroz, F. Vedel, Produzione di metaboliti secondari nelle
N. Frascaria-Lacoste, Forest Genetics 9, piante medicinali in coltura artificiale, F.
111 (2002). Tognoni, A. Pardossi, A. Mensuali Sodi,
eds. (Aracne editrice, Roma, 2007), pp.
[9] D. D. Hinsinger, M. Gaudeul, A. Couloux, J. 287297.
Bousquet, N. Frascaria-Lacoste, Molecular
Phylogenetics and Evolution 77, 223 [20] T. Kirisits, M. Matlakova, S. Mottinger-
(2014). Kroupa, E. Halmschlager, F. Lakatos, Plant
Pathology 59, 411 (2010).
[10] A. M. Gray, Flora of Tasmania Online, M.
F. Duretto, ed. (Tasmanian Herbarium, [21] J. Schumacher, EPPO Bulletin 41, 7
Shiny green composite leaves with 7-13 slender leaflets. (2011).
Tasmanian Museum & Art Gallery, Hobart,
(Copyright Javier Martin, commons.wikimedia.org: PD)
2009), p. 4. [22] T. Kowalski, Forest Pathology 36, 264
[11] P. R. Grard, et al., Journal of (2006).
Biogeography 40, 835 (2013). [23] A. Turb, et al., Disturbances of EU forests
Habitat and Ecology [12] M. Thomasset, J. F. Fernndez-Manjarrs, caused by biotic agents - final report,
Tech. Rep. KH-32-13-151-EN-N (2011).
G. C. Douglas, N. Frascaria-Lacoste, T.
The narrow-leaved ash is a fast-growing and light-demanding R. Hodkinson, Climate Change, Ecology Final Report prepared for European
and Systematics, T. R. Hodkinson, M. B. Commission (DG ENV).
tree, occurring mostly in temperate mild climates, with an annual Jones, S. Waldren, J. A. N. Parnell, eds. [24] M. Trmolires, J. M. Snchez-Prez, A.
precipitation between 400 and 800mm. It grows well on moist (Cambridge University Press, 2011), pp. Schnitzler, D. Schmitt, Plant Ecology 135,
320344. 59 (1998).
soils, in temporary flooded lowlands, but also on well-drained [25] T. M. Dugdale, T. D. Hunt, D. Clements,
slopes, although there it suffers more competition from other tree Proceedings of the 19th Australasian
Weeds Conference (2014), pp. 190193.
species. It prefers aerated or only moderately compacted soils,
with a pH range between 5 and 83 . It is particularly susceptible This is an extended summary of the chapter. The full version of
to frosts, which damage winter flowers and seeds in spring, this chapter (revised and peer-reviewed) will be published online at
https://w3id.org/mtv/FISE-Comm/v01/e0101d2. The purpose of this
limiting its northern distribution. However, it can survive in areas summary is to provide an accessible dissemination of the related
main topics.
colder than its natural range when planted, although it is difficult
This QR code points to the full online version, where the most
to disperse seeds in these conditions11, 12 . In central Europe, updated content may be freely accessed.
Please, cite as:
the Pannonian Basin and Balkans, narrow-leaved ash occurs Caudullo, G., Houston Durrant, T., 2016. Fraxinus angustifolia in
mainly in the lowlands, in riparian and floodplain forests along Isolated tree in the Spanish countryside during winter. Europe: distribution, habitat, usage and threats. In: San-Miguel-
(Copyright Alfonso San Miguel: CC-BY)
Ayanz, J., de Rigo, D., Caudullo, G., Houston Durrant, T., Mauri, A. (Eds.),
European Atlas of Forest Tree Species. Publ. Off. EU, Luxembourg, pp.
e0101d2+
skin. In central Europe it has been widely used as an ornamental rapidly spread across several US states, primarily through natural
tree along roads and city streets. In many countries it has some dispersal and the transport of firewood and other wood products
ethnic, cultural, and mythological significance6 . containing ash bark. In infected areas, the pest has caused very high
mortality of the North American ash species, particularly green ash
(Fraxinus pennsylvanica), black ash (Fraxinus nigra), and white ash
Threats and Diseases (Fraxinus americana)5, 23, 24 . The bacterium Pseudomonas syringae
Since it was observed on a large scale in Poland in 1992, the subsp. savastanoi pv. fraxini and the fungus Nectria galligena can
ash dieback phenomenon has spread to other countries in eastern, cause cankers on common ash trees, which adversely affect their
Dark purple anthers of ash flowers, which are without petals.
(Copyright David Nicholls, www.naturespot.ork.uk: AP) northern, and central Europe. In many European countries it has economic value in managed stands. The most severe infections
since caused the death of over 90% of all ash trees18, 19. The fungus occur in extreme habitats8, 16, 20.
Hymenoscyphus fraxineus, also known as Chalara fraxinea, a name
Importance and Usage designating its asexual stage, is primarily responsible for this invasive
The elasticity, hardness, and pressure, shock and splintering disease, causing crown dieback, and this fungus is potentially subject References
resistance of common ash wood make it economically highly to expansion in the European temperate oceanic ecological zones20, [1] H. H. Ellenberg, Vegetation Ecology of [13] D. D. Hinsinger, M. Gaudeul, A. Couloux, J.
valuable, and commercially more important than that of the two 21
. It is visible as a reddish discolouration of the bark in the lower Central Europe (Cambridge University Bousquet, N. Frascaria-Lacoste, Molecular
Press, 2009), fourth edn. Phylogenetics and Evolution 77, 223
other native ash species in Europe (Fraxinus angustifolia and portion of the stem, and eventually often kills the tree. In young trees, [2] A. F. Mitchell, A field guide to the trees of (2014).
Fraxinus ornus). The wood is much-used for tool handles and death can occur in the same growing season in which the symptoms Britain and northern Europe (Collins, 1974). [14] A. Plira, M. Heuertz, EUFORGEN technical
[3] O. Johnson, D. More, Collins tree guide guidelines for genetic conservation and
sports equipment, and also in earlier times, before the widespread first become visually noticeable. Even if older trees might resist (Collins, 2006). use for common ash (Fraxinus excelsior),
Tech. rep., Bioversity International (2003).
use of steel, for weapon handles, agricultural implements, longer, they will be greatly weakened and susceptible to other lethal [4] A. Praciak, et al., The CABI encyclopedia of
[15] G. Marigo, J.-P. Peltier, J. Girel, G. Pautou,
forest trees (CABI, Oxfordshire, UK, 2013).
carriages and car and boat frames. Furthermore, its straight grain diseases or pests22 . The emerald ash borer (Agrilus planipennis) is [5] D. Dobrowolska, et al., Forestry 88, 552
Trees 15, 1 (2000).
[16] G. Kerr, Forestry 68, 63 (1995).
and consistency, with sapwood and hardwood differing little, a beetle native to Asia and eastern Russia. While its adults graze on (2011).
[6] D. Boshier, et al., Ash species in Europe: [17] U. Bohn, et al., Karte der natrlichen
make it very valuable for veneers, furniture, and flooring. Stem ash leaves, the emerald ash borer larvae feed on the phloem, which biological characteristics and practical Vegetation Europas; Map of the
Natural Vegetation of Europe
forking is undesirable and can be caused by frost injury, water can kill the tree. It was observed in western Russia and Sweden in guidelines for sustainable use (Oxford
(Landwirtschaftsverlag, 2000).
Forestry Institute, University of Oxford,
stress, or animal browsing. Older trees can also develop so-called 2007, which has caused concern that the species will spread to United Kingdom, 2005). 128 pp. [18] J. P. Skovsgaard, I. M. Thomsen, I. M.
Skovgaard, T. Martinussen, Forest
'black heart, a non-fungal staining of the wood, which reduces other European countries and cause damage to ash trees. In North [7] E. Wallander, Plant Systematics and
Pathology 40, 7 (2010).
Evolution 273, 25 (2008).
the wood value5, 16 . Traditionally the leaves have been used as America, the emerald ash borer was first discovered in 2002, after [8] P. Wardle, Journal of Ecology 49, 739 [19] R. Bakys, R. Vasaitis, P. Barklund, I.
Thomsen, J. Stenlid, European Journal of
animal fodder during winter, while the bark was used to tan calf probably entering from Asia in shipping material. Since then it has (1961).
Forest Research 128, 51 (2009).
[9] E. Hultn, M. Fries, Atlas of North European
vascular plants (North of the Tropic of [20] J. D. Janse, Forest Pathology 11, 425
Cancer), Vols. I-III. (Koeltz scientific books, (1981).
Autoecology diagrams based on harmonised 1986). [21] D. de Rigo, et al., Scientific Topics Focus 2,
Field data in Europe (including absences) Observed presences in Europe field observations from forest plots. mri10a15+ (2016).
[10] EUFORGEN, Distribution map of common
ash (Fraxinus excelsior) (2009). [22] T. Kowalski, Forest Pathology 36, 264
www.euforgen.org. (2006).
[11] H. Meusel, E. Jager, S. Rauschert, E. [23] N. A. Straw, D. T. Williams, O. Kulinich, Y. I.
Average temperature of the coldest month (C)
with a straight trunk, sinuous branches directed upwards and > 75%
Chorology
frequently forked, and abundant root suckers at the base. The buds Native
Distribution natural range it has been widely planted and now is commonly
Habitat and Ecology
In comparison with the other two European ashes, common present and naturalised in other European countries up to 50in The manna ash occurs typically in Mediterranean temperate
ash (Fraxinus excelsior) and narrow-leaved ash (Fraxinus latitude. There are isolated populations in eastern Spain and in colline-mountain forests. In the northern part of its range and in
angustifolia), the manna ash has the smallest range, which South Turkey to western Syria and Lebanon4, 6-9 . Like other ash higher elevations it is found commonly on south-facing slopes,
covers southern-western Europe, from South-East France, species, manna ash shows great morphological variations inside where it can find the warmth required to grow. In central and
through Italy, the Mediterranean isles, the Balkan peninsula, up its natural distribution and several subspecies and varieties have eastern Europe it grows principally on calcareous soils, while in
to western Turkey. It is present from sea level up to 1500m in been described. Two subspecies are actually recognised and southern areas it also grows on silicate substrates, and does best
altitude (South Tyrol). The northern limits of its natural distribution accepted: the manna ash (Fraxinus ornus subsp. ornus) and the on soils with a pH around neutral. The mean annual precipitation
are the southern edge of the Alps and the Hungarian central Taurus flowering ash (Fraxinus ornus subsp. cilicica) an endemic needed is between 500 and 650mm, although it has a good
Transylvanian mountains (Bihar Mountains), but it is also present species with a scattered population in the Taurus Mountains of drought resistance, storing water in densely branched roots,
in more northern countries as an ornamental tree. Outside its Southern Turkey4, 10, 11 . and reducing transpiration. The root system is widely developed,
requiring gravelly, well-aerated and drained soils. It thrives better
on poor soil, suffering the competition of other broadleaved trees
on richer ones4, 7, 12 . Thanks to its plasticity, fast germination
and fast growth when young, this species easily colonises new
Uncertain, no-data habitats. The ability to resprout after cutting makes it also
well adapted to grow in areas disturbed by animal browsing,
Marginal/no presence < 5%
wildfires, landslides and logging4 . This tree species is not long-
Low presence 5% - 10% lived and rarely reaches 100 years7. This ash is found in several
Mid-low presence 10% - 30% forest communities, typically in mixed broadleaved forests as
a tree and also as a shrub in the understorey. It is associated
Medium presence 30% - 50%
with Mediterranean oaks (Quercus pubescens, Q. cerris, Q.
Mid-high presence 50% - 70% frainetto, Q. coccifera, Q. infectoria), chestnut (Castanea sativa),
High presence 70% - 90% hornbeams (Carpinus spp.), hop hornbeam (Ostrya carpinifolia)
and maples (Acer spp.). In Greece and Turkey it can be found
Very-high presence > 90%
in maquis belts with other deciduous or evergreen broadleaved
shrubs, and sometimes in mixed conifer forests with Lebanon
cedar (Cedrus libani), black pine (Pinus nigra) and occasionally
with Mediterranean firs (Abies spp.)4, 7, 13 .
Annual precipitation (mm)
Map 2: High resolution distribution map estimating the relative probability of presence. Inflorescences of narrow flowers with four linear petals.
(Copyright Ettore Balocchi, www.flickr.com: CC-BY)
References
[1] A . F. Mitchell, A field guide to the trees [16] J. E. Norris, A. Di Iorio, A. Stokes, B. C.
of Britain and northern Europe (Collins, Nicoll, A. Achim, Slope Stability and
1974). Erosion Control: Ecotechnological
[2] O. Johnson, D. More, Collins tree guide Solutions, J. E. Norris, et al., eds. (Springer
(Collins, 2006). Netherlands, 2008), pp. 167210.
[3] M. Goldstein, G. Simonetti, M. Watschinger, [17] G. Evans, Herb Tree 39, 24 (2014).
Mature dry and brown samaras in autumn. Alberi dEuropa (A. Mondadori, 1995). [18] A. Galati, G. Migliore, C. Scaffidi Saggio,
[4] D. Boshier, et al., Ash species in Europe: Colture artificiali di piante medicinali -
(Copyright Aldo De Bastiani, www.actaplantarum.org: AP) Produzione di metaboliti secondari nelle
biological characteristics and practical
guidelines for sustainable use (Oxford piante medicinali in coltura artificiale, F.
Forestry Institute, University of Oxford, Tognoni, A. Pardossi, A. Mensuali Sodi,
United Kingdom, 2005). 128 pp. eds. (Aracne editrice, Roma, 2007), pp.
Threats and Diseases [5] H. J. Elwes, A. Henry, The Trees of Great
287297.
[19] R. Schicchi, L. Camarda, V. Spadaro, R.
Britain and Ireland Vol. 4 (Privately
No serious threats have been documented for manna ash. printed, Edinburgh, 1909). Pitonzo, Quaderni di Botanica ambientale
e applicata 17, 151 (2006).
It is susceptible to the fungus Hymenoscyphus pseudoalbidus, [6] C. Thbaud, M. Debussche, Journal of
[20] R. Bakys, R. Vasaitis, P. Barklund, I.
Biogeography 18, 7 (1991).
also known as Chalara fraxinea, which causes massive diebacks [7] D. Bartha, Enzyklopdie der Holzgewchse:
Thomsen, J. Stenlid, European Journal of
Forest Research 128, 51 (2009).
of common ash and narrow-leaved ash in Europe20 . However, Handbuch und Atlas der Dendrologie,
[21] D. de Rigo, et al., Scientific Topics Focus 2,
A. Roloff, H. Weisgerber, U. M. Lang, B.
this ash does not seem to be a natural host of the pathogen, Stimm, P. Schtt, eds. (Wiley-Vch Verlag, mri10a15+ (2016).
Weinheim, 1996), vol. 3. [22] T. Kirisits, M. Matlakova, S. Mottinger-
as its vulnerability was tested with artificial inoculations on Kroupa, T. L. Cech, E. Halmschlager,
[8] H. Meusel, E. Jager, S. Rauschert, E.
seedlings21, 22 . Other generalist and manna ash pathogens have Weinert, Vergleichende Chorologie der Proceedings of the Conference of IUFRO
Zentraleuropischen Flora (Gustav Fischer Working Party 7.02.02, 11-16 May 2009,
been observed, but in most cases they were in balance with the Verlag Jena, 1978). Eirdir, Turkey, N. Grlevik, ed. (2009), pp.
Compound leaves comprising 5-9 ovate leaflets 7-10 cm 97119.
host, e.g. the cauliflower gall mite (Aceria fraxinivora)23, 24 , or [9] P. Csontos, J. Tams, T. Kalapos, Acta
long showing red autumn colours. Botanica Hungarica 43, 127 (2001). [23] K. Krutler, T. Kirisits, Journal of
in weakened plants for climatic reasons, e.g. the wood cankers (Copyright Stefano Zerauschek, www.flickr.com: AP) Agricultural Extension and Rural
[10] P. H. Davis, Flora of Turkey and the Development 4, 261 (2012).
East Aegean Islands, vol. 6 (Edinburgh
University Press, 1984). [24] M. Anthony, R. Sattler, C. Cooney-Sovetts,
Canadian Journal of Botany 61, 1580
Autoecology diagrams based on harmonised [11] M. Ylmaz, H. Serin, H. Zengin, G. Zengin, (1983).
Field data in Europe (including absences) Observed presences in Europe field observations from forest plots. Orman Mhendislii 46, 24 (2009).
[25] J. Kollr, Acta entomologica serbica 16,
[12] R. Popovi, M. Koji, B. Karadi, Bocconea 115 (2011).
5, 431 (1997).
[26] A. Alves, B. T. Linaldeddu, A. Deidda, B.
[13] C. Yceda, A. Gezer, H. Fakir, Scientific Scanu, A. J. L. Phillips, Fungal Diversity 67,
Average temperature of the coldest month (C)
[15] H.-D. Vlasin, L. Holonec, Bulletin UASVM [28] T. Wallmann, R. Stingl, Neilreichia 6, 183
Horticulture 71, 330 (2014). (2011).
grey bark. The leaves are up to 10cm long, simple, alternate, 25% - 50%
50% - 75%
coriaceous and glabrous. Their upper surface is dark-green and > 75%
Chorology
glossy and the lower surface is yellowish and matt. With an ovate, Native
elliptic or oblong shape, the leaf margin may be undulate with
spines, especially in the lower part of the tree. Flowers are small
(6mm in diameter), white and placed in axillary cymes. Holly is
normally dioecious and flowers between May and August2, 3 . The
fruit is a bright red drupe of 7-12mm size2 . Its seeds ripen in
late autumn and usually last throughout the winter, when birds,
rodents and larger herbivores eat them2-5 .
Tree form of the holly; it usually does not exceed 10 m tall.
Distribution (Copyright Sean MacEntee, www.flickr.com: CC-BY)
[10] A.-M. Skou, F. Toneatto, J. Kollmann, Plant Mediterranea 53, 480 (2014).
Ecology 213, 1131 (2012). [22] L. B. Orlikowski, G. Szkuta, Acta Mycologica
[11] J.-C. Rameau, D. Mansion, G. Dum, Flore 39, 19 (2004).
forestire franaise, Plaines et collines,
Annual precipitation (mm)
stage1, 3 . The bark is silver-grey and smooth between deep, wide Chorology
Actual range
fissures4 . The leaves are 20-45cm long, with 5 to 9 leaflets, the
ones from the apex being larger compared with those from the
base of the leaf4 . Crushed leaves have a scent like shoe-polish4 .
The crown diameter of the common walnut is larger in relation to
its stem diameter than any other broadleaf tree species used in
Europe5 . The fruit ripens during hot summers and is a large rounded
nut of 4-5cm and weighing up to 18g6 . It may be propagated both
by seeds and also vegetatively. It can hybridise and it has been
found that the hybrids between common walnut and black walnut
(Juglans nigra) have good vigour and form3 .
Map 1: Plot distribution and simplified chorology map for Juglans regia.
Frequency of Juglans regia occurrences within the field observations as
reported by the National Forest Inventories. The chorology of the actual
spatial range for J. regia is derived after Fornari et al. and the Botanical
Society of Britain & Ireland12, 24 .
Distribution
Common juniper has the widest distribution range of all
conifers6, 10 , and can be found throughout the Northern Hemisphere:
in North America, Europe, and Asia. It is the most northerly of
the juniper species and one of the most northerly conifers in the Map 1: Plot distribution and simplified chorology map for Juniperus communis.
Frequency of Juniperus communis occurrences within the field observations
world10 . In Europe, it can be found from northern Scandinavia11 as reported by the National Forest Inventories. The chorology of the native
to parts of southern Spain, although at more southern latitudes spatial range for J. communis is derived after Meusel and Jger19 .
it is usually confined to mountain areas. Juniper grows in low
elevations in pasture lands and abandoned fields, as well as at
high elevations, above the treeline in Eurosiberian mountains7. Importance and Usage
Juniper is amongst the most useful multi-purpose shrub
Habitat and Ecology species worldwide. Containing a large number of essential oils13 , Fleshy berry-like seed cones; they become purple-blue in colour when mature.
(Copyright Ettore Balocchi, www.flickr.com: CC-BY)
Common juniper is a typical shrub species of poor soils and extracts from its twigs, leaves, and berries (the blue-black seed
harsh environments7. It is drought and cold tolerant but requires cones) have been used as traditional remedies against urinary
plenty of light2 . It can grow on acidic sandy or calcareous soils12 and infections2 , dermatitis14 , or as a diuretic15 . The wood has even
favours free-draining soils and rocky outcrops. In many areas, juniper been shown suitable for artificial bone implants16 . The twigs,
is considered to be a pioneer species, able to colonise bare terrain leaves, and especially the berries represent an important food
and a range of soil types12 . The subspecies Juniperus communis ssp. source for several small and large animals, such as birds, deer,
alpina occurs in a narrow band above or north of the treeline. elk, cattle, horses and sheep2 , and humans use the berries for
References
[1] A. F. Mitchell, A field guide to the trees [11] T. H. DeLuca, O. Zackrisson, Plant and Soil
of Britain and northern Europe (Collins, 294, 147 (2007).
1974). [12] J. G. B. Oostermeijer, B. De Knegt, Plant
[2] P. A. Thomas, M. El-Barghathi, A. Polwart, Species Biology 19, 175 (2004).
Journal of Ecology 95, 1404 (2007). [13] R. P. Adams, Biochemical Systematics and
[3] B. Beikircher, S. Mayr, Plant, Cell & Ecology 26, 637 (1998).
Environment 31, 1545 (2008). [14] C. Cavaleiro, E. Pinto, M. J. Goncalves, L.
[4] R. P. Adams, R. N. Pandey, Biochemical Salgueiro, Journal of Applied Microbiology
Systematics and Ecology 31, 1271 100, 1333 (2006).
(2003). [15] R. A. Halberstein, Annals of Epidemiology
[5] R. P. Adams, A. E. Schwarzbach, Phytologia 15, 686 (2005).
95(2), 179 (2013). [16] K. A. Gross, E. Ezerietis, Journal of
Juniper in shrub form grown in a rocky habitat in the Ligurian Alps (Savona, North Italy). [6] A. Farjon, D. Filer, An Atlas of the Worlds Biomedical Materials Research 64A, 672
(Copyright Giovanni Caudullo: CC-BY) Conifers: An Analysis of their Distribution, (2003).
Biogeography, Diversity and Conservation [17] S. Vichi, M. R. Aumatell, S. Buxaderas, E.
Autoecology diagrams based on harmonised Status (Brill, 2013). Lpez-Tamames, Analytica Chimica Acta
Field data in Europe (including absences) Observed presences in Europe field observations from forest plots. [7] D. Garcia, R. Zamora, J. M. Gomez, P. 628, 222 (2008).
Jordano, J. A. Hodar, Journal of Ecology [18] F. Cooper, R. E. Stone, P. McEvoy, T. Wilkins,
88, 435 (2000). N. Reid, The conservation status of
[8] L. O. Pedro, A. Montserrat, T. Salvador, juniper formations in Ireland, Department
Annals of Botany 89, 205 (2002). of Environment, Heritage and Local
Average temperature of the coldest month (C)
The Complete Reference (Timber Press, - Band I, II, III (Gustav Fischer Verlag,
2009). Jena, 1998).
1-2.5cm long and 1-2.5mm wide, with two waxy, white shallow
stomata furrows above and a ridge below and a spiny tip. This
species is dioecious. The male plants have solitary pollen cones in
the leaf axils. They are yellow and egg-shaped, with three to seven
pollen sacs below. Female plants have axillary berry-like seed cones
known as galbulus. They are approximately spherical, brown-red in
colour and 7-12mm long, maturing in two years. They do not open Berry-like orange seed cones; they are brown-red when mature.
(Copyright Stefano Zerauschek, www.flickr.com: AP)
and end in three small triangle-shaped protuberances. Inside are
one to three brown triangular-ovoid seeds1-4 . Importance and Usage
The prickly Juniper is suitable for cultivation as an ornamental
shrub in southern Europe, where a number of cultivars, especially
with more pendulous foliage, are commonly planted in gardens
Map 1: Plot distribution and simplified chorology map for Juniperus oxycedrus.
and parks6 . Its wood is resistant and hard, highly valued for
Frequency of Juniperus oxycedrus occurrences within the field observations as making furniture and other carpentry items2 . Essential oils
reported by the National Forest Inventories. The chorology of the native spatial are extracted from the branches and leaves by destructive
range for J. oxycedrus is derived after Jalas andSuominen, and Klimko, etal.10, 11 .
distillation, especially in France and Turkey6, 7. This oil of cade is
used for medicinal purposes2 , such as to prepare empyreumatic
sea. They are principally associated with coastal grass and shrub oil8 . It has antiseptic and antiparasitic properties. Rectified cade
vegetation and in the clearings with Mediterranean pine woods: oil is also used as a fragrance component in soaps, detergents,
Aleppo pine (Pinus halepensis) Turkish pine (Pinus brutia), Italian creams, lotions and perfumes9 .
stone pine (Pinus pinea) and maritime pine (Pinus pinaster). The
inland subspecies are found at higher elevations in the maquis Threats and Diseases
and garrigue shrubland or open sclerophyll woods dominated by No important threats have been identified for the prickly
holm oak (Quercus ilex), mastic (Pistacia lentiscus) and European juniper. The populations in the natural range are stable and
hornbeam (Carpinus betulus), as well as appearing in montane in some areas this juniper is abundant. However, the coastal
and wetter forests with cedar of Lebanon (Cedrus libani), black subpopulations are more scattered than the past, especially in
pine (Pinus nigra) and other junipers (Juniperus foetidissima and Spain and around the Adriatic Sea, due to the impacts of urban
Juniperus excelsa)5, 6 . and tourist developments2, 5, 6 .
Mediterranean coastal vegetation with pricky juniper shrubland.
(Copyright Stefano Zerauschek, www.flickr.com: AP)
Distribution
This juniper is native to the Mediterranean region and
widespread from Morocco and Portugal, to Lebanon and Syria,
reaching Kurdistan in Iran, Iraq and the Caucasus mountains.
There are four commonly accepted subspecies: Juniperus
oxycedrus subsp. oxycedrus has the largest range set in the
inlands as well as in the coasts of the species distribution range,
Juniperus oxycedrus subsp. macrocarpa commonly throughout
the coasts of the Mediterranean and Black Sea, Juniperus
oxycedrus subsp. badia in the inlands of North Algeria and Iberian
Peninsula, Juniperus oxycedrus subsp. transtagana in lowlands
and coasts of central Portugal and South-West Spain1, 3, 5 .
[5] A. Farjon, D. Filer, An Atlas of the Worlds [11] M. Klimko, et al., Flora - Morphology,
Conifers: An Analysis of their Distribution, Distribution, Functional Ecology of Plants
Biogeography, Diversity and Conservation 202, 133 (2007).
Status (Brill, 2013).
Annual precipitation (mm)
coastal pines (Pinus pinea, Pinus pinaster, Pinus brutia and S.Mazzoleni, G.D. Pasquale, M.Mulligan,
P.D. Martino, F.Rego, eds. (John Wiley &
Pinus halepensis) most often in plantations but also in natural Sons, Ltd, Chichester, UK, 2004), pp. 1-12.
habitats10, 18, 19 .On cliffs Phoenician juniper forms typical
scrubland, thriving on dry, rocky and often limestone substrates, re-establishment .
18, 20 of the chapter. The full version of
This is an extended summary
this chapter (revised and peer-reviewed) will be published online at
characterised by harsh conditions and called arborescent https://w3id.org/mtv/FISE-Comm/v01/e012f63. The purpose of this
matorrals. On coastal cliffs, plants are exposed to the sea summary is to provide an accessible dissemination of the related
main topics.
spray, sea winds, and severe winter storms followed by drought This QR code points to the full online version, where the most
updated content may be freely accessed.
summers, developing as a short shrub and shaped by the wind. Please, cite as:
Instead, mountain populations can reach high elevations and Caudullo, G., de Rigo, D., 2016. Juniperus phoenicea in Europe:
Large Phoenician juniper on limestone xeric soil in Milos (Aegean Islands, Greece) distribution, habitat, usage and threats. In: San-Miguel-Ayanz,
are adapted to a more continental climate. Usually they grow on J., de Rigo, D., Caudullo, G., Houston Durrant, T., Mauri, A. (Eds.),
(Copyright Pavel Burk, www.biolib.cz: PD)
European Atlas of Forest Tree Species. Publ. Off. EU, Luxembourg,
pp. e012f63+
then it becomes broad, rounded, and often irregular. The bark is 25% - 50%
50% - 75%
thin, dark brown, grey-brown at maturity, scaly and exfoliating in > 75%
Chorology
strips. Leaves are light green, 2mm long, acute scaly, appressed, Native
covering completely the twigs. This tree species is dioecious.
Flowers in male trees are clustered in 3-4mm yellow spherical
cones. Female trees have almost undetectable flowers that ripen
into 7-8mm berry-like dark-purple fleshy cones2 .
Berry-like dark-blue seed cones: they mature over a period of around 18 months.
Distribution (Copyright Gaston Aitor: CC-BY)
The Spanish juniper is endemic to South-Western Europe and viable seeds9 . Despite its tolerance to harsh climatic conditions,
North Africa3 . European juniper woodlands are mainly in Spain, a large contraction of the Spanish juniper range is expected as a
covering about 600000 ha4 , but also occur in French Alps and result of climate change15 . In Morocco heavy grazing and browsing
Pyrenees, in Corsica and the Italian Alps, with a total of 1500 ha5 , pressures have caused damage and prevented regeneration5 .
in Morocco in the High and Middle Atlas mountains, with an
area of 20000 ha, and in Algeria in limited areas of the Aures
mountains6 . In Spain, this species is concentrated mainly in the
Eastern part of the Northern Plateau and the Iberian Range, but Map 1: Plot distribution and simplified chorology map for Juniperus thurifera.
Frequency of Juniperus thurifera occurrences within the field observations
it found also in the Betic Range, the Ebro River Valley and the as reported by the National Forest Inventories. The chorology of the native
Cantabrian Range7. spatial range for J. thurifera is derived after several sources16-18 .
References
[1] J. Ruiz de la Torre, Flora Mayor (Organismo [9] R. Alonso, J. M. Barrio, S. Roig, Selvicultura
Autnomo Parques Nacionales & Direccin de Juniperus thurifera (Instituto Nacional
General para la Biodiversidad, 2006). de Investigacin y Tecnologa Agraria y
[2] C. J. Earle, The gymnosperm database Alimentaria (Espaa), 2008), pp. 229258.
(2015). http://www.conifers.org [10] L. DeSoto, J. M. Olano, V. Rozas, M. De la Cruz,
[3] J. do Amaral Franco, Flora Iberica: plantas Applied Vegetation Science 13, 15 (2010).
vasculares de la Peninsula Iberica e [11] A. Farjon, The IUCN Red List of Threatened
Islas Baleares, Volume 1: Lycopodiaceae- Species (2013), pp. 42255/0+.
Papaveraceae, S. Castroviejo, et al., eds. [12] G.Escribano-Avila, etal., PLoS ONE 7,
(Real Jardn Botnico, CSIC, Madrid, 1998), e46993 (2012).
pp. 181188.
[13] Council of the European Union, Official
[4] R. Alonso Ponce, O. Sanchez Palomares, Journal of the European Union 35, 7 (1992).
S. Roig Gomez, E. Lopez Senespleda, J.
M. Gandullo Gutierrez, Las estaciones [14] C. Muoz, V. Prez, P. Cobos, R. Hernndez,
ecolgicas actuales y potenciales de los G. Snchez, Sanidad forestal: gua en
sabinares albares espaoles, Monografas imagenes de plagas, enfermedades y otros
INIA. Serie Forestal n. 19 (Instituto agentes presentes en los bosques (Mundi-
Nacional de Investigacin y Tecnologa Prensa, Madrid, 2003).
Agraria y Alimentara, 2010). [15] M. Benito Garzn, R. Snchez de Dios, H.
[5] T. Gauquelin, V. Bertaudiere, N. Montes, Sainz Ollero, Applied Vegetation Science
Young Spanish juniper stand developing after agricultural use abandonment and decreasing grazing pressure. 11, 169 (2008).
W. Badri, J.-f. Asmode, Biodiversity &
(Copyright Gaston Aitor: CC-BY) Conservation 8, 1479 (1999). [16] J. Jalas, J. Suominen, Atlas Florae
[6] N. Montes, V. Bertaudiere, Thuriferous Europaeae: distribution of vascular plants
Autoecology diagrams based on harmonised juniper (Juniperus thurifera L.) in morocco: in Europe Vol. 2 Gymnospermae (Pinaceae
Field data in Europe (including absences) Observed presences in Europe field observations from forest plots. an endangered species (2005). Accessed to Ephedraceae) (Committee for Mapping
on January 2015. the Flora of Europe and Societas Biologica
Fennica Vanamo, Helsinki, 1973).
[7] A. Gastn Gonzlez, C. Soriano Martn,
Investigacin Agraria: Sistemas y Recursos [17] O. de Bols, J. Vigo, Flora dels pasos
Forestales 15, 9+ (2006). catalans, vol I-IV (Barcino, Barcelona,
Average temperature of the coldest month (C)
1984-2001).
[8] A. Gastn Gonzlez, J. I. Garca Vias, El
Estudio del hbitat climtico para la seleccin [18] Anthos, Information System of the plants
de especies la restauracin de la vegetacin of Spain (Real Jardn Botnico, CSIC -
Annual precipitation (mm)
Larix decidua and other larches in Europe: distribution, habitat, usage and threats
F. Da Ronch, G. Caudullo, W. Tinner, D. de Rigo between 650 and 1900m in elevation; the Poland larch (Larix
decidua var. polonica) with patchy often isolated stands growing
The European larch (Larix decidua Mill.) is a pioneer, very long-lived, fast-growing coniferous tree, which occurs in the in central-south Poland between 180 and 650m1, 4, 11 . The
central and eastern mountains of Europe, forming open forests or pasture woods at the upper tree limits. Larch is the Carpathian and the Poland larches are often grouped together
only deciduous conifer in Europe as an adaptation to continental alpine climates. In fact, it is able to tolerate very cold or divided in geographic subspecies by several authors2, 11 . In
temperatures during winter and, by losing its needles, avoids foliage desiccation. It is a transitional species, colonising north-western Europe (Great Britain, Scandinavia) larches have
open terrain after natural disturbances. It forms pure stands but more often it is found with other alpine tree species, been widely cultivated since 16th century and naturalised in
which tend to replace it if no other disturbances occur. Thanks to its adaptability and the durability of its wood, the some cases1, 12, 13 . More recently, these plantations have been
European larch represents an important silvicultural tree species in the alpine regions, planted even outside its natural largely superseded first by the Japanese larch (Larix kaempferi)
ranges. Its wood is largely used for carpentry, furniture and pulp for paper. In lower altitudes or with high precipitation or Siberian larch (Larix sibirica) and then by larch hybrids (i.e.
rates, larch is more susceptible to fungal diseases. Outbreaks of insect defoliators, principally caused by thelarch bud Larix x marschlinsii)2 . The European larch was also introduced in
moth(Zeiraphera diniana), can limit tree development, with economic losses in plantations, but they rarely lead to the southern Canada and the north-eastern United States from the
death of the trees. mid-19th century14, 15 , and in New Zealand, where it is classified
The European larch (Larix decidua Mill.) is a large deciduous as a naturalised and invasive species16 .
coniferous tree that reaches 45 m, rarely over 50 m, and a
lifespan of 600-800 years in optimal conditions1 . Like other tree
Frequency
< 25%
Habitat and Ecology
species, larches in the highest elevations are more slow-growing 25% - 50% The European larch is a light-demanding, pioneer species of the
50% - 75%
and long-living, reaching even more than 1000 years in age, so > 75% mountain and subalpine regions. This conifer has a large ecological
making this species suitable for dendrochronology studies2, 3 . The
Chorology
amplitude. In the Alps and Tatra Mountains it grows in continental
Native
trunk is monopodial, straight or curved at base in slopes, with a climates, with cold, dry and snowy winters. In Poland and in the
diameter of 1-1.5 (2)m and fissured bark from reddish brown to Sudeten Mountains larch thrives at lower altitudes in sub-continental
light grey4 . Young plants are very flexible and are not damaged climates with a more temperate influence1, 11 . It needs light in all
by avalanches1 . The needles are clustered in bunches of 20-40, stages of development, colonising disturbed soils (avalanches,
flexible, 1.5-4cm long and 1mm wide. The colour is light green and landslide, livestock grazing, etc.) and forming open woodlands1 . In
turns yellow in autumn before falling2 . The larch is a monoecious lower elevations it is a transitional tree, performing as coloniser
unisexual species: the male cones (5-10mm) are sulphur yellow, better than other mountain tree species in poor to medium nutrient
with a reddish margin, hanging on the longer branches; the female sites8, 17, while in the subalpine belt it forms more stable forests in
cones are pink-red or dark purple when immature and turn a light pure or mixed stands10. It grows on well-drained soils, not tolerating
green with purple margins at maturity. The seed cones are about waterlogging, with a pH range from neutral to acid12 . The larch is
2.5cm long and persist in the plant up to 10 years; when old they very cold and wind tolerant during winter (dormant period), and it
turn to a grey colour and fall along with the small branches. The has a cold hardiness limit of around -30C18 . Its deciduous habit
seeds are 4-5mm long, greyish in colour5 . confers a significant advantage by reducing desiccation damage on
Map 1: Plot distribution and simplified chorology map for Larix decidua. foliage during winter8 . In the Alps at higher elevations larch forms the
Distribution Frequency of Larix decidua occurrences within the field observations as
reported by the National Forest Inventories. The chorology of the native
upper tree limit, occurring in pure forests in the Italian, French and
European larch is discontinuously distributed in the spatial range for L. decidua is derived after Wagner et al.41 . Southern Swiss Alps, while farther north this species is more often
mountains of southern, central and Eastern Europe, from south- found in mixed stands with other alpine tree species, principally the
eastern France and south-western Italy to eastern Poland and under debate. Three main varieties are recognised: the Alpine Swiss stone pine (Pinus cembra), but also green alder (Alnus viridis)
central Romania6-8 . It has a broad vertical range, forming forests larch (Larix decidua var. decidua), living in a compact area that and dwarf mountain pine (Pinus mugo). In the lower elevations it can
between 180m (in Poland) to 2500m (central Alps, south- includes the Alpine mountains continuing up to east Austria and be found with Norway spruce (Picea abies) and silver fir (Abies alba),
western Alps), but reaching very high elevations where it can northern Slovenia between 250m (Austria) and 2300m (western while lower down with beech (Fagus sylvatica) on poor soils and in
found in small groups or single trees in sheltered sites9-12 . The Alps) of elevation; the Carpathian larch (Larix decidua var. open and disturbed areas4, 9, 10, 15 . In the Carpathian Mountains larch
species is divided in different geographic varieties, sometimes carpatica) with three more patchy populations, in the Sudeten occurs usually with Norway spruce and Swiss stone pine or Scots
given the status of subspecies, and their classification is still Mountains, the Tatra Mountains and in Southeast Carpathians, pine (Pinus sylvestris), and also with fir and beech. It only sporadically
Uncertain, no-data
Marginal/no presence < 5%
Low presence 5% - 10%
Mid-low presence 10% - 30%
Medium presence 30% - 50%
Mid-high presence 50% - 70%
High presence 70% - 90%
Very-high presence > 90%
Map 2: High resolution distribution map estimating the relative probability of presence for the whole genus Larix. Large isolated larch; this tree can in some cases grow to over 50m.
(Copyright Giovanni Caudullo: CC-BY)
forms the timberline belt, typically in the High Tatra Mountains with
Swiss stone pine, a vegetation similar to Alpine larch woods1, 19-21 .
In Poland it occurs in lowlands growing in small groups or even as
individual trees within pine-birch and oak-birch forests1, 21, 22 . These Uncertain, no-data
ecotypes are less light demanding and able to grow in the shade of Tundra, cold desert
other tree species11 .
Negligible survivability
Importance and Usage Low survivability
The larch is an economically and traditionally important
Mid-low survivability
timber tree in Europe, thanks to its fast-growing nature, high
adaptability and its durable wood8 . The heartwood ranges Medium survivability
from yellow to a medium reddish brown. The narrow sapwood Mid-high survivability
is nearly white or pale yellow and is clearly demarcated from
High survivability
the heartwood. The wood is hard, strongly fragrant and is valued
for its durability, due to its concentration in tannins (up to 10%)
and resin content (about 2.6%)1 . It is also durable under water.
In fact it is largely used for carpentry and naval constructions8 .
In the European mountain areas its wood has been traditionally
used for building wooden houses23 , for producing furniture and
fine floors and many weatherproof outdoor objects such as
fences, gates, benches and tables, wooden roof shingles and
water troughs for cattle11 . Moreover, larch is used for pulpwood
and its good fibre characteristics (also for high-quality printing
paper23), for extracting tannin from bark and resin from wood,
and also as an ornamental tree, appreciated for airy foliage
turning to bright yellow in autumn8 . The larch turpentine, also
known as Venice turpentine, is obtained by distillation of larch
resin and it has been used in traditional medicine as antitussive
and expectorant action for colds, and more recently as industrial
solvent, for paint and wax preparation, or as a source of organic
compounds (e.g. camphor, rosin, etc.). The larch essential oil is
still used in aromatherapy and as perfume1 . In some Alpine areas
larch wood is still used to manufacture the Alphorn, a wooden
horn, 3-4m long, end-curved, played with its end resting on the
ground, which is able to produce very low and strong notes with
great effort from the musician. It was originally used in central
European Alps for communications among village communities
through the valleys; now it is a traditional Alpine instrument24,25 . Map 3: High resolution map estimating the maximum habitat suitability for the whole genus Larix.
Adult plant has thick and plated greyish bark with deep fissures.
(Copyright Stefano Zerauschek, www.flickr.com: AP)
Leaves turn to yellow in autumn before falling: the only deciduous European conifer.
(Copyright Dave Durrant: CC-BY)
References
Threats and Diseases [1] T. Geburek, Enzyklopdie der Holzgewchse: [20] T. Zielonka, J. Holeksa, P. Fleischer, P.
While the larch heart-root system may offer a good resistance Handbuch und Atlas der Dendrologie, A. Kapusta, Journal of Vegetation Science
Roloff, H. Weisgerber, U. M. Lang, B. Stimm, 21, 31 (2010).
to windthrow26 , the species appears less resistant to rockfall26, 27 P. Schtt, eds. (Wiley-Vch Verlag, Weinheim, [21] A. Farjon, D. Filer, An Atlas of the Worlds
2002), vol. 3. A. Farjon, A handbook of the Conifers: An Analysis of their Distribution,
even if the thick bark may offer a lower rockfall-mortality rate when worlds conifers (Brill, 2010). Biogeography, Diversity and Conservation
compared with spruce26 . Larches are vulnerable to Ips typographus [2] J. E. Eckenwalder, Conifers of the World: The Status (Brill, 2013).
Complete Reference (Timber Press, 2009). [22] M. Ferchmin, Memorabilia Zoologica 32,
and to other species of the Ips genus, such as Ips cembrae. These [3] P. Schtt, H. J. Schuck, B. Stimm, Lexikon 149 (1979).
bark beetles are also associated as vectors of important fungal der Baum- und Straucharten: Das [23] V. Chalupa, Trees III, Y. P. S. Bajaj, ed.
Standardwerk der Forstbotanik (Nikol, (Springer Berlin Heidelberg, 1991), vol.
pathogens28-32 . The larch canker, caused by Lachnellula wilkommii, Hamburg, 2002). 16 of Biotechnology in Agriculture and
is a fungal disease, which causes cankers distorting branches and [4] A. Farjon, A handbook of the worlds Forestry, pp. 446-470.
conifers (Brill, Leiden, 2010). [24] B. Geiser, Das Alphorn in der Schweiz (Paul
stems.It is considered the most destructive disease of the larch [5] J. Silba, Encyclopedia Coniferae, Haupt Publisher, Bern, 1976).
in Europe, particularly at lower altitudes and on inadequate, badly Phytologia Memoirs VIII (Harold N. [25] C. Vignau, Modernity, Complex Societies,
Moldenke and Alma L. Moldenke, Corvallis, and the Alphorn (Lexington Books,
drained sites8 . Other fungal diseases are the leaf cast fungus, Oregon, 1986). Maryland, 2013).
Meria laricis, which may cause significant defoliation, the root rot [6] H. Meusel, E. Jager, S. Rauschert, E. [26] J. E. Norris, A. Di Iorio, A. Stokes, B. C.
Weinert, Vergleichende Chorologie der Nicoll, A. Achim, Slope Stability and
Heterobasidion annosum12 and the velvet-top fungus Phaeolus Zentraleuropischen Flora (Gustav Fischer Erosion Control: Ecotechnological
Needle-like leaves in spring: they are arranged in clusters of 20-40 in the branchlets. Verlag Jena, 1978).
schweinitzii causing butt rot. Defoliations, sometimes heavy, can (Copyright Stefano Zerauschek, www.flickr.com: AP)
Solutions, J. E. Norris, et al., eds. (Springer
[7] J. Jalas, J. Suominen, Atlas Florae Netherlands, 2008), pp. 167210.
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in Europe Vol. 2 Gymnospermae (Pinaceae
laricella) or thelarch bud moth(Zeiraphera diniana). Tree deaths to Ephedraceae) (Committee for Mapping
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Management 215, 183 (2005).
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[9] P. Ozenda, Die Vegetation der Alpen
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among the most serious pests affecting young coniferous forests
[10] H. H. Ellenberg, Vegetation Ecology of [31] R. Kirschner, D. Begerow, F. Oberwinkler,
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Forest Ecosystems, pp. 177227.
[34] P. Nola, M. Morales, R. Motta, R. Villalba,
promoted larch presence with more open woodlands where the [12] P. S. Savill, The silviculture of trees used in Trees 20, 371 (2006).
larch regeneration was guaranteed. Today larches are mostly British forestry (CABI, 2013).
[35] R. Burgess, Risks of Exotic Forest Pests
[13] H. J. Elwes, A. Henry, The Trees of Great and Their Impact on Trade (The American
in forest reserves and are no longer logged, leading through Britain and Ireland Vol. 2 (Privately Phytopathological Society, 2001).
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[36] J. I. Barredo, et al., EPPO Bulletin 45, 273 (2015).
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Chronicle 78, 822 (2002). [37] CABI, Hylobius abietis (large pine weevil)
(2015). Invasive Species Compendium.
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Species (2014), pp. 34161/0+.
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Field data in Europe (including absences) Observed presences in Europe Jones, Flora of New Zealand Vol. 4. [39] J. Do Amaral Franco, Flora Europea.
observations from forest plots for Larix kempferi. Volume 1. Psilotaceae to Platanaceae, T.
Naturalised Pteridophytes, gymnosperms,
dicotyledons (D.S.I.R., Christchurch, 1988). G. Tutin, et al., eds. (Cambridge University
Press, 1993), p. 40, second edn.
[17] R. Del Favero, I boschi delle regioni alpine
italiane (Cleup, Padova, 2004). [40] G. R. Stairs, 13th Northeastern Forest
Average temperature of the coldest month (C)
[18] P. Bannister, G. Neuner, Conifer Cold Albany, New York, August 12-13, 1965
Hardiness, F. Bigras, S. Colombo, eds. (1966), pp. 3036.
(Springer Netherlands, 2001), vol. 1 of
Tree Physiology, pp. 321. [41] S. Wagner, T. Litt, M.-F. Snchez-Goi, R.
Annual precipitation (mm)
Distribution
In spite of the controversy generated on its origin, most authors
agree that wild olives are native to Minor Asia7. From the eastern
Map 1: Plot distribution and simplified chorology map for Olea europaea.
parts of the Mediterranean basin, olive trees spread west through Frequency of Olea europaea occurrences within the field observations as
Greece, Italy, France Spain and Portugal following the coasts8, 9. reported by the National Forest Inventories. The chorology of the native
Nowadays olive cultivations and selection of cultivars are expanding spatial range for O. europaea is derived after Meusel andJger38 .
in many areas outside its natural ranges, and even in other Maturing fruits: these fleshy drupes become black when ripening.
(Copyright Michael Wunderli, www.flickr.com: CC-BY)
continents, such as Australia, South and North America (Argentina, domestication of the oleaster seems to have occurred in the Near-
Chile, United States), South Africa and even in exotic places like East during the early Neolithic period, and then it was successively References
Hawaii8 . Of its six subspecies, only three are naturally distributed [1] S. Rhizopoulou, American-Eurasian Journal [19] P. Hanelt, ed., Mansfelds Encyclopedia
propagated to western areas of Mediterranean basin18 . Historically of Agricultural & Environmental Science 2, of Agricultural and Horticultural Crops
in Europe: subsp. europaea in the Mediterranean basin (Greece, this species has always been appreciated first for its fruits and the 382 (2007). (Springer, 2001).
Italy, Spain, Portugal, France, Cyprus, Slovenia and Malta) and some [2] G. S. Sibbett, L. Ferguson, Olive Production [20] B. K. Shishkin, et al., Flora of the USSR -
wood, and then for the oil. The fruit is edible and all parts contain Manual (Universityt of California, Volume XVIII: Metachlamydeae, vol. 18 of
Atlantic enclaves in South-West Europe; subsp. guanchica in the non-drying oil. Pickled, canned or otherwise prepared table olives are Agriculture & Natural Resources, Oakland, Flora of the USSR (Keter Press, Jerusalem,
2004), second edn. 1970).
Canary Islands; and subsp. cerasiformis in the Madeira archipelago. eaten as relish or used in bread, soups, salads, etc.19. The olive wood [3] G. C. Martin, The Woody Plant Seed [21] C. Breton, et al., Comptes Rendus Biologies
Of the other three, subsp. maroccana occurs in Morocco, subsp. is heavy and very tough, used for high-end furniture, inlays, turned Manual, F. T. Bonner, R. P. Karrfalt, 332, 1059 (2009).
eds., Agriculture Handbook 727 (U.S. [22] H. M. Roche, M. J. Gibney, A. Kafatos, A.
laperrinei in Algeria, Sudan and Niger, and subsp. cuspidata from objects, and handcraft20. It is also appreciated as firewood because it Department of Agriculture, Forest Service, Zampelas, C. M. Williams, Food Research
2008), pp. 753756.
South Africa throughout East Africa, Arabia to South West China. burns even when wet21 . The olive oil has several uses, for eating and International 33, 227 (2000).
[4] P. Vargas, S. Talavera, Flora Iberica: [23] H. M. Roche, et al., American Journal of
The Mediterranean subspecies includes the oleaster (Olea europaea cooking, as well as for ointment, lighting (burning without smoke), plantas vasculares de la Peninsula Clinical Nutrition 68, 552 (1998).
Ibrica e Islas Baleares, Volume 11
subsp. europaea var. sylvestris), the wild form, and cultivated olive and medical uses15, 20, 21 . Virgin olive oil is an important component Gentianaceae-Boraginaceae, S. Talavera, [24] European Commission, EU olive oil farms
et al., eds. (Real Jardn Botnico, CSIC, report Based on FADN data (Directorate-
(Olea europaea subsp. europaea var. europaea)4, 8, 10, 11 . of the Mediterranean diet, valued for its beneficial properties for Madrid, 2012), pp. 136139. General for Agriculture and Rural
human health thanks to the high amounts of unsaturated fatty Development, Brussels, 2012).
[5] M. Agnoletti, ed., Italian Historical Rural
Habitat and Ecology acids22, 23. Mediterranean countries produce more than 70% of the Landscapes, vol. 1 (Springer Netherlands, [25] C. M. Breton, P. Warnock, A. J. Bervill,
Olive Germplasm - The Olive Cultivation,
Dordrecht, 2013).
This species is a typical component of the thermo- total world supply of olive oil. About 95% of the European production [6] J. M. Alcantara, P. J. Rey, F. Valera, A. M.
Table Olive and Olive Oil Industry in Italy, I.
Muzzalupo, ed. (InTech, 2012), chap. 1.
Mediterranean climate, characterised by warm, dry summers Sanchez-Lafuente, Ecology 81, 1937 (2000).
[26] S. Rickard, The New Ornamental Garden
[7] G. Bartolini, R. Petruccelli, H. D. Tindall,
and rainy, cool winters, which corresponds generally to the U. G. Menini, Classification, origin,
(CSIRO Publishing, Melbourne, 2011).
coasts of the Mediterranean Sea up to 200m in elevation12 . The diffusion and history of the olive (Food & [27] E. Rugini, C. De Pace, P. Gutirrez-Pesce, R.
Agriculture Organization, Rome, 2002). Muleo, Wild Crop Relatives: Genomic and
olive tree is a thermophile species and is adapted to tolerate [8] A. Chiappetta, I. Muzzalupo, Olive
Breeding Resources: Temperate Fruits, C.
Kole, ed. (Springer-Verlag, Berlin, 2011),
drought and salinity stress1, 8, 13 . It grows on a wide range of Germplasm - The Olive Cultivation, Table pp. 79117.
Olive and Olive Oil Industry in Italy, I.
soils4 , but prefers sandy loam soils of moderate depth14 . The wild Muzzalupo, ed. (InTech, 2012), chap. 2. [28] V. Sergeeva, R. Spooner-Hart, N. G. Nair,
Australasian Plant Disease Notes 3, 143
form frequently thrives as one of the common constituents of [9] A. P. De Candolle, Origine des Plantes (2008).
Cultivees (Germer Baillire, Paris, 1883).
maquis and garrigue scrub formations on poor soils and slopes. [10] G. Besnard, R. R. de Casas, P.-A. Christin, P.
[29] J. Mercado-Blanco, F. J. Lpez-Escudero,
Plant and Soil 355, 17 (2012).
It colonises secondary habitats, such as the edges of cultivation Vargas, Annals of Botany 104, 143 (2009).
[30] J. R. Viruega, J. Moral, L. F. Roca, N.
[11] P. S. Green, Kew Bulletin 57, 91 (2002).
or abandoned orchards, spread by bird-dispersed seed, but also Navarro, A. Trapero, Plant Disease 97,
[12] Food and Agriculture Organization of 1549 (2013).
propagating vegetatively by root suckers3, 15, 16 . It can be found the United Nations, Global Ecological [31] G. Marchi, C. Viti, L. Giovannetti, G. Surico,
Zoning for the Global Forest Resources
in the sclerophyllous evergreen vegetation, along with carob Assessment 2000 - Final Report (Food
European Journal of Plant Pathology 112,
101 (2005).
(Ceratonia siliqua) mastic (Pistacia lentiscus), myrtle (Myrtus and Agriculture Organization of the United
Nations, Forestry Department, Rome, [32] G. E. Haniotakis, IOBC/WPRS Bulletin 28,
communis), junipers (Juniperus spp.), etc.17. Italy, 2001). 1 (2005).
[13] G. Lpez Gonzlez, Los rboles y arbustos [33] E. Rugini, M. Mencuccini, R. Biasi,
M. Altamura, Protocol for Somatic
Importance and Usage de la Pennsula Ibrica e Islas Baleares
(Mundi-Prensa, Madrid, 2006), second edn. Embryogenesis in Woody Plants, Jain, P.
Gupta, eds. (Springer Netherlands, 2005),
The olive is one of the most emblematic and economically [14] D. H. R. Spennemann, L. R. Allen, Australian vol. 77 of Forestry Sciences, pp. 345360.
Old cutivated olive tree for fruit production near Galatas (Peloponnese Journal of Experimental Agriculture 40,
important crop trees of the Mediterranean regions15. The peninsula, Greece). 889 (2000). [34] G. Besnard, P. Henry, L. Wille, D. Cooke, E.
Chapuis, Heredity 99, 608 (2007).
(Copyright Miltos Gikas, www.flickr.com: CC-BY) [15] D. Zohary, M. Hopf, E. Weiss, Domestication
of Plants in the Old World (Oxford [35] D. de Rigo, et al., Scientific Topics Focus 2,
Autoecology diagrams based on harmonised University Press, Oxford, 2000). mri10a15+ (2016).
Field data in Europe (including absences) Observed presences in Europe field observations from forest plots. [16] J. M. Caballero, C. del Ro, El Cultivo del [36] R. Baker, et al., EFSA Journal 13, 3989+
Olivo, D. Barranco, R. Fernndez-Escobar, (2015).
L. Rallo, eds. (Mundi-Prensa, Madrid, [37] G. P. Martelli, D. Boscia, F. Porcelli, M.
2008), pp. 93125, 6th edn. Saponari, European Journal of Plant
[17] U. Bohn, et al., Karte der natrlichen Pathology pp. 19 (2015).
Average temperature of the coldest month (C)
Distribution
The European hop-hornbeam is endemic to temperate
West Eurasia: more in detail, its native range includes Middle
Europe (South-East Switzerland and South Austria), Southern
Europe (South-East France and Corsica, Italy, Sardinia, Sicily), the Map 1: Plot distribution and simplified chorology map for Ostrya carpinifolia.
Frequency of Ostrya carpinifolia occurrences within the field observations
Balkan area (Albania, Bosnia and Herzegovina, Bulgaria, Croatia, as reported by the National Forest Inventories. The chorology of the native
Macedonia, Montenegro, Serbia, Slovenia, Greece, probably spatial range for O. carpinifolia is derived after Meusel and Jger4 .
extinct in Hungary) European Turkey, Western Asia (Anatolia,
Syria, Lebanon) and Caucasian countries (Georgia, Armenia, important role in all the Balkan-Illyrian area up to 900m in
Cis- and Transcaucasian districts of the Russian Federation)3-6 . elevation, occasionally reaching 1300-1400m. Here it takes part Male catkins are produced in spring and are 5-10 cm long.
Palynological evidence suggests that it was able to spread in species-rich shrubberies (the so-called shibljak): together with (Copyright Gianluca Nicolella, www.actaplantarum.org: AP)
northwards and colonise Europe only after the last glaciation; i.e. Cornus mas, C. sanguinea, Corylus avellana, Cotinus coggygria,
7000 years ago7. Cotoneaster nebrodensis, Juniperus oxycedrus, Paliurus spina-
christi, Syringa vulgaris and, in the warmest sites, also Coronilla Importance and Usage
Habitat and Ecology emerus and Pistacia terebinthus, it usually colonises open places The wood of the hop-hornbeam is very hard and heavy, difficult
This hop-hornbeam is a stenohydric plant8 , which means where it forms the understorey of Pinus nigra forests and deciduous to work2, 27. It has been used in the past for different purposes,
that it shows rather constant transpiration and osmotic pressure sub-Mediterranean forests, where it may represent one of the especially in rural areas, for making small items and charcoal28-30 .
values also under moderate drought stress conditions9 . Thus, dominating species together with Carpinus orientalis, Fraxinus It tends to crack when dried, so it is not appreciated for industrial
it is able to colonise windy and sunny slopes, but it is mostly ornus, and Quercus pubescens13 . These plant communities purposes, although it still represents an excellent firewood27, 31 . For
found in rainy areas or under wet microclimatic conditions (e.g. may be ascribed to the alliance of Ostryo-Carpinion orientalis, this very purpose, in central Italy most hop-hornbeam woodlands
deep and humid ravines and canyons) where air humidity is and similar species-assemblages are rather widespread on the are still intensely exploited as coppices16 . The ability to colonise dry
constantly available1 . This explains why in the northernmost part hillsides and mountains of central and northern Italy14-16 , former areas and shallow lime- and magnesium-rich soils makes this tree
of its range this species behaves as a light-demanding pioneer Yugoslavia17 and of the continental part of Greece18-20 , while species suitable for the reforestation of many degraded sites32 .
that prefers sunny and warm places, while in the southernmost those of Calabria, Corsica, Sardinia and Sicily21-24 may better It is also used to form hedges and as an ornamental tree along
countries it grows better in semi-shaded and more humid be referred to the alliance Pino laricionis-Quercion congestae roadsides27. Hop hornbeam is one of the hosting trees of the white
sites2, 10 . The European hop hornbeam often grows in rocky areas and those of South-East France to Quercetalia pubescentis25 . truffle (Tuber magnatum)33 .
and on shallow and poorly developed soils, mainly on limestone3 , European hop-hornbeam seems to have played an important
but also on volcanic11 and gypsum rock outcrops12 . It plays an role within the mature forest communities of the Near East26 .
Uncertain, no-data
Tundra, cold desert
Negligible survivability
Low survivability
Mid-low survivability
Medium survivability
Mid-high survivability
High survivability
References
[1] S. Pignatti, I boschi dItalia: sinecologia e [13] A. arni, et al., Plant Biosystems 143, 1 [26] S. Bottema, W. Van Zeist, Prhistoire du [31] D. Bartha, Enzyklopdie der Holzgewchse:
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[2] S. Korkut, B. Guller, Bioresource Technology [14] A. Hofmann, Studia Geobotanica 2, 217 lespace depuis les origines jusquau VI A. Roloff, H. Weisgerber, U. M. Lang, B.
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juin 1980), J. Cauvin, P. Sanlaville, eds., Weinheim, 2001).
[3] K. Browicks, J. Zieliski, Chorology of [15] D. Lausi, R. Gerdol, F. Piccoli, Studia Colloques Internationaux du CNRS n
trees and shrubs in south-west Asia and Geobotanica 2, 41 (1982). [32] M. Guidi, P. Piussi, Revue de Gographie
598 (Editions du Centre National de la Alpine 81, 95 (1993).
adjacent regions, vol. 1 (Polish Scientific [16] C. Blasi, G. Filibeck, L. Rosati, Fitosociologia Recherche Scientifique, Paris, 1981), pp.
Publishers, Warszawa, Pozna, 1982). 43, 3 (2006). 111132. [33] L. Bertini, et al., Microbiological Research
[4] H. Meusel, E. J. Jger, Plant Systematics 161, 59 (2006).
[17] R. Lakui, D. Pavlovi, S. Redi, Glasnik [27] M. Goldstein, G. Simonetti, M. Watschinger,
and Evolution 162, 315 (1989). Republickog Zavoda za Zastitu Prirode i Alberi dEuropa (A. Mondadori, 1995). [34] B. Pikur, et al., European Journal of Forest
[5] P. Uotila, Euro+Med Plantbase - the Prirodnjackog Muzeja Titogradu 15, 103 Research 130, 235 (2011).
[28] A. J. Panshin, C. de Zeeuw, Textbook of
information resource for Euro- (1982). Wood Technology: Structure, Identification, [35] G. Goidnich, Manuale di patologia
Mediterranean plant diversity (2009). [18] D. Voliotis, Acta Botanica Hungarica 31, Properties, and Uses of the Commercial vegetale, vol. 2 (Edagricole, Bologna,
http://www.emplantbase.org. 339 (1985). Woods of the United States and Canada, 1964).
[6] K. Shaw, S. Roy, B. Wilson, The IUCN Red vol. 1 of McGraw-Hill series in forest [36] D. Ottonello, Il Naturalista siciliano 10,
[19] A. Boratyski, K. Browicz, J. Zieliski, resources (Mcgraw-Hill College, 1980),
List of Threatened Species (2014), pp. Chorology of trees and shrubs in Greece 107 (1987).
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[7] K. J. Willis, Endeavour 20, 110 (1996). Dendrology, Kornik, Poland, 1992). [29] J. H. Flynn, A Guide to Useful Woods of the e Boschi 42, 54 (1991).
World (King Philip Publishing Co., Portland,
[8] R. Del Favero, O. Andrich, G. De Mas, [20] E. Milios, Silva Gandavensis 65, 128 Maine, 1994). [38] D. de Rigo, et al., Scientific Topics Focus 2,
C. Lasen, D. Poldini, La vegetazione (2000). mri10a15+ (2016).
forestale del Veneto - Prodromi di [30] H. A. Alden, Hardwoods of North America,
[21] J. Gamisans, La vgtation de la Corse General Technical Report FPL; GTR-83 (U.S. [39] M. Jurc, S. Bojovi, B. Komjanc, J. Kr,
tipologia forestale (Regione del Veneto, (Edisud, Aix-en-Provence, 1999). Biologia 64, 130 (2009).
Dipartimento per le Foreste e lEconomia Department of Agriculture, Forest Service,
Montana, Venezia, 1990). [22] S. Brullo, F. Scelsi, G. Spampinato, La Forest Products Laboratory, Madison, WI, [40] CABI, Scolytus intricatus (European oak
vegetazione dellAspromonte - studio 1995). bark beetle) (2015). Invasive Species
[9] V. De Micco, G. Aronne, P. Baas, Trees 22, fitosociologico (Laruffa, Reggio Calabria, Compendium. http://www.cabi.org
643 (2008). 2001).
[10] G. Venturella, P. Mazzola, F. M. Raimondo, [23] G. Bacchetta, G. Iiriti, L. Mossa, C.
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biogographie des forts du bassin (2012).
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[25] G. Lapraz, Bulletin de la Socit Botanique
[12] P. Marino, V. Ilardi, Atti del 102 Congresso de France. Lettres Botaniques 130, 137
della Societ Botanica Italiana, Palermo (1983).
26-29 settembre 2007, G. Venturella, F. M.
Raimondo, eds. (2007).
Isolated tree in winter. This species is a small tree rarely exceeding 25m.
(Copyright Stefano Zerauschek, www.flickr.com: AP)
underneath living individuals have been dated with radiocarbon to 25% - 50%
50% - 75%
the early Holocene, about 9500 years ago, suggesting vegetative > 75%
Chorology
survival (by re-sprouting from the roots) over millennia4 . The Native
crown is regularly conic, columnar, with whorled, short and stout
branches, the upper level ascending and the lower drooping. Buds
are reddish brown, 5mm long with an acute apex. Needles are
1-2.5cm long, 4-angled in cross section, rigid, light to dark green
with fine white speckled lines. The species is monoecious, with
unisexual flowers usually appearing at an age of 20-30 years,
but up to 40 years in dense stands. Male flowers are located
principally at the base of the preceding years shoot, 1-2.5cm
long, globular, crimson then yellow when mature. Female flowers
are located at the tip of the shoot, dark red, 5cm long, erect
before pollination, becoming pendent afterwards. Cones are
cylindrical, 12-15cm long, green before maturity, turning brown Majestic isolated spruce in a mountain field in Leskova Dolina
Map 1: Plot distribution and simplified chorology map for Picea abies. (Snenik, South Slovenia).
in autumn. When dry the cones open to disperse 4mm winged (Copyright Stefano Zerauschek, www.flickr.com: AP)
Caption: Frequency of Picea abies occurrences within the field observations
seeds. The bark is orange brown and the wood is creamy white as reported by the National Forest Inventories. The chorology of the native
and easy to work1, 5-8 . spatial range for P. abies is derived after EUFORGEN39 . (United States, Canada and Japan) and in the southern hemisphere
(South Africa, Tasmania and New Zealand)6, 16 .
Distribution distribution there are a great number of varieties and forms,
Norway spruce is the main species in the Boreal and subalpine which can be considered as normal patterns of variation within Habitat and Ecology
conifer forests, from Central (in mountains) to Northern and a widespread species. Historically cultivated since of the 18th Spruce forests dominates in the Boreal zone of N and N-E
Eastern Europe up to the Ural Mountains, where the species century, Norway spruce plantations, even outside natural ranges Europe; in central Europe spruce forests cover wide areas of the
merges with Siberian spruce (Picea obovata), which is sometimes (e.g. in the temperate lowland areas), have changed natural forests montane (mostly planted) and the sub-alpine zones, in lowlands
considered as a sub-species of Picea abies5, 9-13 . Its elevation range into artificial ones2, 14 . Now it is locally naturalised in many areas of it is more mixed with other species2 . The natural distribution
goes from sea level in Northern Europe up to above 2400m in Europe outside its native range, including Britain and the Pyrenees shows continental tendencies but thanks to its climatic tolerance
the Alps, where it grows in a stunted form14, 15 . Due to its large Mountains. It was also introduced in other countries outside Europe it grows even in extreme oceanic climates9, 17. Norway spruce is
a secondary coloniser, but can be both a pioneer and a climax
species. It shows good yield and quality performance under
very different site conditions, and has been favoured over long
periods by silviculture, especially in the lowlands, but also in
Uncertain, no-data mountain areas18 . Shade-tolerant, it can survive for decades
under a closed canopy, fast growing after 5-10 years. It does
Marginal/no presence < 5%
not grow close to coasts when exposed to salt winds, nor does
Low presence 5% - 10% it like summer drought or waterlogged conditions2, 7. Although it
Mid-low presence 10% - 30% can occur on most substrates, it is most common and widespread
on acidic soils, preferring nutritious deep soils with enough fresh
Medium presence 30% - 50%
moisture8, 9 . Spruce shows a noticeable soil-acidifying ability19 . In
Mid-high presence 50% - 70% the Boreal forests it grows with birch (Betula spp.) and European
High presence 70% - 90% aspen (Populus tremula), with willow (Salix spp.) alongside streams
and lakes. In the Alps, when it is not in pure stands, it occurs
Very-high presence > 90%
with European larch (Larix decidua) and Swiss stone pine (Pinus
cembra) on higher elevations (ca. 1800-2100m), with European
beech (Fagus sylvatica) and European silver fir (Abies alba) under
fresh conditions at intermediate altitudes (800-1800m), and with
Scots pine (Pinus sylvestris) in drier conditions9, 15 . Stages and
reproductive processes are regulated by climatic conditions, in
particular by temperature, which become more important in higher
Map 2: High resolution distribution map estimating the relative probability of presence. Droplets of resin are common over the bark of the trunk and branches.
(Copyright Agnieszka Kwiecie, commons.wikimedia.org: CC-BY)
Dead spruces killed by the bark beetle Ips typographus near Bo Dar inside
Top side of a violin made with spruce wood. the protected area Krunohorsk plat (Ostrov, North-West Czech Republic).
(Copyright Takeshi Kuboki, www.flickr.com: CC-BY) (Copyright Ji Berkovec, commons.wikimedia.org: PD)
fungi31, 35, 36 . Starting from the 1980s, spruce forests have shown range . Due to its preferences for cool and moist climatic
2, 8
symptoms of decline in mountainous areas of central Europe conditions this economically very valuable species may become
including yellowing, loss of needles, die-back of branches and severely affected under global warming conditions37. European
reduced growth. Air pollution has often been used to explain alternatives to Norway spruce are mostly fir species such as
this14 . Health problems in central European forests have reduced Abies alba (e.g. Mediterranean or dry inner Alpine provenances)
its popularity for reforestation, particularly outside its natural which can tolerate significantly warmer and drier conditions)37, 38 .
Old Tjikko, claimed to be the oldest tree in the world, a 9 550-year-old spruce
possibly surviving by root re-sprouting in Fulufjllet Mountain (Dalarna, Sweden).
(Copyright Karl Brodowsky, commons.wikimedia.org: CC-BY)
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[13] A. Farjon, D. Filer, An Atlas of the Worlds
Conifers: An Analysis of their Distribution, [35] S. Prospero, Ecology of Armillaria
Biogeography, Diversity and Conservation cepistipes: population structure, niches,
Status (Brill, 2013). pathogenicity and interactions with
Armillaria ostoyae, Masters thesis, ETH
[14] G. Jansson, et al., Forest Tree Breeding Zurich, Switzerland (2003).
Pure Norway spruce forest by Lake Carezza in the western Dolomites (Bolzano, North-East Italy). in Europe, L. E. Pques, ed. (Springer
(Copyright Son of Groucho, www.flickr.com: CC-BY) Netherlands, 2013), vol. 25 of Managing [36] D. Morrison, K. Mallett, Canadian Journal
Forest Ecosystems, pp. 123176. of Plant Pathology 18, 194 (1996).
[15] H. H. Ellenberg, Vegetation Ecology of [37] H. Bugmann, et al., Toward Quantitative
Autoecology diagrams based on harmonised Scenarios of Climate Change Impacts
Field data in Europe (including absences) Observed presences in Europe field observations from forest plots. Central Europe (Cambridge University
in Switzerland, C. Appenzeller, et al.,
Press, 2009), fourth edn.
eds. (OCCR, FOEN, MeteoSwiss, C2SM,
[16] J.-C. Svenning, F. Skov, Ecology Letters 7, Agroscope and ProClim, Bern, Switzerland,
565 (2004). 2014), pp. 7988.
[17] P. S. Savill, The silviculture of trees used in
Average temperature of the coldest month (C)
and between 2 and 4cm long4 . Seed cones are from 5 to 10cm
long, composed of papery scales with wavy, irregularly toothed
margins and producing seeds from 2 to 3mm long with a wing of
around 8mm in length9 .
Distribution
Sitka Spruce is native to the west coast of North America
where it extends along the north Pacific coast from southern
Alaska to northern California4, 5, 9 . It was introduced in Europe
in the 1800s and it is now planted in more than 16 countries
worldwide10 . The majority of the area planted is in the United
Kingdom where it comprises over 25% of the national forest
area11 and is now the most widely planted conifer3, 12 , and Ireland
Coastal temperate mixed forest dominated by Sitka spruce and western hemloc (52% of national forest area3) but it is also important in Denmark
(Tsuga heterophylla) in Misty Fiords National Monument (Ketchikan, Alaska).
(Copyright Kimberly Vardeman, commons.wikimedia.org: CC-BY) (comprising 16% of the softwood timber harvest13). Sitka Spruce
References
[1] A. Farjon, A handbook of the worlds [11] Forestry Commission, Forestry Statistics
conifers (Brill, Leiden, 2010). 2014 (Economics and Statistics, Forestry
[2] P. S. Savill, The silviculture of trees used in Commission, 231 Corstorphine Road,
British forestry (CABI, 2013). Edinburgh, 2014).
[3] S. Lee, D. Thompson, J. K. Hansen, Forest [12] G. F. Peterken, Forest Ecology and
Tree Breeding in Europe, L. E. Pques, Management 141, 31 (2001).
ed. (Springer Netherlands, 2013), vol. [13] J. Moore, Wood properties and uses
25 of Managing Forest Ecosystems, pp. of Sitka spruce in Britain (Forestry
177227. Commission, 2011).
[4] J. E. Eckenwalder, Conifers of the World: [14] G. Halldrsson, T. Benedikz, O. Eggertsson,
The Complete Reference (Timber Press, E. S. Oddsdttir, H. skarsson, Forest
2009). Ecology and Management 181, 281
[5] A. S. Harris, Sitka Spruce (Picea sitchensis (2003).
(Bong.) Carr.), Agriculture Handbook 654 [15] L. Augusto, J. Ranger, D. Binkley, A. Rothe,
(U.S. Department of Agriculture, Forest Annals of Forest Science 59, 233 (2002).
Service, Washington, DC., 1990), pp. [16] C. P. Quine, D. C. Malcolm, Canadian Journal
Ripening mature seed cones; they are usually 5 to 10 cm long. 513529. of Forest Research 37, 1787 (2007).
(Forestry Commission, www.forestry.gov.uk: Crown Copyright) [6] A. F. Mitchell, A field guide to the trees [17] A. Askenfelt, et al., The Science of String
of Britain and northern Europe (Collins, Instruments (Springer New York, New York,
Autoecology diagrams based on harmonised 1974). NY, 2010).
Field data in Europe (including absences) Observed presences in Europe field observations from forest plots. [7] A. Farjon, The IUCN Red List of Threatened [18] J. E. Norris, J. R. Greenwood, Proceedings
Species (2013), pp. 42337/0+. of the 10th IAEG International Congress,
[8] OECD, Safety Assessment of Transgenic IAEG2006 (2006), pp. 744+.
Organisms (OECD Publishing, 2006), vol. 2 [19] J. I. Barredo, et al., EPPO Bulletin 45, 273
of OECD Consensus Documents. (2015).
Average temperature of the coldest month (C)
[9] A. Praciak, et al., The CABI encyclopedia of [20] CABI, Hylobius abietis (large pine weevil)
forest trees (CABI, Oxfordshire, UK, 2013). (2015). Invasive Species Compendium.
[10] R. L. Deal, P. Hennon, R. OHanlon, D. http://www.cabi.org
Annual precipitation (mm)
cylindrical and finally very open6 . It grows commonly in a curved > 75%
Chorology
or contorted shape, but in protected areas can grow straight and Native
large and edible (7x10mm)1, 7. Genetically the Arolla pine is close Outside its natural range, it is planted in parks and arboretums
to the Siberian stone pine (Pinus sibirica) and they can hybridise. especially in northern Europe8 .
Swiss stone pine mixed with larch (Larix decidua) in open subalpine
Map 2: High resolution distribution map estimating the relative probability of presence. woodland near Morgex (Valle dAosta, North-West Italy).
(Copyright Giovanni Caudullo: CC-BY)
References
[1] F. Bussotti, Pines of Silvicultural [13] A. Farjon, D. Filer, An Atlas of the Worlds
Importance, CABI, ed. (CABI, Wallingford, Conifers: An Analysis of their Distribution,
UK, 2002), pp. 5052. Biogeography, Diversity and Conservation
Swiss stone pines at the limit of tree vegetation along the Aletsch Glacier (Valais, Switzerland). [2] A. Farjon, The IUCN Red List of Threatened Status (Brill, 2013).
(Copyright Jo Simon, www.flickr.com: CC-BY) Species (2013), pp. 42349/0+. [14] B. Huntley, Journal of Vegetation Science
[3] M. Ulber, F. Gugerli, G. Bozic, EUFORGEN 1, 507 (1990).
Importance and Usage particular ski runs, ski lifts, roads and parking lots2 . Damage Technical Guidelines for genetic
conservation and use for Swiss stone pine
[15] A. Farjon, A handbook of the worlds
conifers (Brill, Leiden, 2010).
Arolla pine is not an important timber tree economically, as it to young seedlings by grazing animals can create gaps in the (Pinus cembra) (Bioversity International, [16] S. Boden, P. Pyttel, C. S. Eastaugh, iForest -
2004). Biogeosciences and Forestry 3, 82 (2010).
grows slowly and with irregular shapes, so forestry practises tend age distribution3 . Deep snow layers lasting until late winter or [4] F. H. Schweingruber, C. Wirth, Old trees [17] P. Vittoz, B. Rulence, T. Largey, F.
to favour other species in alpine habitats, such as the larch2, 15 . This spring causes browning needle diseases by the snow mold fungi and the meaning of old (Springer, Berlin Frelchoux, Arctic, Antarctic, and Alpine
Heidelberg, 2009), pp. 3554. Research 40, 225 (2008).
pine has as a more ecological and protection function for slopes Phacidium infestans, Gremmeniella abietina and Herpotrichia [5] L. Susmel, Monti e boschi 11/12 (1954). [18] R. Del Favero, I boschi delle regioni alpine
and valleys against avalanches and soil erosion1-3 . The wood has juniperi, which lead to mortality of young plants (seedlings)22-24 . [6] J. E. Eckenwalder, Conifers of the World: italiane (Cleup, Padova, 2004).
In the Alps, the larch bud moth Zeiraphera diniana has a species The Complete Reference (Timber Press, [19] L. Masutti, Italian Journal of Forest and
yellowish sapwood and reddish heartwood with a strong aromatic 2009). Mountain Environments 57, 437 (2002).
odour and it is very high quality, light, easy to work and durable, as form genetically differentiated and specialised for defoliating the [7] C. J. Earle, The gymnosperm database [20] A. Rolando, L. Carisio, Journal fr
Arolla pine25 . However, the sporadic outbreaks do not influence (2015). http://www.conifers.org Ornithologie 144, 69 (2003).
it is resistant to woodworm8 . It was overused for centuries in alpine [8] P. Schtt, Enzyklopdie der Holzgewchse:
the presence of the pine or its dominance in mixed forest with [21] A. A. Ali, C. Carcaillet, B. Talon, P. Roiron,
areas as firewood, and for furniture and building construction. Handbuch und Atlas der Dendrologie, J.-F. Terral, Journal of Biogeography 32,
European larch26, 27. A. Roloff, H. Weisgerber, U. M. Lang, B. 1659 (2005).
Now it is principally used for handicraft (turnery, carvings, toys, Stimm, P. Schtt, eds. (Wiley-Vch Verlag,
Weinheim, 1994), vol. 3. [22] M.-H. Li, J. Yang, Annals of Forest Science
marquetry) and in a minor way it is still used in carpentry, traditional [9] G. G. Goncharenko, V. E. Padutov, A. E. Silin,
61, 319 (2004).
houses, and flooring. Pine nuts are tasty and rich in nutrients8 . They Plant Systematics and Evolution 182, [23] M. Schneider, C. R. Grnig, O. Holdenrieder,
121 (1992). T. N. Sieber, Mycological Research 113,
were used more in the past: now Arolla pine is rarely cultivated [10] W. B. Critchfield, E. L. Little, Geographic
887 (2009).
for its seeds, as it is difficult to harvest due to the soft, resinous distribution of the pines of the world, [24] J. Senn, Forest Pathology 29, 65 (1999).
no. 991 (U.S. Dept. of Agriculture, Forest [25] I. Emelianov, J. Mallet, W. Baltensweiler,
and closed cone scales1, 2 . Cones can be used to flavour alcoholic Service, Washington, D.C., 1966). Heredity 75, 416 (1995).
distillates1 . From cones, needles, buds and branches an essential [11] J. Jalas, J. Suominen, Atlas Florae [26] P. Nola, M. Morales, R. Motta, R. Villalba,
Europaeae: distribution of vascular plants Trees 20, 371 (2006).
oil can be extracted and used in natural medicine and as essence. in Europe Vol. 2 Gymnospermae (Pinaceae
to Ephedraceae) (Committee for Mapping [27] R. Motta, M. Morales, P. Nola, Annals of
Some cultivars with different habits and needle colours have been the Flora of Europe and Societas Biologica Forest Science 63, 739 (2006).
selected for ornamental purposes, used principally in Northern and Fennica Vanamo, Helsinki, 1973). [28] EUFORGEN, Distribution map of swiss
[12] H. Meusel, E. Jger, eds., Vergleichende stone pine (Pinus cembra) (2010).
Eastern Europe, where late frosts are less frequent15 . Chorologie der Zentraleuropischen Flora www.euforgen.org.
- Band I, II, III (Gustav Fischer Verlag,
Threats and Diseases Jena, 1998).
Pinus halepensis and Pinus brutiain Europe: distribution, habitat, usage and threats
A. Mauri, M. Di Leo, D. de Rigo, G. Caudullo
Pinus halepensis Miller (Aleppo pine) and Pinus brutia Ten. (Turkish or Calabrian pine) are drought tolerant and fast
growing coniferous species native of the Mediterranean region. P. halepensis widely covers the Mediterranean coasts
concentrating in the western side of the basin, while P. brutia is located mainly on the eastern coasts. They are commonly
found in coastal zones, and because of their drought tolerance, are well adapted to dry summer conditions. They are
among the species most affected by wildfires in Europe, although they are fire resilient trees due to the high production
of serotinous cones that favour a quick post-fire regeneration. These species have been widely planted between the
thirties and seventies in Mediterranean areas for soil protection and wind breaks near the coasts.
Aleppo pine (Pinus halepensis Miller) and Turkish pine (Pinus
brutia Ten.) are two systematically close tree species, which can Frequency
naturally hybridize where they co-occur. Although some authors < 25%
25% - 50%
consider them as subspecies, in this chapter they are described as 50% - 75%
two separate pines1 . P. halepensis and P. brutia reach heights up to > 75%
Chorology
20 and 35m respectively1, 2 . The diameter of the trunk ranges from Native
Map 2: High resolution distribution map estimating the relative probability of presence for Pinus halepensis. Male flowers of Pinus halepensis.
Copyright Victor M. Vicente Selvas, commons.wikimedia.org: PD)
Frequency
< 25%
25% - 50%
50% - 75% Uncertain, no-data
> 75%
Chorology Tundra, cold desert
Native
Negligible survivability
Low survivability
Mid-low survivability
Medium survivability
Mid-high survivability
High survivability
Map 1-B: Plot distribution and simplified chorology map for Pinus brutia.
Caption: Frequency of Pinus brutia within the field observations as reported
by the National Forest Inventories. The chorology of the native spatial range
for P. brutia is derived after Critchfield and Little, and EUFORGEN24, 26 .
[13] M. Vennetier, et al., Sciences Eaux & (Pinus brutia) (2008). www.euforgen.org.
Territoires, la revue dIrstea 44, 49 (2005).
a big tree closely related to Pinus mugo, as shown by molecular > 75%
Chorology
markers which indicate the absence of species differentiation4, 5; Native
Map 2: High resolution distribution map estimating the relative probability of presence. Cluster of male pollen cones at the top of the shoot.
(Copyright Crusier, commons.wikimedia.org: CC-BY)
Open forest of Pyrenean pine (Pinus uncinata) in the karst Larra-Belagua massif (Navarra, North Spain).
(Copyright Alfonso San Miguel: CC-BY)
this plant has a great role in preventing torrents and avalanche Threats and Diseases
erosions on high mountains. Even though it grows at high altitudes, the dwarf mountain
The wood is elastic but hard, suitable for manufacturing pine is threatened by some pathogenic fungi associated with
small items and valuable as fuel. There is a large number of root-rot, but significantly affecting the living trees in adjacent
cultivars used in horticulture and it is grown in gardens for forests26 . Other fungi colonise the needles, the bark and needles27.
decorative planting2, 6, 9, 24 . Pine needles are filled with vitamin Insect pests are not dangerous. But actually the biggest threat
C and carotene. Beverages made out of them are recommended for Pinus mugo is humans. Pine scrublands were cut and burnt
to reinforce the immune system, if one has a cold, and to cure in order to enlarge pastures, especially since the Middle Ages
scurvy. Needles should be used fresh, if possible moments expansion of mountain animal husbandry. Given its low stature
after collecting, because they can completely lose their healing and scrubland density, dwarf pine habitus may favour the spread
properties after a year2, 12 . Syrups and liquors are commonly of fires; hence, frequent human-caused fires may eliminate
obtained with cones and buds. Essential oil distilled from the them28 . This is why the dwarf pine has become extinct on several
leaves exhibits good antioxidative activity in lipophilic media25 . mountains in Central Europe and the Balkan Peninsula, although
in recent decades land use changes have allowed the reversed
process of invasion by dwarf pine in abandoned grasslands29, 30 .
References
[1] H. Gaussen, V. H. Heywood, A. O. Chater, [19] R. Avigliano, G. D. Anastasio, S. Improta,
Flora Europaea, Volume 1: Psilotaceae M. Peresani, C. Ravazzi, Journal of
to Platanaceae, T. G. Tutin, et al., eds. Quaternary Science 15, 789 (2000).
(Cambridge University Press, 1993), pp. [20] H. Walter, S.-W. Breckle, Ecological
4044, second edn. Systems of the Geobiosphere (Springer
[2] M. Vidakovic, Tree Physiology 12, 319 Berlin Heidelberg, 1989), pp. 1140.
(1993). [21] J. ibk, D. Dt, I. ibkov, D. Pukajov,
[3] A. Farjon, A handbook of the worlds Phytocoenologia 38, 221 (2008).
conifers (Brill, Leiden, 2010). [22] A. Dini, M. Jankovi, Vegetacija Srbije,
[4] M. Heuertz, et al., Journal of Biogeography D. kori, ed. (Srpska Akademija Nauka
37, 541 (2010). i Umetnosti, Beograd, 2006), vol. 2, pp.
[5] I. Monteleone, D. Ferrazzini, P. Belletti, 201211.
Silva Fennica 40, 391 (2006). [23] V. Stefanovi, Fitocenologija: sa pregledom
[6] G. Krssmann, Manual of Cultivated umskih fitocenoza Jugoslavije (Svjetlost,
Conifers (Timber Press, Portland, 1985). Sarajevo, 1986).
[7] W. Prus-Gowacki, E. Bajus, H. Ratyska, [24] W. Dallimore, A. B. Jackson, A Handbook
Acta Societatis Botanicorum Poloniae 63, of Coniferae and Ginkgoaceae (Edward
269 (1998). Arnold Ltd, London, 1961), third edn.
[8] F. Boguni, S. Siljak-Yakovlev, E. Muratovi, [25] J. Grassmann, S. Hippeli, R. Vollmann, E. F.
F. Pustahija, S. Medjedovi, Annals of Elstner, Journal of Agricultural and Food
Forest Science 68, 179 (2011). Chemistry 51, 7576 (2003).
[9] J. Hamernk, I. Musil, Journal of Forest [26] M. Bendel, F. Kienast, D. Rigling, H.
Science 53, 253 (2007). Bugmann, Canadian Journal of Forest
Research 36, 2666 (2006).
[10] H. H. Ellenberg, Vegetation Ecology of
Central Europe (Cambridge University [27] M. Uupli, Patologija: umskog i
Press, 2009), fourth edn. ukrasnog drvea (umarski fakultet
Univerziteta u Sarajevu, Sarajevo, 1996).
[11] G. Hegi, ed., Illustrierte Flora von
Mitteleuropa, vol. 1 (Lehmann, Mnchen, [28] B. Leys, et al., Quaternary Science Reviews
1906). 90, 60 (2014).
[12] B. Jovanovi, Dendrologija (Univerzitetska [29] S. Dullinger, T. Dirnbck, G. Grabherr,
tampa, Beograd, 2000). Arctic, Antarctic, and Alpine Research 35,
434 (2003).
[13] K. Christensen, Nordic Journal of Botany
7, 383 (1987). [30] I. Horvat, V. Glava, H. H. Ellenberg,
Scrubland of dwarf mountain pines in the karstic Valley of Five Polish Lakes (Dolina Piciu Staww Polskich) Vegetation Sdosteuropas, vol. 4 of
[14] Z. Debreczy, I. Racz, Conifers Around the Geobotanica selecta (Gustav Fischer
in Tatra National Park (Gmina Bukowina Tatrzaska, South Poland). World: Conifers of the Temperate Zones Verlag, Jena, 1974).
(Copyright Nova, commons.wikimedia.org: CC-BY) and Adjacent Regions, vol. 1 (Dendropress,
Budapest, 2011). [31] W. B. Critchfield, E. L. Little, Geographic
Autoecology diagrams based on harmonised distribution of the pines of the world,
Field data in Europe (including absences) Observed presences in Europe field observations from forest plots.
[15] C. Carcaillet, A. A. Ali, N. Fauvart, P. Roiron, no. 991 (U.S. Dept. of Agriculture, Forest
J.-F. Terral, Travaux scientifiques du Parc Service, Washington, D.C., 1966).
national de la Vanoise 24, 57 (2009).
[32] J. Jalas, J. Suominen, Atlas Florae
[16] A. Farjon, The IUCN Red List of Threatened Europaeae: distribution of vascular plants
Species (2013), pp. 18153856/0+. in Europe Vol. 2 Gymnospermae (Pinaceae
Average temperature of the coldest month (C)
Distribution Map 1: Plot distribution and simplified chorology map for Pinus nigra.
Frequency of Pinus nigra occurrences within the field observations as
The past distribution of black pine in Europe is difficult reported by the National Forest Inventories. The chorology of the native
to reconstruct. This because past occurrences based on both spatial range for P. nigra is derived after EUFORGEN48 .
pollen and charcoal (widely used to reconstruct past species
distribution) cannot be easily recognised at the species level8 . spread conifer species in the Balkans and Asia Minor. Its widest
However, more localized studies mainly based on macrofossils distribution worldwide is in Turkey, with more than 2.5 million
suggest that large populations of black pine were already present hectares12 . Outside Europe, it was also introduced in the United
during the late Pleistocene and the Holocene in the north-western States (where it is known as Austrian pine) in 175913 , and has now Black pine grown on vulcanic rocky soils in Etna Vulcane (Sicily, South Italy).
Mediterranean basin (see9 for a review). These populations are become naturalised in parts of New England and the Great Lake (Copyright Alfie Ianni, www.flickr.com: CC-BY)
thought to have followed a substantial decrease during the States14 . As a result of climate warming the future distribution of
Holocene as a consequence of climate warming at the onset of black pine is thought to change considerably but the response is Habitat and Ecology
the Holocene as well as increased human activities during the likely to be different depending on the geographic region15 . In the Black pine stands exist at altitudes ranging from 350m in
last millennia9 . This led to the current fragmented distribution Mediterranean regions climate warming increases water stress Italy to 2200m in the Taurus Mountains, the optimal altitudinal
of black pine extending from North-Western Africa through and thus has a negative influence on the growth of this species16, range being between 800 to 1500m5 . It can grow on a variety
southern Europe to Asia Minor10, 11 . Black pine presently covers 17
, whereas in central Europe climate amelioration is thought to of soils, from podzolic sands to limestone, often dependent on
more than 3.5 million hectares4 , making it one of the most wide- lead to an expansion18, 19 . Two main subspecies of black pine region and climate21 . The Austrian pine subspecies is more able
to tolerate exposed chalk and limestone than Corsican pine23 .
However Corsican Pine is more often found in coastal areas as
it is more resistant to salt wind than most other pine species21 .
Black pine can grow in both extremely dry and humid habitats
Uncertain, no-data
with considerable tolerance of temperature fluctuations11 . It is a
Marginal/no presence < 5% light-demanding species, but it shows higher shade tolerance than
Low presence 5% - 10% Scots pine (Pinus sylvestris)24 . It is resistant to drought and wind4 .
It grows in pure stands or in association with other broadleaved or
Mid-low presence 10% - 30%
conifer species, in particular Pinus sylvestris. It is also commonly
Medium presence 30% - 50% found in association with other pines such as dwarf mountain
Mid-high presence 50% - 70% pine (Pinus mugo), Aleppo pine (Pinus halepensis), Italian stone
pine (Pinus pinea) and Heldreich pine (Pinus heldreichii)4, 21, 22 .
High presence 70% - 90%
Very-high presence > 90%
Management 175, 103 (2003). [28] CABI, Pinus pinaster (maritime pine)
[13] R. Ala, et al., Regiones de procedencia (2015). Invasive Species Compendium.
Pinus pinaster Aiton (Ministerio de Medio http://www.cabi.org
Annual precipitation (mm)
long, heavy, with easily detachable wings and ineffective for wind hybridisation with other pines is known10, 14, 15 . It is considered a
dispersal. Stone pine presents mast seeding with a significant heliophilous, xerophilous and thermophilous pine16 , that can
variation in seeds production across the years1-4 . withstand slight shade during its first stages but which requires
abundant light for fructification in maturity17. Concerning annual
Distribution average precipitation, the minimum requirement is for around
The natural range of stone pine is uncertain and difficult to Map 1: Plot distribution and simplified chorology map for Pinus pinea.
Frequency of Pinus pinea occurrences within the field observations as 250mm but the optimum is considered to be 600mm17. Stone
establish due to a long history of planting. This species was reported by the National Forest Inventories. The chorology of the native pine is well adapted to coastal thermo-Mediterranean areas where
largely distributed in Europe during the last thousand years, as spatial range for P. pinea is derived after EUFORGEN25 .
frost damage is not a relevant issue; however it also thrives well
it was introduced throughout the Mediterranean region for its in sandy continental areas of central Spain with wide yearly and
edible seeds1, 5. The earliest evidence of the human use of this daily thermal oscillations and where night frosts are frequent for
species were recently found in Gibraltar and dated to 49 200 several months of the year17, 18 . Concerning the soil there are no
years before present6. Currently, the stone pine has a scattered special requirements; it tolerates calcareous soils4 , but it prefers
distribution around the Mediterranean basin, from Portugal to siliceous and sandy soils with acid or sub-acid reaction. It also
Syria, and it is more abundant in south-western Europe (Iberian Habitat and Ecology presents limitations in clay soils due to its inability to develop a
Peninsula, South France, Italy), where its regeneration is natural. The stone pine, one of the most characteristic trees of proper root system in these conditions. The soil pH in its locations
It is also present along the shores of the Black Sea5, 8-1. This the Mediterranean flora, occupies a broad range of climate and can range from 4 to 917, 19 .
pine occurs mainly on coastal areas, except in Spain and soil conditions along the Mediterranean basin. Despite this, it
Portugal where it grows naturally at some distance from the has been identified as having very low genetic variation and no
sea.
Uncertain, no-data
Marginal/no presence < 5%
Low presence 5% - 10%
Mid-low presence 10% - 30%
Medium presence 30% - 50%
Mid-high presence 50% - 70%
High presence 70% - 90%
Very-high presence > 90%
Stone pine forest on coastal sand dunes near Trafalgar Cape (Cdiz, Andalusia, South-West Spain).
(Copyright Alfonso San Miguel: CC-BY)
References
[1] J. E. Eckenwalder, Conifers of the World: [14] D. M. Richardson, P. W. Rundel, Ecology and
The Complete Reference (Timber Press, Biogeography of Pinus, D. M. Richardson,
2009). ed. (Cambridge University Press, 1998).
[2] C. J. Earle, The gymnosperm database [15] G. G. Vendramin, et al., Evolution 62, 680
(2015). http://www.conifers.org (2008).
[3] O. Johnson, D. More, Collins tree guide [16] J. Retana, et al., Post-Fire Management
(Collins, 2006). and Restoration of Southern European
[4] G. Montero, R. Calama, R. Ruiz-Peinado, Forests, F. Moreira, M. Arianoutsou, P.
Selvicultura de Pinus pinea (Instituto Corona, J. De las Heras, eds. (Springer
Nacional de Investigacin y Tecnologa Netherlands, 2012), vol. 24 of Managing
Agraria y Alimentaria (Espaa), 2008), pp. Forest Ecosystems, pp. 151170.
431470. [17] G. Borrero Fernndez, El pino pionero
[5] A. Farjon, D. Filer, An Atlas of the Worlds (Pinus pinea L.) en Andaluca: Ecologa,
Conifers: An Analysis of their Distribution, distribucin y selvicultura (Consejera de
Biogeography, Diversity and Conservation Medio Ambiente. Direccin General de
Status (Brill, 2013). Gestin del Medio Natural, Sevilla, 2004).
[6] C. Finlayson, et al., Nature 443, 850 [18] M. Pardos, J. Climent, H. Almeida, R. Calama,
(2006). Annals of Forest Science 71, 551 (2014)
[7] M. Barbro, R. Loisel, P. Quzel, D. M. Richardson, [19] A. Cutini, Pines of Silvicultural Importance,
F. Romane, Ecology and Biogeography of Pinus, CABI, ed. (CABI, Wallingford, UK, 2002),
D. M. Richardson, ed. (Cambridge University pp. 329343.
Press, 1998), pp. 153170. [20] R. Calama, G. Madrigal, J. A. Candela, G.
[8] A. Farjon, The IUCN Red List of Threatened Montero, Investigacin Agraria: Sistemas y
Species (2013), pp. 42391/0+. Recursos Forestales 16, 241 (2007).
[9] J. Jalas, J. Suominen, Atlas Florae [21] A. Boutheina, M. H. El Aouni, P. Balandier,
Europaeae: distribution of vascular plants Influence of stand and tree attributes and
in Europe Vol. 2 Gymnospermae (Pinaceae silviculture on cone and seed productions
to Ephedraceae) (Committee for Mapping in forests of Pinus pinea L. in northern
the Flora of Europe and Societas Biologica Tunisia, Options Mditerranennes , Series
Fennica Vanamo, Helsinki, 1973). A: Mediterranean Seminars, No. 105
(CIHEAM, FAO, INIA, IRTA, CESEFOR, CTFC,
[10] B. Fady, S. Fineschi, G. G. Vendramin, Zaragoza, 2013), pp. 914.
EUFORGEN Technical Guidelines for
genetic conservation and use for Italian [22] M. Bracalini, et al., Journal of Economic
stone pine (Pinus pinea) (International Entomology 106, 229 (2013).
Plant Genetic Resources Institute, 2008). [23] I. M. zankaya, S. N. Balay, C. Bucak,
[11] W. B. Critchfield, E. L. Little, Geographic Effects of pests and diseases on stone pine
distribution of the pines of the world, (Pinus pinea L.) conelet losses in Kozak
no. 991 (U.S. Dept. of Agriculture, Forest catchment area, Options Mditerranennes,
Service, Washington, D.C., 1966). Series A: Mediterranean Seminars, No. 105
(CIHEAM, FAO, INIA, IRTA, CESEFOR, CTFC,
[12] G. L. Shaughnessy, Historical ecology of Zaragoza, 2013), pp. 2933.
alien woody plants in the vicinity of cape
town, south africa, Ph.D. thesis, University [24] EPPO, EPPO Global Database (2015).
of Cape Town (1980). https://gd.eppo.int
[13] N. T. Mirov, P. M. Iloff, Journal of the [25] EUFORGEN, Distribution map of italian
American Pharmaceutical Association 44, stone pine (Pinus pinea) (2008).
186 (2006). www.euforgen.org.
Pine nuts fresh from the cone are covered with a black powder.
(Copyright Eran Finkle, www.flickr.com: CC-BY)
and a number of sub-species and varieties exist. However, it can often be easily recognised because of its distinctive Chorology
Native
orange-red coloured bark. Scots pine is a pioneer species, frost and drought tolerant and able to grow on very poor soils,
so it can be found in many ecologically diverse habitats. Its timber is valued for its good strength to weight ratio and it
is both commercially and culturally a very important species in a number of European countries, particularly in the more
northerly regions.
Pinus sylvestris L. (Scots pine) is a medium-sized conifer. It comprising over 20% of the productive forest area9 . The tree
reaches 23-27m in height on average but can attain over 40m1 varies widely in form throughout its range and there is debate
and live for 400 years or more: one tree in Lapland is reported to over how many separate subspecies should be recognised8 . Its
be over 750 years old2 . The bark on the upper part of the stem modern genetic diversity is probably caused by its isolation in a
develops a distinct reddish-orange colour while the lower part is number of glacial refugia during the last ice age10 .
furrowed brown to grey-brown and becomes deeply fissured. Its
blue-green or grey-green needles are in pairs, generally slightly
twisted and are around 5-7cm long3 . They stay on the tree for at
least 2, and in some cases up to 6 years1 .The needles are adapted Map 1: Plot distribution and simplified chorology map for Pinus sylvestris.
to deal with cold and drought, having imbedded stomata and a Frequency of Pinus sylvestris occurrences within the field observations as
waxy layer on the thick-walled epidermis to protect the needle reported by the National Forest Inventories. The chorology of the native
spatial range for P. sylvestris is derived after EUFORGEN32 .
from water loss4 . It is a wind-pollinated species and is normally
monoecious but mature trees may very occasionally bear only
male or only female flowers5 . The male flowers cluster at the
base of new shoots and are yellow or pink; the female flowers
occur at the tips of new, strong shoots and develop a rose-purple
shade. The cones develop the year following pollination and are
conic-oblong 5-8cm in size3, 6 . They require alternating periods of
dry and wet weather to open and shed the winged seeds, which
can be dispersed some way from the parent tree7.
Distribution
Scots pine is the most widespread species of the Pinus genus in
the world, and the second most widespread conifer after common
juniper (Juniperus communis)8 . It occupies a range from Spain in the
Maturing male flowers clastered at the base of a new shoot.
west to the far east of Russia. In terms of latitude it can be found (Copyright Stuart Caie, www.flickr.com: CC-BY)
from northern Scandinavia (70N) to the mountains of Sierra Nevada
in southern Spain (37N). It grows at a wide range of elevations, Habitat and Ecology
from sea level in the northern parts of its range to over 2600m in It is a light-demanding pioneer species and can colonise
the Caucasus. It has also been widely planted in the United States recently disturbed sites if competition and grazing pressure
(where it is referred to as Scotch pine), especially in the Northeast, are low11 . It grows mainly on sunny to partially shaded, usually
Lake States, Central States and the Pacific Northwest7. The upper part of the trunk is a distinctive reddish-orange colour. nutrient-poor sites12 . With a pronounced drought tolerance and
(Copyright Alfonso San Miguel: CC-BY)
In Europe, Scots pine forests now exceed 28 million hectares, also good frost resistance, it is very undemanding as to site
and water supply and can grow on the poorest sandy soils, even
colonising acid highland moors13 . However, it cannot cope with
atmospheric pollution or salty sea winds14 and on fertile sites
it is often outcompeted by other species (usually spruce or
Uncertain, no-data broadleaved species)11, 15 . It requires a period of winter chilling to
Marginal/no presence < 5% break autumn dormancy, and starts to grow in the spring when
temperatures reach about 5C4, 5 . Under conditions of a warming
Low presence 5% - 10%
climate it is likely to increase its presence in the north, but decline
Mid-low presence 10% - 30% in the southern parts of its range16-19 .
Medium presence 30% - 50% It frequently grows in large single-species stands, but across
its huge range it may also be found with most of the boreal
Mid-high presence 50% - 70%
species of Europe and Asia20 . In Europe it can be found growing
High presence 70% - 90% with broadleaved trees such as oaks (Quercus petraea, Quercus
Very-high presence > 90% robur), beech (Fagus sylvatica) and birch (Betula pendula), and
other conifers including spruce (Picea abies), larch (Larix decidua),
fir (Abies alba) and other pines (Pinus nigra, Pinus uncinata)9 , but
no single species or species group is associated with it over its
entire range15 .
Annual precipitation (mm)
Map 2: High resolution distribution map estimating the relative probability of presence. Native Scots pine woodlands in Glen Affric, Scotland.
(Forestry Commission, www.forestry.gov.uk: Crown Copyright)
Uncertain, no-data
Tundra, cold desert
Negligible survivability
Low survivability
Mid-low survivability
Medium survivability
Mid-high survivability
Cones develop in one year and are 5-8 cm long. High survivability
(Copyright Alfonso San Miguel: CC-BY)
References
[1] A. Praciak, et al., The CABI encyclopedia of [17] M. Benito Garzn, R. Snchez de Dios, H.
forest trees (CABI, Oxfordshire, UK, 2013). Sainz Ollero, Applied Vegetation Science
[2] T. H. Wallenius, H. Kauhanen, H. Herva, J. 11, 169 (2008).
Pennanen, Canadian Journal of Forest [18] L. Matas, A. S. Jump, Forest Ecology and
Research 40, 2027 (2010). Management 282, 10 (2012).
[3] O. Johnson, D. More, Collins tree guide [19] G. E. Rehfeldt, et al., Global Change
(Collins, 2006). Biology 8, 912 (2002).
[4] U. K. Krakau, M. Liesebach, T. Aronen, M. [20] D. M. Richardson, Ecology and
A. Lelu-Walter, V. Schneck, Forest Tree biogeography of Pinus (Cambridge
Breeding in Europe, L. E. Pques, ed. University Press, 1998).
(Springer Netherlands, 2013), vol. 25 of [21] J. E. Milner, The tree book : the
Managing Forest Ecosystems, chap. 6, pp. indispensable guide to tree facts, crafts
267323. and lore (Collins & Brown, 1992).
[5] A. Carlisle, A. H. F. Brown, Journal of [22] M. Lindholm, et al., Silva Fennica 34, 317
Ecology 56, 269 (1968). (2000).
[6] A. F. Mitchell, A field guide to the trees [23] B. Langstrm, E. Annila, C. Hellqvist, M.
of Britain and northern Europe (Collins, Varama, P. Niemel, Scandinavian Journal
1974). of Forest Research 16, 342 (2001).
[7] D. D. Skilling, Pinus sylvestris (Scotch [24] T. Virtanen, S. Neuvonen, A. Nikula, M.
Pine), Agriculture Handbook 654 (U.S. Varama, P. Niemel, Silva Fennica 30,
Department of Agriculture, Forest Service, 169 (1996).
Washington, DC., 1990), pp. 10001017.
[25] J. I. Barredo, et al., EPPO Bulletin 45, 273
[8] J. E. Eckenwalder, Conifers of the World: (2015).
The Complete Reference (Timber Press,
2009). [26] CABI, Hylobius abietis (large pine weevil)
(2015). Invasive Species Compendium.
[9] W. L. Mason, R. Ala, Investigacin agraria. http://www.cabi.org
Sistemas y recursos forestales 9, 317 (2000).
[27] R. Kirschner, D. Begerow, F. Oberwinkler,
[10] R. Cheddadi, et al., Global Ecology and Mycological Research 105, 1403 (2001).
Biogeography 15, 271 (2006).
[28] R. Jankowiak, J. Hilszczaski, Acta
Ancient large tree in Sierra de Guadarrama, Spain. [11] C. Mtys, Ackzell, C. J. A. Samuel, Societatis Botanicorum Poloniae 74, 345
(Copyright Alfonso San Miguel: CC-BY) EUFORGEN Technical guidelines for (2011).
genetic conservation and use for Scots
Autoecology diagrams based on harmonised pine (Pinus sylvestris) (2003). [29] D. de Rigo, et al., Scientific Topics Focus 2,
Field data in Europe (including absences) Observed presences in Europe mri10a15+ (2016).
field observations from forest plots. [12] A. Farjon, A handbook of the worlds
conifers (Brill, 2010). [30] L. Giordano, M. Garbelotto, G. Nicolotti, P.
Gonthier, Mycological Progress 12, 127
[13] M. Gardner, The IUCN Red List of (2013).
Threatened Species (2013), pp. 42418/0+.
[31] R. Drenkhan, J. Hantula, M. Vuorinen, L.
Average temperature of the coldest month (C)
[14] P. S. Savill, The silviculture of trees used in Jankovsk, M. M. Mller, European Journal
British forestry (CABI, 2013). of Plant Pathology 136, 71 (2013).
[15] D. Kelly, A. Connolly, Forest Systems 9, [32] EUFORGEN, Distribution map of scots pine
15 (2000).
Annual precipitation (mm)
Distribution
This tree is native in riparian steppe and coastal forest
communities of central and southern Europe. It occurs over a wide
range, from North Africa to Poland and from the Iberian Peninsula Map 1: Plot distribution and simplified chorology map for Populus alba.
Frequency of Populus alba occurrences within the field observations as
to western Siberia and Central Asia2, 6-9 . It was introduced in the reported by the National Forest Inventories. The chorology of the native
United States in the 18th century as a shade and ornamental tree spatial range for P. alba is derived after Isebrands and Richardson16 .
and more recently in all other continents, becoming naturalised in
many areas and invasive in some countries10-14 . the European mountain systems22 . In these critical areas, white
poplar complements key forest ecosystem services such as soil
Habitat and Ecology stabilisation and watershed protection23 . More generally, white
The white poplar occurs spontaneously along river valleys poplar is used for erosion control along river banks and roadsides,
in warm temperate and Mediterranean zones where soil-water windbreaks and land reclamation. Silvoarable agroforestry with Creamy white bark with small black diamonds.
(Copyright Silvano Radivo, www.actaplantarum.org: AP)
is available15, 16 . It is a fast-growing and light demanding tree, this species24 may be exploited in Mediterranean areas with high
colonising woodland edges and open sunny habitats, including potential soil erosion, also considering the effectiveness of its
meadows, wetlands and riparian zones. It grows in most sites, cover-management on erosion rates25, 26 . Thanks to its salt and Threats and Diseases
tolerating from waterlogged to drought habitats and from sandy soil tolerance, it is also used near coasts in windbreaks Like other poplars, it hosts a large number of insects, but only a
acid to strong alkaline soils, but developing in shrub form in against salty winds and for dune stabilisation2, 15 . It is further few of those need to be controlled especially in plantations. Among
extreme conditions17. It performs with high growing rates on exploited for phytoremediation with the hybrids Populus alba x the leaf defoliators, the main ones are the moth Hyphantria cunea
optimal sites, characterised by good water availability and well- tremula and Populus tremula x alba 27. The wood is not of high and the large poplar-leaf beetle Chrysomela populi. Wood diseases
textured soils that have neutral-alkaline pH and which are rich in quality, especially from natural stands, being woolly in texture, of can occasionally be caused by the goat moth Cossus cossus and
nutrients3, 15, 17, 18 . Reproduction is primarily by vegetative means, low flammability, not durable, very light and soft, so it is suitable the longhorn beetles of genus Saperda, even if their xylophagous
through root suckers. They arise from adventitious buds on the only for local and artisan use. However, specialised plantations caterpillars are found mainly in other poplar species. The soil
lateral root system, which can grow as far as 30-50m from the
plant. New plants may also establish through tree fragments,
which can easily root in suitable environments. Suckers from
a single tree can quickly develop a dense and large colony,
shading out competitive vegetation10, 11, 18 . Mature trees also
produce abundant wind-dispersed seeds that may be carried long
distances11 . The white poplar hybridises naturally with Eurasian
aspen (Populus tremula). The resulting hybrids are known as
grey poplars (Populus x canescens), which is a morphologically
intermediate species, but which exhibits more vigour than
either parent2, 4, 5 . This poplar is a common species of floodplain
forests in early- to mid-seral vegetation communities10 . In
central and southern Europe it dominates and co-dominates in
riparian woodland of the central-western Mediterranean zone
with willows (Salix alba, Salix fragilis), black poplar (Populus
nigra), and alder (Alnus glutinosa). In the wooded steppe zone
it is dominant on sandy soils having originated from alluvial
deposits and on sand dunes bordering riverine gallery forests9 .
It occurs also as a secondary species in hygrophilous floodplain
forests of the temperate zone dominated by pedunculate oak
(Quercus robur), ashes (Fraxinus excelsior, Fraxinus angustifolia),
elms (Ulmus spp.) and alder (Alnus glutinosa), and in eastern
Mediterranean riparian forests dominated by oriental plane
(Platanus orientalis)19, 20 .
Distribution
Black poplar has a wide natural distribution. In Europe, it
can be found as far north as the British Isles and down to the
Mediterranean coast. At the southern extreme of its range it can
be found in parts of northern Africa and the Middle East. To the
east its range extends as far as Kazakstan and China1, 7, 9 . It is
also cultivated in India between 26 and 29N latitude and is Map 1: Plot distribution and simplified chorology map for Populus nigra.
Frequency of Populus nigra occurrences within the field observations as
naturalised in both North and South America1 . reported by the National Forest Inventories. The chorology of the native
spatial range for P. nigra is derived after EUFORGEN32 .
Habitat and Ecology
The black poplar is a tree species of floodplain forests5 , several breeding programmes across Europe, especially for
growing in riparian mixed forests together with white poplar obtaining the hybrid Populus x euramericana (P. deltoides x P.
(Populus alba L.), willow (Salix spp.), alder (Alnus spp.), maple nigra) 7, 10-12 . As other fast growing Salicaceae, this species may
(Acer spp.), elm (Ulmus spp.), and sometimes oak (Quercus spp.)7. have a multifunctional role for pollution mitigation , microclimate
It is a pioneer tree species6, 10 , and does not tolerate drought or regulation and improved structural and biological diversity in open
shade1 . It is an opportunistic species able to colonise new sites agricultural landscapes13 . The wide spatial distribution of black
after disturbances, and has a good tolerance to high water levels poplar overlaps with many areas in Europe subject to high erosion
and high temperatures during summer8 . It can be managed easily rates, including moist slopes with high drainage-area within
by coppicing1 . the European mountain systems14 . Here, this tree contributes
to relevant forest ecosystem services such as soil stabilisation
and watershed protection15 . It has also high ecological value in
Importance and Usage riparian floodplain ecosystems, frequently used as a windbreak
Both tree breeders and conservationists are aware of the or to control erosion along riverbanks1 . In Mediterranean areas
importance of black poplar8 . It is a highly valuable tree species with high potential soil erosion16 , silvoarable agroforestry
from an economic point of view: it is used as a parent pool for with this species17 may be exploited even considering the
Uncertain, no-data
Marginal/no presence < 5%
Low presence 5% - 10%
Mid-low presence 10% - 30%
Medium presence 30% - 50%
Mid-high presence 50% - 70%
High presence 70% - 90%
Very-high presence > 90%
Map 2: High resolution distribution map estimating the relative probability of presence. Male catkins with reddish anthers during pollination.
(Copyright Aldo De Bastiani, www.actaplantarum.org: AP)
Isolated poplars with autumn foliage in the rural area near Torresto (Len, North-West Spain).
(Copyright Alfonso San Miguel: CC-BY)
Rome, 2007).
[15] J. E. Norris, A. Di Iorio, A. Stokes, B. C.
Nicoll, A. Achim, Slope Stability and [32] EUFORGEN, Distribution map of black
Erosion Control: Ecotechnological poplar (Populus nigra) (2015).
Annual precipitation (mm)
ovate, with big wave-shaped teeth2, 3 . They flutter in the slightest Chorology
Native
breeze, constantly moving and rustling, so that trees can often be
heard but not seen. In spring the young leaves are coppery-brown
and turn to golden yellow in autumn, making it attractive in all
vegetative seasons1, 2 . The aspen is a dioecious tree. Flowers
are produced in February-March before the leaves appear2, 3 .
Male catkins are 5-10cm long, grey-brown, yellowish in mid-
March when shedding pollen. Female catkins are green, 5-6cm at
pollination, extending 10-12cm long at maturity in early summer
to bear 50-80 capsules each containing numerous tiny seeds
Young trees in Lapland, Finland..
embedded in downy fluff and dispersed by wind4 . The bark is (Copyright Mattivirtala, commons.wikimedia.org: CC0)
greenish-grey, smooth, wrinkled with diamond lenticels1, 2 .
woodland, especially with species casting a relatively light shade,
Distribution such as Scots pine (Pinus sylvestris) and birches (Betula spp.)11 .
This species is native to cool temperate and boreal regions Map 1: Plot distribution and simplified chorology map for Populus tremula. Eurasian aspen trees typically occur as scattered patches in the
Frequency of Populus tremula occurrences within the field observations midst of the conifer-dominated landscape. It is a disturbance-
of Europe and Asia. It is the second most widely distributed tree as reported by the National Forest Inventories. The chorology of the native
in the world, after Scots pine (Pinus sylvestris). Aspens range spatial range for P. tremula is derived after several sources5, 30, 31 . adapted species and is a coloniser of clear disturbed areas such
extends from Iceland and Ireland to Kamchatka, and from north as after fire, clear-cutting, wind-throw or defoliation. In such
of the Arctic Circle in Fennoscandia and Russia (growing at sea age of about 20 years when crown competition increases. After cases it may form large and more continuous stands9, 13 . Though
level), to Spain, Turkey, North Korea and northern Japan (growing that, its growth increment is slower and culminates at about 30 Eurasian aspen may produce enormous numbers of viable seeds,
up to 1900m in the Pyrenees)5-9 . There are also isolated glacial- years of age, and the average lifespan is 50-100 years8, 9, 11 . seedlings find it difficult to establish apart from under suitable
relict populations on the highest elevations of the Atlas Mountains The enormous wide natural range demonstrates its tolerance of environmental conditions or after a forest fire14 . Aspen maintains
in Algeria10 . Due to its wide distribution, many geographical races a high variety of climatic and habitat conditions, such as frost, its existing populations mainly through vegetative replacement
have been differentiated morphologically, and some of these shade, waterlogging, wind and weed competition. It also grows and expansion. The vegetative reproduction is guaranteed by root
forms are considered as sub-species8 . on a wide range of soils, from slightly dry to wet soils of poor suckers, which are abundantly produced on the shallow lateral
to rich nutrient status, although it favours moist soils with a high roots after an individual has been damaged or destroyed, e.g. by
Habitat and Ecology organic matter content and wet conditions8, 11, 12 . Light is more cutting, fire or diseases, leaving an open space exposed to sunlight.
Eurasian aspen is a light-demanding, rapidly growing important than soil conditions, even if, unlike many other poplars, it The trees growing from the suckers form clones and mature stands
broad-leaved tree. Its fast-growing habit continues until the is sufficiently shade-tolerant to be a stable part of a mixed stand reproduce vigorously by this vegetative means. Clones may live
thousands of years if new suckers continuously arise from the
original rootstock4, 9 . Eurasian aspen hybridises naturally with
white poplar (Populus alba) forming the grey poplar (Populus x
canescens), which is intermediate morphologically, but more
Uncertain, no-data vigorous than its parents8 . Artificial hybrids have been produced
with a number of other poplars. In particular Populus tremula x
Marginal/no presence < 5%
tremuloides, the hybrid with the North American quaking aspen
Low presence 5% - 10% (Populus tremuloides), is widely used for large-scale plantations
Mid-low presence 10% - 30% thanks to its stronger vigour and higher growth rates4, 15 .
Medium presence 30% - 50% Importance and Usage
Mid-high presence 50% - 70% Although its commercial importance is limited compared
High presence 70% - 90% with other tree species, aspen is often found to be a keystone
species due to its fundamental ecological importance for other
Very-high presence > 90%
species; e.g. herbivorous, saprophytic invertebrates, fungi and
lichens, birds, etc. It has more host-specific species than any other
boreal tree and is one of the most significant contributors to total
epiphyte diversity in the boreal forest9, 11, 16 . It is an attractive
species for ornamental purposes thanks to the colouration of
foliage8, 11 . The wood is not dense, like other poplars, and it is
mainly used for veneer and pulp for paper production, also for
good quality charcoal and chip-wood8, 11 . It is used as a biomass
crop for energy production because of its rapid growth. As a
pioneer species Eurasian aspen is often used for afforestation of
barren or degraded lands, and it is also planted as a shelterbelt
species thanks to its wind resistance8 . As other fast growing
Annual precipitation (mm)
The foliage flutters in the breeze so the trees are often heard even when not seen.
Map 2: High resolution distribution map estimating the relative probability of presence. (Copyright AnRo0002, commons.wikimedia.org: CC0)
120cm1 . The species mostly develops single, straight trunks 25% - 50%
50% - 75%
with a thin, smooth purplish-grey bark that becomes grey-brown > 75%
Chorology
with horizontal fissuring and peeling when old. Young trees grow Native
with a strong apical control developing a straight trunk and an
erect-pyramidal coniferous crown shape, becoming broader
and rounded on single old trees or conical in individuals in forest
stands2, 3 . Young shoots are shiny, pale grey to purplish-brown,
and have large, reddish brown and protruding ovoid-ellipsoid,
glabrous winterbuds at the branch ends arranged in whorl-
like form. Stem wounds produce a resin-like, amber coloured
odourless gum4, 5 . The leaves change colour from light green
in spring over dark green in summer, and to yellow, orange-red,
scarlet or pink in autumn. They are alternate, pendulous, simple Cutivated cherry tree in orchard.
and elliptic-ovate to obovate acute in shape. The leaf margins (Copyright Tara2, commons.wikimedia.org: CC-BY)
However, it favours deep fertile soils with a good water supply. The
tree does not tolerate heavy clays, waterlogged or poorly drained
sites and can be sensitive to drought1, 2, 5 . Main habitat type is
semi-shade, open deciduous woodland or scrubland especially at
edges, glades and clearings, where this tree essentially occurs as
a rare and scattered pioneer species1, 2 . The pioneer colonisation
strategy is realized as a first generation establishment via
seedling recruitment, potentially followed by sometimes extensive
Map 2: High resolution distribution map estimating the relative probability of presence. vegetative growth via root suckering. With its ability for coppice
Cherres are ripe from late spring until summer and are
dispersed principally by birds.
(Copyright Steven Gill, www.flickr.com: CC-BY)
Distribution
This plum tree is native to south-eastern Europe (Balkan
Peninsula, Crimea), western and middle Asia (Caucasus, Iran,
Iraq). It has been widely cultivated for its fruits in Asia Minor and
the Caucasus for millennia, spreading in the Mediterranean area
and Balkans since 200 BC and later in the rest of Europe. More
recently, it is present on all continents, naturalised widely outside Reddish fleshy cherries can be used for a variety of culinary purposes.
(Copyright Phil Sellens, www.flickr.com: CC-BY)
its native range throughout temperate areas1-7. In Australia
and in New Zealand it is considered a weed species8-10 . Due
to its high variability and easy hybridisation with other Prunus Importance and Usage
species, different geographical subspecies have been described It is considered, probably together with the blackthorn
White flowers blossoming in spring.
and its taxonomic subdivision is rather confusing and still under (Prunus spinosa), to be one of the ancestral lineages leading (Copyright Jevgnijs lihto, www.flickr.com: CC-BY)
revision7, 11 . to the cultivated European plum (Prunus domestica)5, 14 . Unripe
fruits are used in sour soups, ripe fruits are eaten fresh or used by Agrobacterium tumefaciens . For this reason, some varieties
17
Habitat and Ecology to make non-alcoholic or fermented and distilled alcoholic are used as rootstock in grafting other fruit species and cultivars
This species has a broad area inside the temperate zone. beverages3 . Anthocyanin composition, phenolic content and of genus Prunus, such as plums, apricots and peaches3, 5, 6 . It
Natural populations are characterised by high variability in respect antioxidant activities of wild red and purple varieties of the is commonly cultivated as an ornamental tree for its colourful
of vigour, temperature tolerance, ripening time and disease cherry plum were tested and show potential for being developed leaves and numerous and delicate flowers. There are several
resistance, which makes it easily adaptable to a variety of sites12 . into a source of healthy fruit drinks due to their high antioxidant hybrids or varieties mainly with purple foliage: e.g. 'Pissardii with
The cherry plum is very hardy, with modest demands, except for activity. The fruit peel could be used as a resource to extract pink flower buds, white flowers and purple leaves, 'Nigra smaller
light. It can be found in open areas, such as forest margins, open natural pigments15 . This species is of great genetic importance plants with pink flowers and deep red-purple leaves18 .
woodlands, riversides and disturbed sites1-3, 6 . This tough plant is for horticultural breeding7. Genetic studies identified cherry plum
Threats and Diseases
also frost and drought tolerant, and wind resistant. It thrives on genotypes that are highly resistant to all root-knot nematodes
Together with other Prunus species, Cherry plum is an
a wide range of soil types, including those that are gravelly and of the genus Meloidogyne16 . Furthermore, these genotypes are
overwintering host species for the damson hop aphid Phoradon
sandy or poor in nutrients, but not compacted soils7, 13 . resistant to the expression of root crown gall consecutive infection
humuli, an important pest of hops Humulus lupulus19 . This plum
tree is also a natural host of plum pox virus, the causal agent
of Sharka disease, a serious economic threat for the production
of temperate stone fruits, such as apricots, plums and peaches.
Planted as an ornamental tree, it becomes a potential reservoir
of this virus20, 21 .
References
[1] D. A. Webb, Flora Europea. Volume 2. [11] A. Horvath, H. Christmann, F. Laigret,
Rosaceae to Umbelliferae, T. G. Tutin, Botany 86, 1311 (2008).
et al., eds. (Cambridge University Press, [12] D. Nikolic, V. Rakonjac, Genetika 39, 333
1968), pp. 7780. (2007).
[2] O. Polunin, Flowers of Europe: A Field [13] P. Duchovskis, V. Stanys, A. Sasnauskas,
Guide (Oxford University Press, 1969). C. Bobinas, Acta Horticulturae 734, 299
[3] T. Svulescu, ed., Flora Republicii Populare (2007).
Romine, vol 4 (Editura Academiei [14] B.-E. van Wyk, Food Plants of the World:
Romne, Bucureti, 1956). An Illustrated Guide (Timber Press, 2005).
[4] O. Johnson, D. More, Collins tree guide [15] Y. Wang, X. Chen, Y. Zhang, X. Chen,
(Collins, 2006). Journal of Food Science 77, C388 (2012).
[5] A. Horvath, H. Christmann, F. Laigret, [16] A. C. Lecouls, et al., Theoretical and
Botany 86, 1311 (2008). Applied Genetics 99, 328 (1999).
[6] B. G. Sutherland, R. Cerovi, T. P. Robbins, [17] M.-J. Rubio-Cabetas, J.-C. Minot, R. Voisin,
K. R. Tobutt, Euphytica 166, 385 (2009). D. Esmenjaud, European Journal of Plant
[7] P. Hanelt, ed., Mansfelds Encyclopedia Pathology 107, 433 (2001).
of Agricultural and Horticultural Crops [18] G. Szekely, D. Vioiu, Journal of
(Springer, 2001). Horticulture, Forestry and Biotechnology
[8] C. J. Webb, W. R. Sykes, P. J. Garnock- 15, 169 (2011).
Jones, Flora of New Zealand Vol. 4. [19] S. P. Worner, G. M. Tatchell, I. P. Woiwod,
Naturalised Pteridophytes, gymnosperms, Journal of Applied Ecology 32, 17 (1995).
dicotyledons (D.S.I.R., Christchurch, 1988).
[20] Elibuyuk, Phytoparasitica 34, 347 (2006).
[9] J. M. B. Smith, Telopea 3, 145 (1988).
[21] J. Sochor, P. Babula, V. Adam, B. Krska, R.
[10] D. R. Given, New Zealand Journal of Kizek, Viruses 4, 2853 (2012).
Botany 20, 221 (1982).
Europe and the Middle East. More recently this plant has been 25% - 50%
50% - 75%
used in horticulture as a frost-resistant rootstock for cherry > 75%
Chorology
plants. The mahaleb cherry, or St. Lucies cherry, (Prunus mahaleb Native
L.) is a deciduous shrub or small tree, reaching 10m tall. The
bark is dark brown, smooth and glossy1-4 . The young twigs are Ovate simple leaves with pointed tips and finely toothed margins.
glandular with yellowish-grey hairs, becoming later brownish and (Copyright Andrey Zharkikh, www.flickr.com: CC-BY)
Distribution
The bird cherry is the most widely distributed species of the
genus Prunus and can be found throughout northern Europe,
central Asia and as far east as Siberia, Northern China and
Japan1 . Its northern limit coincides broadly with the shores of the
Arctic Ocean, and in the south it has been recorded in Morocco5 . Map 1: Plot distribution and simplified chorology map for Prunus padus.
Frequency of Prunus padus occurrences within the field observations as
The white flowers arranged in pendulous racemes are insect pollinated.
reported by the National Forest Inventories. The chorology of the native
Habitat and Ecology spatial range for P. padus is derived after several sources16-22 .
(Copyright AnRo0002, commons.wikimedia.org: CC0)
in its natural and optimal habitat in North western America can > 75%
Uncertain, no-data
Marginal/no presence < 5%
Low presence 5% - 10%
Mid-low presence 10% - 30%
Medium presence 30% - 50%
Mid-high presence 50% - 70%
High presence 70% - 90%
Very-high presence > 90%
Coastal Dougal fir (Pseudotsuga menziesii var. menziesii) forest in Mount Tamalpais State Park (Marin County, California).
(Copyright Miguel Vieira, www.flickr.com: CC-BY)
robur), the wood is inferior, and only useful for rough work such
as shuttering or fuelwood1 . The leaves are dark green above and
grey-felted underneath6 ; they are variable in size and shape but
are normally 9-12cm long and 3-5cm wide, with 7-9 pairs of
triangular lobes6 . The leaves turn yellow to gold in late autumn
and drop off or persist in the crown until the next spring, especially
on young trees3 . The twigs are long and pubescent, grey or
olive-green, with lenticels. The buds, which are concentrated Large shade tree in agricultural area near Altamura (Bari, South Italy).
(Copyright Vito Buono, www.actaplantarum.org: AP)
on the tip of the twigs, are egg-shaped and hairy and, typically,
they are surrounded by long twisted whiskers6 . The flowers are can grow in a wide range of soil types including weakly acid14 ,
monoecious and wind-pollinated, appearing in April-May. The pseudogley12 , or even shallow calcareous soils, as long as they
fruit is a large acorn stalkless, 2-3.5(5)cm long and 2cm broad. are not too dry1 . When established it devlops a taproot and deep
The acorn cup is densely covered with bristles5 . Turkey oak acorns Map 1: Plot distribution and simplified chorology map for Quercus cerris. lateral root branches, helping it to remain windfirm3 . It is light-
Frequency of Quercus cerris occurrences within the field observations as reported
mature over a two year period, but the acorn crop is abundant by the National Forest Inventories. The chorology of the native spatial range for
demanding but can grow under a light woodland canopy1 . It has
and it germinates readily and can be easily propagated1, 3, 7. Q. cerris is derived after Meusel andJager; and Jalas andSuominen25, 26 . many pioneer characteristics, including good germination rates of
seeds and fast early growth. It also has a high resprouting capacity,
Distribution Apennines and covers around 280000 ha over the peninsula, making it particularly suitable for coppicing and pollarding3 .
The range of this species extends from southern Europe frequently occurring together with Hungarian oak (Q. frainetto)9 .
to Asia Minor3 . Across its distribution range, it is particularly It is also widely distributed in Slovenia, most frequently in the
present in the Balkan and Italian Peninsulas3 . The western sub-mediterranean regions of Kras, Brkini and Tolminsko, but it
limit of its natural range is France and its northern limit is in also grows on warm and dry steep slopes in the continental parts
Germany, continuing eastward through Austria, Switzerland, of the country9 . In the case of a warming climate, the species
eastern Czech Republic, Slovakia and Hungary3 . It is one of 12 is expected to show a range shift North11 . Turkey oak has been
native oak species in Albania. In Bulgaria it occupies drier and introduced in some other European countries including the UK
moderately rich habitats in the plain and hilly regions8 , where it and France3 , and it is also planted in North America4 , Ukraine,
forms large forests with other oak species (e.g. Quercus frainetto, Argentina and New Zealand3 .
Quercus pubescens) and other mixed broadleaves including field
maple (Acer campestre), elm (Ulmus minor) Oriental hornbeam Habitat and Ecology
(Carpinus orientalis) and manna ash (Fraxinus ornus)9 . It is also Turkey oak has a good adaptability to a variety of different
important in Hungary, where it forms over 11% of the forested site conditions. It is relatively tolerant to drought (more than the
area in the country10 . In Italy, it grows from sea level up to the other oak species of the same region)3, 12 , air pollution9, 13 and
Uncertain, no-data
Marginal/no presence < 5%
Forest dominated by Turkey oak in the Koutnjak Forest Park
Low presence 5% - 10% near Belgrade (Slovenia).
(Copyright Stefanst, commons.wikimedia.org: PD)
Mid-low presence 10% - 30%
Medium presence 30% - 50%
Mid-high presence 50% - 70%
Importance and Usage
The wood of Turkey oak has relatively few uses due to
High presence 70% - 90% its tendency to crack and its lower technological quality1 . It is
Very-high presence > 90% frequently used as firewood, having almost the same calorific
value as hornbeam or beech12, 15 . In past years the wood was
used for railway sleepers9, and it is still used for timber production
Annual precipitation (mm)
Map 2: High resolution distribution map estimating the relative probability of presence. Dark-green leaves with 7-9 pairs of lobes.
(Copyright Enrico Romani, www.actaplantarum.org: AP)
in the eastern part of its range, where the wood quality is at its
best3 . It has a useful role in soil conservation, erosion control
and reforestation of bare soils because of its ability to establish
and grow quickly in a range of soil types3 . Turkey oak is also Uncertain, no-data
often planted in urban areas as an ornamental tree as it is an Tundra, cold desert
attractive and well-formed tree1, 3 . The acorns and young coppice
Negligible survivability
shoots represent an important source of food for animals in
Mediterranean agro-silvopastoral systems3 . It is used in traditional Low survivability
Mediterranean medicine for numerous purposes, including anti- Mid-low survivability
infective treatments, and there is some evidence that it could be
Medium survivability
used against the pathogen Staphylococcus aureus16 .
Mid-high survivability
High survivability
Red female flower with fleshy stigmas blossoming with new leaves in spring.
(Copyright Graziano Propetto, www.actaplantarum.org: AP)
Grey bark with long fissures showing pinkish-orange colours in the cracks.
(Copyright Stefano Zerauschek, www.flickr.com: AP)
References
[1] P.S. Savill, The silviculture of trees used in [15] F.Clinovschi, Dendrologie (Editura
British forestry (CABI, 2013). Universitatii Suceava, 2005).
[2] A.DiIorio, B.Lasserre, G.S. Scippa, [16] G.H. Hobby, etal., Journal of
D.Chiatante, Tree Physiology 27, 407 (2007). Ethnopharmacology 144, 812 (2012).
[3] A. Praciak, et al., The CABI encyclopedia of [17] E.Amorini, M.Biocca, M.C. Manetti,
forest trees (CABI, Oxfordshire, UK, 2013). E.Motta, Annals of Forest Science 53,
[4] E.F. Gilman, D.G. Watson, Fact sheet ST- 731 (1996).
544: Quercus cerris - turkey oak (1994). [18] D.Jurc, N.Ogris, Plant Pathology 55, 299
[5] O.Johnson, D.More, Collins tree guide (2006).
(Collins, 2006). [19] S.Moricca, A.Ragazzi, Phytopathology 98,
[6] A.F. Mitchell, P.Dahlstrom, E.Sunesen, 380 (2008).
C.Darter, A field guide to the trees of [20] A.Ragazzi, etal., Phytopathologia
Britain and northern Europe (Collins, 1974). Mediterranea 40, 165 (2001).
[7] E.W. Jones, Journal of Ecology 47, 169 [21] A.Vannini, R.Valentini, Tree Physiology 14,
(1959). 129 (1994).
[8] S.Yurukov, P.Zhelev, Schweizerische [22] K.Schnrogge, etal., Galling Arthropods
Leaf gall caused by the wasp (Andricus quercuscalicis) on Zeitschrift fur Forstwesen 152, 52 (2001). and Their Associates, K.Ozaki, J.Yukawa,
pedunculate oak (Quercus robur): Turkey oak is the alternate [9] M.Bozzano, J.Turok, Mediterranean Oaks T.Ohgushi, P.Price, eds. (Springer Japan,
Stalkless acorns with cup covered by bristles. 2006), pp. 91101.
host completing the life cycle of this wasp. Network : report of the second meeting,
(Copyright Graziano Propetto, www.actaplantarum.org: AP) 2-4 May 2002 - Gozo, Malta (IPGRI, Rome, [23] D.deRigo, etal., Scientific Topics Focus 2,
(Copyright Somepics, commons.wikimedia.org: CC0)
Italy, 2003). mri10a15+ (2016).
Autoecology diagrams based on harmonised [10] R.Solymos, Annales des Sciences [24] K.V. Tubby, J.F. Webber, Forestry 83, 451
Field data in Europe (including absences) Observed presences in Europe field observations from forest plots. Forestires 50, 607 (1993). (2010).
[11] T.Hlsny, J.Holua, P.tpnek, [25] H.Meusel, E.Jager, eds., Vergleichende
M.Turni, N.Polk, Journal of Forest Chorologie der Zentraleuropischen Flora
Science 57, 422 (2011). - Band I, II, III (Gustav Fischer Verlag,
Jena, 1998).
Average temperature of the coldest month (C)
distinctive: up to 25cm long, widest close to the apex, with many > 75%
Chorology
deep-cut lobes (more than any other oaks2). The base of the leaf Native
Leaves are the most deeply lobed of the oak species: here just
starting to show autumn colours.
(Copyright Franco Caldararo, www.actaplantarum.org: AP)
the Balkan Peninsula. Despite its name, this oak is not native
to Hungary, although it is present sporadically as an introduced
Map 1: Plot distribution and simplified chorology map for Quercus frainetto.
Frequency of Quercus frainetto occurrences within the field observations
species8 . It is also present in north-west Turkey and southern Italy10
as reported by the National Forest Inventories. The chorology of the native in form of scattered patches along the pre-Apennine ridges11, 12 . As
spatial range for Q. frainetto is derived after Meusel and Jger10 .
a response to future expected warming its future distribution is
predicted to expand in Spain, France and Northern Italy13 .
(individual flowers are either male or female, but both sexes can
be found on the same plant) and are wind-pollinated. The acorns Habitat and Ecology
are up to 25mm long and egg shaped. In common with other oak Hungarian oak is a meso-xerophilous species, meaning
species it has high fructification rates that occur around every that it occupies a climate that is a transition between the typical
5-8 years3, 4 . The acorn cup is sessile and covered with long Mediterranean climate and a continental climate with hot
overlapping scales and hairs5, 6 . summers and harsh winters14 . It is light demanding and cannot
tolerate shading14 . It can grow in heavy acidic soils and tolerates
Distribution some water-logging15 . This species can form pure stands or more
Paleoecological evidence suggests that Hungarian oak was frequently it occurs mixed with hop hornbeam (Ostrya carpinifolia),
already present in the Peloponnese more than 6000 years ago7. It oriental hornbeam (Carpinus orientalis), South European flowering
Dark grey bark formed by small plates.
(Copyright Franco Caldararo, www.actaplantarum.org: AP) is indeed considered native to south-eastern Europe8 as an element ash (Fraxinus ornus) and Turkey oak (Quercus cerris)8 . This tree has
of the sub-Mediterranean flora9, with its widest distribution in a narrower ecological amplitude than that of Turkey oak in most
respects16 . It is more drought-tolerant than the Turkey oak but less
so than some other more Mediterranean oak species, such as holm
oak (Quercus ilex)17.
Uncertain, no-data
Marginal/no presence < 5%
Low presence 5% - 10%
Mid-low presence 10% - 30%
Medium presence 30% - 50%
Mid-high presence 50% - 70%
High presence 70% - 90%
Very-high presence > 90%
Mature acorn in scaled cup, which is hairy and sessile (stalkless) on twigs.
(Copyright Franco Caldararo, www.actaplantarum.org: AP)
Barrels made from Hungarian oak wood for conserving and flavouring wine.
(Copyright Elin, www.flickr.com: CC-BY)
References
[1] K. G. Kostadinov, Gorskostopanska Nauka [13] A. Baselga, M. B. Arajo, Ecography 32,
pp. 4155 (1984). 55 (2009).
[2] A. F. Mitchell, A field guide to the trees [14] G. Chatziphilippidis, G. Spyroglou,
of Britain and northern Europe (Collins, Sustainable Forest Management, H.
1974). Hasenauer, ed. (Springer Berlin Heidelberg,
[3] V. L. Sork, Plant Ecology 107-108, 133 2006), pp. 373395.
(1993). [15] T. G. M. Sanders, R. Pitman, M. S. J.
[4] I. Jerkovi, Z. Marijanovi, Molecules 15, Broadmeadow, iForest - Biogeosciences
3744 (2010). and Forestry 7, 61 (2014).
[5] F. Clinovschi, Dendrologie (Editura [16] R. Popovi, M. Koji, B. Karadi, Bocconea
Universitatii Suceava, 2005). 5, 431 (1997).
[6] O. Johnson, D. More, Collins tree guide [17] M. N. Fotelli, K. M. Radoglou, Constantinidou,
(Collins, 2006). Tree Physiology 20, 1065 (2000).
[7] S. Jahns 2, 187 (1993). [18] K. Kitikidou, E. Milios, E. Tsirekis, E. Pipinis,
Forest dominated by Hungarian oak in protected area forest near Foloi in Polyphagous caterpillar of the gypsy moth (Lymantria dispar). A. Stampoulidis, iForest - Biogeosciences
western Peloponnese peninsula (Elis, South Greece). (Copyright echoe69, www.flickr.com: CC-BY) [8] D. Bartha, Enzyklopdie der Holzgewchse: and Forestry 8, 53 (2015).
Handbuch und Atlas der Dendrologie,
(Copyright Huskarl, commons.wikimedia.org: PD)
A. Roloff, H. Weisgerber, U. M. Lang, B. [19] A. Vannini, R. Valentini, N. Luisi, Annales
Stimm, P. Schtt, eds. (Wiley-Vch Verlag, des Sciences Forestires 53, 753 (1996).
Autoecology diagrams based on harmonised Weinheim, 1998). [20] D. de Rigo, et al., Scientific Topics Focus 2,
Field data in Europe (including absences) Observed presences in Europe field observations from forest plots. [9] L. Corcuera, J. J. Camarero, E. Gil-Pelegrn, mri10a15+ (2016).
Trees 16, 465 (2002). [21] C. M. Brasier, Annales des Sciences
[10] H. Meusel, E. J. Jger, Plant Systematics Forestires 53, 347 (1996).
and Evolution 162, 315 (1989). [22] A. Sall, L. M. Nageleisen, F. Lieutier, Forest
Average temperature of the coldest month (C)
and shallowly cracked into small, square, thin plates5 . The twigs Chorology
Native
and the buds are grey-tomentose1 . Very variable in the shape,
the leaf is generally lanceolate to oval, 3-7cm long, thick but
not rigid, cuneate or rounded at the base, with 1-2cm woolly
petioles. The margins are waved or sinuate, but they can be
dentate or in some case spinose on young trees or sprouts1, 5 .
They unfold in spring silvery-white then pale yellow, covered with
dense hairs. Soon leaves become rough and shiny blackish-green
on the upper side, grey and densely pubescent on the lower5 . Leaf
lifespan ranges from less than 1 year to 4 years, with turnover
rates changing according to leaf position and environmental
factors6 . This is species is monoecious, blossoming in May-
June with new leaf growth6 . After dry summers new leaves in
autumn can appear2 . The male flowers are in dense pendulous
catkins 4-7cm long, pale green, then opening in a mass of yellow Map 1: Plot distribution and simplified chorology map for Quercus ilex.
Frequency of Quercus ilex occurrences within the field observations as
stamens very visible the against silvery-grey leaves. The female reported by the National Forest Inventories. The chorology of the native
flowers are minute, 2-3 on short and erect peduncles at the axil spatial range for Q. ilex is derived after Meusel andJager7.
on one leaf, green-grey and pubescent 2, 5 . The fruit is an acorn
ripening in the first year, brown in colour, 1.5-2cm long, one third
to half enclosed in a light green cupule with appressed scales, and Distribution
hanged on short peduncles2, 5 . Mature acorns fall in November- The natural distribution of holm oak occurs principally in the
January with high productions every 4-6 years6 . central-western part of the Mediterranean basin, covering from
Portugal and Morocco, to the Aegean Islands and western Turkey, Male catkins with yellow stamens producing pollen.
expanding also northward up to northern Italy and France. It also (Copyright Franco Giordana, www.actaplantarum.org: AP)
Map 2: High resolution distribution map estimating the relative probability of presence. Acorns half covered by the light green cupule.
(Copyright Giancarlo Pasquali, www.actaplantarum.org: AP)
References
[1] F. Clinovschi, Dendrologie (Editura [7] L. S. Minckler, Silvical Characteristics of Pin
Universitatii Suceava, 2005). Oak, USDA Forest Service (1957).
[2] A. F. Mitchell, A field guide to the trees [8] J. H. Carey, Quercus palustris. Fire Effects
of Britain and northern Europe (Collins, Information System (1992). http://www.
1974). feis-crs.org/feis
[3] E. F. Gilman, D. G. Watson, Fact sheet ST- [9] D. M. Rizzo, M. Garbelotto, J. M. Davidson,
555: Quercus palustris - pin oak (1994). G. W. Slaughter, S. T. Koike, Phytophthora
[4] R. A. McQuilkin, Quercus palustris ramorum and sudden oak death in
Muenchh. Pin Oak, Agriculture Handbook california: I. host relationships, Tech. rep.
654 (U.S. Department of Agriculture, (2002).
Forest Service, Washington, DC., 1990), pp. [10] C. Brasier, J. Rose, S. Kirk, S. Denman,
13661377. J. Webber, Comparative host range and
[5] O. Johnson, D. More, Collins tree guide aggressiveness of phytophthora ramorum
(Collins, 2006). and phytophthora kernoviae sp. nov. on
north american and european trees, Tech.
[6] T. G. M. Sanders, R. Pitman, M. S. J. rep. (2005).
Broadmeadow, iForest - Biogeosciences
and Forestry 7, 61 (2014).
Low valley crops in the mountain region near Graz (Styria, Austria).
(Copyright Christinamari, www.flickr.com: CC-BY)
Map 2: High resolution distribution map estimating the relative probability of presence. Leaves can persist on the tree during winter.
(Copyright Stefano Zerauschek, www.flickr.com: AP)
[43] S.Rivas-Martnez, etal., Itinera Geobotanica Deciduous Forests around the Northern
15, 5 (2002). Hemisphere, E.O. Box, K.Fujiwara, eds.,
Geobotany Studies (Springer International
Annual precipitation (mm)
Deeply lobed leaf with velvety stallate hairs on both sides giving an
ashy-green colour.
(Copyright Juan Ignacio Garca Vias: CC-BY)
Quercus robur and Quercus petraea in Europe: distribution, habitat, usage and threats
E. Eaton, G. Caudullo, S. Oliveira, D. de Rigo these oaks can mix, compete and naturally hybridise with other
Mediterranean oaks, such as Quercus pubescens and Quercus
Quercus robur L., (pedunculate oak) and Quercus petraea (Matt.) Liebl., (sessile oak) are common broadleaved tree frainetto, even if at relatively low rates12 . Both oaks occur at
species in Europe, found from Scandinavia to the Iberian Peninsula. The two species are quite similar in appearance and higher elevations in southern regions: Q. robur is recorded to grow
have a broadly overlapping range. Oak trees have cultural significance for people throughout Europe and the trees or up to 1300m in the Alps13, while Q. petraea is more montane
leaves are frequently used in national or regional symbols. Oak trees can live for more than 1000 years and grow to and in southern Turkey can reach over 2000m4, 14, 15 . Due to the
be 30 to 40m in height. The wood from oaks is hard and durable and has been valued for centuries. It is favoured for substantial human interest and usage of the species over many
construction and for wine and spirit barrels; historically it was a major source of ship timbers. Recently, concerns have centuries, there is widespread disturbance in their distribution, and
arisen about the fate of oaks in the face of Acute Oak Decline, a little understood syndrome. the structure of their original forests is highly uncertain16 . Q. robur
Quercus robur L., known as pedunculate or English oak, and has been introduced into the United States for timber production
Quercus petraea (Matt.) Liebl., known as sessile oak, are large, and in some areas it has naturalised; more recently they have been
rugged, deciduous broadleaved trees, native to most of Europe. Frequency
< 25%
exported into other continents as ornamental trees17, 18 .
Individuals can be very long-lived (over 1000 years in some cases) 25% - 50%
Uncertain, no-data
Marginal/no presence < 5%
Low presence 5% - 10%
Mid-low presence 10% - 30%
Medium presence 30% - 50%
Mid-high presence 50% - 70%
High presence 70% - 90%
Very-high presence > 90%
Map 2-A: High resolution distribution map estimating the relative probability of presence for Quercus robur. Acorn of pedunculate oak (Quercus robur) with long stalk.
(Copyright Graham Calow, www.naturespot.ork.uk: AP)
Map 3: High resolution map estimating the maximum habitat suitability for Quercus robur.
are frequently managed either as high forest or as coppice with mechanical stresses)10, 31, 32 . Furthermore promoting an abundant
standards23-27. Pedunculate and sessile oaks are amongst the and uniform natural regeneration is also crucial10, 15, 20, 33, 34 .
most economically important deciduous forest trees in Europe, In coppices, oaks provide a valuable source of firewood and
providing high quality hardwood for construction and furniture charcoal, and in the past the bark has been much used in the
manufacture2 . Q. petraea wood is largely indistinguishable from tanning of leather10 . Several cultivars have been selected for
that of Q. robur and is particularly appreciated for its straight ornamental purposes, especially from Q. robur, and exported
grain, its durability thanks to its hardness, and its high tannin all over the world. Oaks are particularly appreciated as park or
content, which makes it resistant to insect and fungal attacks28, 29 . roadside trees for their size and shade. Quercus robur Fastigiata
Oaks have traditionally been used in timber-framed buildings, is one of the most common cultivars, large in size and with a
as well as for fencing, gates and mining timber, and in the past columnar cypress-like habit; Quercus petraea Laciniata has
it was the most important wood used in the manufacture of long narrow and deeply incised leaves4 . These tree species also
wooden sailing vessels7. Furniture, floor-boards, panelling, joinery have an important ecological role, as they support many species
and veneer are also important uses of the wood10 . As the wood of insects such as moths, wood-boring beetles and gall-forming
is resistant to liquids, it has been used for barrels for wines and hymenoptera, and the acorns provide a valuable food source for
spirits, where the flavour imparted by the wood is often much many birds and mammals, such as jays, mice, squirrels and pigs10 .
desired13, 30 . The most valuable oak wood has narrow rings and is
produced in high mixed forests on fertile sites with long economic
rotations (about 160 years of age for Q. petraea, about 130
years for Q. robur)2 . Successful oak silviculture requires particular
attention, selecting the proper deciduous tree species mixture,
proportion and density, which strongly influence the wood quality
with regard to tree diameters, ring widths and the presence of
wood knots formed by lateral epicormic shoots. Choice of site
and management is also important to minimise the number of
trees with shake (the development of circular or radial cracks
through the timber, which substantially reduces its value and
Large pedunculate oak (Quercus robur) in Dunkeld Hilton park (Scotland). which is influenced by a number of factors including soil type and
(Forestry Commission, www.forestry.gov.uk: Crown Copyright)
300 year old sessile oak (Quercus petraea) in Rigney (eastern France).
(Copyright Arnaud 25, commons.wikimedia.org: PD)
reaching them7, 10 . In recent years, oak processionary moth squirrels10 . Pedunculate and sessile oaks are vulnerable to
(Thaumetopoea processionea) has spread from its native Lymantria dispar and moderately susceptible to Cryphonectria
habitat in southern Europe further north35 . This caterpillar parasitica37. They both suffer because of root pathogens of the
defoliates oaks and sheds micro hairs that are a serious irritant oomycete genus Phytophthora (P. cinnamomi, P. ramorum, P.
to the human respiratory system, eyes and skin. Knopper gall quercina)37. Phytophthora ramorum has been known to cause
wasps (Andricus quercuscalicis) cause some damage to acorn extensive damage and mortality in North America, known as
crops36 . Young oak trees often have their bark stripped by grey Sudden Oak Death. Although this pathogen has been detected
in Europe, it has not yet had a substantial effect on native
European oaks and it is under observation38 . Acute Oak Decline Bark of a sessile oak (Quercus petraea).
is a new syndrome affecting principally pedunculate and sessile (Copyright Stefano Zerauschek, www.flickr.com: AP)
Uncertain, no-data
Marginal/no presence < 5%
Low presence 5% - 10%
Mid-low presence 10% - 30%
Medium presence 30% - 50%
Mid-high presence 50% - 70%
High presence 70% - 90%
Very-high presence > 90%
Frequency
< 25%
25% - 50%
50% - 75%
> 75%
Chorology
Native
Annual precipitation (mm)
References
[1] MonumentalTrees.com, Monumental trees [23] P.S. Johnson, S.R. Shifley, R.Rogers, eds.,
(2015). The ecology and silviculture of oaks (CABI,
[2] A.Praciak, etal., The CABI encyclopedia of Wallingford, 2002).
forest trees (CABI, Oxfordshire, UK, 2013). [24] E.Hochbichler, Annales des Sciences
[3] I.ColinPrentice, H.Helmisaari, Forest Forestires 50, 583 (1993).
Pedunculate oak (Quercus robur) in winter near Havr village (Mons, Belgium). Ecology and Management 42, 79 (1991). [25] G.K. Kenk, Annales des Sciences
(Copyright Jean-Pol Grandmont, commons.wikimedia.org: CC-BY) [4] O.Johnson, D.More, Collins tree guide Forestires 50, 563 (1993).
(Collins, 2006). [26] R.Solymos, Annales des Sciences
[5] G.Aas, Annales des sciences forestires Forestires 50, 607 (1993).
50, 107s (1993). [27] G. Decocq, Biodiversity & Conservation 9,
[6] J.Jensen, A.Larsen, L.R. Nielsen, 1467 (2000).
J.Cottrell, Annals of Forest Science 66, [28] EN 350-2, Durability of wood and wood-
706 (2009). based products. Natural durability of
[7] E.W. Jones, Journal of Ecology 47 (1959). solid wood. Guide to natural durability
and treatability of selected wood species
[8] A.F. Mitchell, P.Dahlstrom, E.Sunesen, of importance in Europe (European
C.Darter, A field guide to the trees of Committee for Standardization, Brussels,
Britain and northern Europe (Collins, 1994).
1974).
[29] L.Meyer, etal., International
[9] I.Bossema, Behaviour 70, 1 (1979). Biodeterioration & Biodegradation 86,
[10] P.S. Savill, The silviculture of trees used in 79 (2014).
British forestry (CABI, 2013). [30] T.Garde-Cerdn, C.Ancn-Azpilicueta,
[11] A.Ducousso, S.Bordacs, EUFORGEN Trends in Food Science & Technology 17,
Technical Guidelines for genetic 438 (2006).
conservation and use for pedunculate [31] A.J.E. Price, Shake in oak: an evidence
and sessile oaks (Quercus robur/Quercus review (Forestry Commission, 231
petraea) (Bioversity International, 2003). Corstorphine Road, Edinburgh, 2015).
[12] A.L. Curtu, O.Gailing, R.Finkeldey, BMC [32] R.A. Mather, P.S. Savill, Forestry 67, 119
Evolutionary Biology 7, 218+ (2007). (1994).
[13] A.Roloff, H.Weisgerber, U.M. Lang, [33] E.Guilley, J.-C. Herv, G.Nepveu, Forest
B.Stimm, Bume Mitteleuropas: Von Aspe Ecology and Management 189, 111
bis Zirbelkiefer. Mit den Portrts aller (2004).
Bume des Jahres von 1989 bis 2010.
(Wiley-VCH, 2010). [34] G.Kerr, R.Harmer, Forestry 74, 467
(2001).
[14] H.Meusel, E.Jager, eds., Vergleichende
Chorologie der Zentraleuropischen Flora [35] E.Wagenhoff, R.Blum, K.Engel, H.Veit,
- Band I, II, III (Gustav Fischer Verlag, H.Delb, Journal of Pest Science 86, 193
Jena, 1998). (2013).
[15] G.Aas, Enzyklopdie der Holzgewchse: [36] M.J. Crawley, C.R. Long, Journal of
Handbuch und Atlas der Dendrologie, Ecology 83, 686 (1995).
A.Roloff, H.Weisgerber, U.M. Lang, [37] D.deRigo, etal., Scientific Topics Focus 2,
B.Stimm, P.Schtt, eds. (Wiley-Vch Verlag, mri10a15+ (2016).
Weinheim, 2000), vol.3. [38] N.J. Grnwald, M.Garbelotto, E.M. Goss,
[16] R. Bunce, Trees and Their Habitats: An K.Heungens, S.Prospero, Trends in
Ecological Guide to Some European Trees Microbiology 20, 131 (2012).
Grown at Westonbirt Arboretum (Forestry [39] S.Denman, N.Brown, S.Kirk, M.Jeger,
Commission Research and Development J.Webber, Forestry 87, 535 (2014).
Division, Edinburgh, 1982).
[40] J.N. Gibbs, B.J.W. Greig, Forestry 70,
[17] USDA NRCS, The PLANTS database (2015). 399 (1997).
National Plant Data Team, Greensboro,
USA, http://plants.usda.gov. [41] K.Sohar, S.Helama, A.Lnelaid, J.Raisio,
H.Tuomenvirta, Geochronometria 41
[18] K.C. Nixon, C.H. Muller, Flora of North (2014).
America North of Mexico, Flora of North
America Editorial Committee, ed. (New [42] C.deSampaioe Paiva Camilo-Alves,
York and Oxford, 1997), vol.3. M.I. daClara, N.M. deAlmeidaRibeiro,
European Journal of Forest Research 132,
[19] H.H. Ellenberg, Vegetation Ecology of 411 (2013).
Forest dominated by sessile oak (Quercus petraea) in Sierra Ancares (North Western Spain). Central Europe (Cambridge University
(Copyright Alfonso San Miguel: CC-BY) Press, 2009), fourth edn. [43] D. B. Stojanovi, T. Levani, B. Matovi,
S. Orlovi, European Journal of Forest
[20] G.Aas, Enzyklopdie der Holzgewchse: Research 134, 555 (2015).
Autoecology diagrams based on harmonised field Handbuch und Atlas der Dendrologie,
Field data in Europe (including absences) Observed presences in Europe A.Roloff, H.Weisgerber, U.M. Lang, [44] R.Siwkcki, K.Ufnalski, Forest Pathology
observations from forest plots for Quercus petraea. B.Stimm, P.Schtt, eds. (Wiley-Vch Verlag, 28, 99 (1998).
Weinheim, 2012), vol.3. [45] EUFORGEN, Distribution map of
[21] European Environment Agency, European pedunculate oak (Quercus robur) (2008).
forest types. categories and types for www.euforgen.org.
Average temperature of the coldest month (C)
Uncertain, no-data
Marginal/no presence < 5%
Low presence 5% - 10%
Mid-low presence 10% - 30%
Medium presence 30% - 50%
Mid-high presence 50% - 70%
High presence 70% - 90%
Very-high presence > 90%
The exterior of Apollos Spacecraft Lunar Module Adapter was
covered by a thin layer of cork painted white to minimize thermal
stresses during launch and ascent.
(Copyright NASA, commons.wikimedia.org: PD)
Annual precipitation (mm)
Map 2: High resolution distribution map estimating the relative probability of presence. Detail of the thick cork bark as extracted from the oak.
(Copyright Sallyofmayflower, commons.wikimedia.org: CC-BY)
Debarking operations for cork production in a plantation near Dehesa with oaks managed for cork production in Navalcn (Toledo, Central Spain).
Jimena de la Frontera (Andalucia, Spain). (Copyright Alfonso San Miguel: CC-BY)
(Copyright Alfonso San Miguel: CC-BY)
References
[1] L.Gil, M.C. Varela, EUFORGEN Technical [9] L.Gil, Materials 2, 776 (2009).
Guidelines for genetic conservation [10] A.Chapman, Englands Leonardo: Robert
and use for cork oak (Quercus suber) Hooke and the Seventeenth-Century
(Bioversity International, 2008). Scientific Revolution (CRC Press, 2004).
[2] A.Praciak, etal., The CABI encyclopedia of [11] Council of the European Union, Official
forest trees (CABI, Oxfordshire, UK, 2013). Journal of the European Union 35, 7
[3] J.Aronson, J.S. Pereira, J.G. Pausas, Cork (1992).
Oak Woodlands on the Edge: Ecology, [12] A. Costa, H. Pereira, M. Madeira, Annals of
Adaptive Management, and Restoration, Forest Science 67, 204 (2010).
The Science and Practice of Ecological
Restoration Series (Island Press, 2012). [13] V.Accio, M.Holmgren, F.Rego, F.Moreira,
Grazing bulls in a dehesa with cork oaks near Casas Alcobaza (Extremadura, West Spain). G.M.J. Mohren, Agroforestry Systems 76,
(Copyright Pablo, www.flickr.com: CC-BY)
[4] G.Allard, etal., State of Mediterranean 389 (2009).
forests 2013 (FAO, 2013). 177 pp.
[14] J.Herrera, Forest Ecology and
[5] D.Magri, etal., Molecular Ecology 16, Management 76, 197 (1995).
Autoecology diagrams based on harmonised 5259 (2007).
Field data in Europe (including absences) Observed presences in Europe field observations from forest plots. [15] D.deRigo, etal., Scientific Topics Focus 2,
[6] M.N. Bugalho, M.C. Caldeira, J.a.S. mri10a15+ (2016).
Pereira, J.Aronson, J.G. Pausas, Frontiers
in Ecology and the Environment 9, 278 [16] C.M. Brasier, Annales des Sciences
(2011). Forestires 53, 347 (1996).
Average temperature of the coldest month (C)
in some locations3 . It has a typical life span of about 60-100 25% - 50%
50% - 75%
Habitat and Ecology
years4 , although the current longevity record in Europe is for more > 75% Black locust is a light-demanding pioneer species and
than 300 years5 . The tree is usually bent-stemmed with greyish- grows either as an upright, single- or multi-stemmed tree or, in
brown to dark brown bark, becoming longitudinally fissured with harsh environments, as a multi-stemmed shrub. It tolerates a
age. The leaves are composed, pinnate, 10-30cm long, usually remarkably diverse range of soil conditions, being only limited by
with a pair of spines at the base which persist on young shoots. low soil aeration and waterlogging. On steep slopes, black locust
The leaflets are commonly in 2-12 pairs, usually opposite, with an is usually less vigorous19. The tree is easily damaged by extreme
additional one at the end of the rachis. Leaf blades are oblong, frosts in winter and presents low adaptability to arid conditions6, 20.
elliptic or ovate, 2-51.5-2.5cm, with entire margin. Black locust is Climate warming is expected to favour its further expansion21 . Its
a monoecious species: the hermaphrodite scented flowers have a reproduction is primary asexual through horizontal root elongation22 .
white to cream corolla with yellow spots inside, up to 2cm long and It is able to produces a high number of root suckers and therefore
grouped in pendent, many-flowered, axillary racemes 10-20cm exhibits the capacity for clonal growth, up to an area of 100 square
long6 . Black locust is a monoecious species: the hermaphrodite, metres23. Disturbance favours clonal growth and causes an increase
white and fragrant flowers are arranged in pendulous racemes 10- in the number of suckers19. The invasion of black locust is well
20cm long6 . The fruit is a legume, 5-10cm long dark brown pods documented in early succession habitats, like abandoned gravel-
hanging in winter and containing 4-10 seeds, mainly dispersed by sand pits and landfills, brownfields, secondary forests, coppiced
gravity and wind7. Fruiting takes place once or twice per year at a Map 1: Plot distribution and simplified chorology map for Robinia pseudoacacia.
Frequency of Robinia pseudoacacia occurrences within the field observations
forests, lowland pastures, roadsides, and burned sites6 . As with
tree age from 6 to 30-40 years8 . as reported by the National Forest Inventories. other species of the Fabaceae (syn. Leguminosae) family, the roots
of black locust host rhizobia bacteria in symbiotic relationship within
Distribution name to Jean Robin, who, with his son Vespasien, was among the structures called root nodules. These bacteria have the capacity to
Black locust is widespread across Europe, occurring from Sicily first arborists to introduce and cultivate this tree in Europe5. In the late take di-nitrogen gas (N2) out of the air and convert it to a form usable
in Italy to South Norway and longitudinally from the Portugal littoral 18th and early 19th century extensive planting of black locust trees to the host plant (nitrogen fixation). This process makes nitrogen
regions up to the Caucasus9-11 . Core areas in the alien range of this started in Central Europe. Currently it occurs in 42 European countries available to other plants, mainly through mineralization of leaf litter,
species are in sub-Mediterranean to warm continental climates, where and is naturalised in 3213, covering with pure and mixed stands, for and improves the quality of the soils. Black locust can add between
a rather high heat-sum is available7 and prolonged drought is rare12 . example, 400000ha in Hungary14, 200000ha in France15, 250000ha 23 and 300kg of nitrogen in a hectare every year6 . So stands of
It can be found from sea level up to 1640m in the Southern Alps in Romania16 and 230000ha in Italy17. Although accounted among black locust may generate substantial soil alterations, increasing the
(Prosser F., pers. comm., July 1, 2015). The species was introduced the 100 most invasive alien species in Europe11, only a few countries total soil nitrogen, the litter and the organic carbon, also changing
in Europe in the early 17th century. Linnaeus dedicated the genus have policies and initiatives to tackle it18. soil pH and decreasing the total phosphorus6 . Its adaptability and
the capacity of transforming ecosystem processes are the reason
for its adverse effects on biodiversity. However, inferior competitive
ability in later successional stages is reported where stands have
been left unmanaged for a sufficient time24 . Black locust commonly
Uncertain, no-data contributes to the composition of deciduous woodlands, mainly oak
Marginal/no presence < 5% and riparian woods, which are the most frequently invaded12 . Stands
dominated by black locust are cultural forest communities, which
Low presence 5% - 10%
can totally differ from native forest vegetation25. As such, they have
Mid-low presence 10% - 30% been either classified within the phytosociological separate class
Medium presence 30% - 50% Robinietea or included in other units, as a derivative community26 .
Several associations have been described in Central Europe, among
Mid-high presence 50% - 70%
which Balloto-Robinietum (diagnostic species include Ballota
High presence 70% - 90% nigra, Bromus sterilis, Dactylis glomerata) on poor aeolian sandy
Very-high presence > 90% soils, Solidagino-Robinietum (diagnostic species include Solidago
Map 2: High resolution distribution map estimating the relative probability of presence. Black locust in Berlin, Germany.
(Copyright Giovanni Trentanovi: CC-BY)
more1 . It is not long lived: usually surviving only 20-30 years2 . 25% - 50%
50% - 75%
The deeply fissured bark is dark grey with corky ridges. The leaves > 75%
Chorology
are long and narrow (lanceolate), silver-grey on the top side and Native
with dense silky white hairs on the underside, giving the tree a
distinctive pale appearance3, 4 . The yellow male catkins are up
to 5cm long; the female ones are shorter and greenish-yellow,
becoming fluffy white, and attract pollinating insects3, 3 .
Map 1: Plot distribution and simplified chorology map for Salix alba.
Frequency of Salix alba occurrences within the field observations as reported
by the National Forest Inventories. The chorology of the native spatial range
White willow in a riparian forest near Bingen (Rhineland-Palatinate,
for S. alba is derived after Meusel and Jger, and Skvortsov20, 21 .
West Germany).
(Copyright Willow, commons.wikimedia.org: CC-BY)
Habitat and Ecology areas in Europe with high erosion rates, in particular in moist
The white willow is a riparian species of temperate climates, slopes with high drainage-area within the European mountain
with mild winters, warm summers with short drought periods1 . It systems9 . In these critical areas, the white willow contributes to
typically grows near water on the banks of rivers and lakes or by mitigate erosion, and more generally it is also useful for erosion
ponds, streams, wet hollows and ditches5 . It occurs from sea level control and stabilising the banks of waterways, as well as for
to 2400m in the most southerly parts of its range1 . It tolerates a ecosystem restoration and phytoremediation10-12 .
wide variety of soil types, preferring sandy, silty or calcareous soils,
but the roots need access to water. It is strongly light-demanding
Threats and Diseases
and does not tolerate shade. It can be pollarded or coppiced and can This species is particularly susceptible to watermark disease,
readily reproduce from suckers or from adventitious roots6 . Adults caused by the bacterium Erwinia salicis13, 14 . Diseased trees suffer
of white willow are noticeably tolerant to inundation, performing from wilting, withering, and browning of leaves and tips of new
better even in comparison with flood-tolerant taxa such as Ulmus shoots, followed by die-back of affected branches and eventually
laevis, Ulmus minor and Quercus robur7. the death of the tree. Wood from affected trees becomes brittle
Silver-grey lanceolate leaves with hairy upper side. and unsuitable for use. The white willow is also attacked by the
(Copyright Rosendahl, commons.wikimedia.org: PD)
Importance and Usage Asian longhorned beetle (Anoplophora glabripennis), despite
The timber of white willow has a number of commercial uses. showing noticeable resistance and thus potentially acting
Distribution These include wooden kitchen utensils, archery bows, hoops, wicker as overwintering reservoir of the beetle15-17. It suffers from
The white willow is widely distributed throughout Europe baskets, canoes, and some construction use (e.g. log cabins)1 . The occasional but severe outbreaks of the gypsy moth (Lymantria
except in the northern regions, though Asia Minor and central variety Salix alba var. caerulea is known as cricket bat willow, dispar)15, 18, 19 . Furthermore, it is susceptible to Phytobia barnesi,
Russia to Chinese borders. In the west at its southernmost extent as its timber is particularly suited for this purpose8 . There are a which impairs the wood quality, and to the defoliators Porthetria
it can be found around the Mediterranean basin, including North number of non-wood products, including tannin from the bark and obfuscate and Yponomeuta rorella17.
Africa (Morocco and Algeria). In the north its range includes the salycin (from which aspirin was produced), but these generally
British Isles, the Netherlands and the eastern Baltic coast (Latvia have little commercial importance now2 . A number of cultivars References
and Lithuania). Its natural range is difficult to determine as it has with attractive stem colours have been bred for ornamental [1] J.G. Isebrands, J.Richardson, Poplars [14] J.G. Turner, J.M.L. Davis, K.Guven,
and willows: trees for society and the Proceedings of the Royal Society of
been extensively cultivated1, 2 . purposes. The white willow distribution range overlaps with many environment (CABI ; FAO, 2014). Edinburgh. Section B. Biological Sciences
[2] A. Praciak, et al., The CABI encyclopedia of 98, 105 (1992).
forest trees (CABI, Oxfordshire, UK, 2013). [15] D.deRigo, etal., Scientific Topics Focus 2,
[3] A.F. Mitchell, P.Dahlstrom, E.Sunesen, mri10a15+ (2016).
C.Darter, A field guide to the trees of [16] M.Faccoli, R.Favaro, M.T. Smith, J.Wu,
Britain and northern Europe (Collins, 1974). Agricultural and Forest Entomology 17,
[4] O.Johnson, D.More, Collins tree guide 188 (2015).
(Collins, 2006). [17] V. de Tillesse, L. Nef, J. Charles, A. Hopkin,
[5] R.V. Lansdown, The IUCN Red List S. Augustin, Damaging poplar Insects
of Threatened Species (2014), pp. - Internationally important species
203465/0+. (International Poplar Commission, FAO,
Rome, 2007).
[6] H.Koop, Vegetatio 72, 103 (1987).
[18] G.Camerini, Bulletin of Insectology 62,
[7] A.Schnitzler, The Botanical Review 63, 21 (2009).
40 (1997).
[19] M.J. Lechowicz, Y.Mauffette, Revue
[8] J.Wilkinson, Annals of Botany 5 (1941). dEntomologie du Quebec 31, 43 (1986).
[9] C.Bosco, D.deRigo, O.Dewitte, J.Poesen, [20] H.Meusel, E.Jger, eds., Vergleichende
P.Panagos, Natural Hazards and Earth Chorologie der Zentraleuropischen Flora
System Science 15, 225 (2015). - Band I, II, III (Gustav Fischer Verlag,
[10] P.S. Savill, The silviculture of trees used in Jena, 1998).
British forestry (CABI, 2013). [21] A.K. Skvortsov, Willows of Russia and
[11] Y.Kuzovkina, M.Quigley, Water, Air, and Adjacent Countries: Taxonomical and
Soil Pollution 162, 183 (2005). Geographical Revision (English translation
of 1968 Russian edition) (University of
[12] J.Ball, J.Carle, A.DelLungo, Unasylva 56, Joensuu, Finland, 1999).
3 (2005).
[13] W. R. Day, The watermark disease of the
cricket bat Willow (Salix caerulea), Oxford
Forestry Memoirs n. 3 (Clarendon Press,
Oxford, 1924).
Limestone cliffs of the Praia da Marinha near Lagoa (Algarve, South Portugal).
(Copyright Miguel Vieira, www.flickr.com: CC-BY)
well-developed fibrous root system1 . The trunks are small, 25% - 50%
50% - 75%
occasionally reaching a diameter of 40cm4 . The bark is smooth > 75%
Chorology
at the beginning, forming a thin rhytidome, irregularly cracked2 . Native
The twigs are greenish, thick and with grey hairs. The leaves are
alternate, broadly elliptic and up to 5-12cm long. The upper side
of the leaf is green and glossy, while the underside is covered
by densely and softy white-downy hairs1, 5 . The goat willow is a
dioecious species and catkins appear in early spring (usually in
March or April, depending on the site conditions). Male catkins have
spreading yellow stamens, while female catkins are greenish and
insect-pollinated. The catkins are erect, with approximately 100-
200 flowers in each female catkin and 200-300 flowers in each
male catkin, respectively. Catkins are produced abundantly and
they appear before the leaves3 . This willow naturally hybridises
with a number of other Salix species, producing fertile descendants
with intermediate characteristics. Hence, this makes it difficult
Map 1: Plot distribution and simplified chorology map for Salix caprea.
to determinate taxonomical limits and to assign individuals to a
Frequency of Salix caprea occurrences within the field observations as
particular species within the Salix genus6, 7. reported by the National Forest Inventories. The chorology of the native
spatial range for S. caprea is derived after several sources9, 52-56 .
Distribution
This species is native to cool temperate and boreal regions types17. Compared with other willow species, the goat willow is Goat Willow in Hesse, Germany.
of Europe and Asia, occurring in a large range of habitats across more sensitive to continuous flooding18 , avoiding the saturated (Copyright Willow, commons.wikimedia.org: CC-BY)
Europe and Asia8 . Its wide distribution spreads from Spain to soils of wetlands19 . Regarding its demand for light, it is a very
China, from Turkey and northern Iran to Fennoscandia and shade intolerant species, occurring in open areas with full sun1 , Importance and Usage
Siberia reaching up to 70north latitude1, 9-11 . It has been also but it is one of the few willow species able to grow in forest This species has a wide variety of uses. The scientific name
introduced in eastern North America and is now naturalised12 . understories8 . Unlike almost all other willows, the goat willow can caprea actually means goat, which probably derives from the
only occasionally be propagated vegetatively by cuttings8, 20 . fact that its leaves are used as fodder27. In northern Europe, it
Habitat and Ecology In boreal forests, it is found in the dominant Norway spruce
The goat willow is a pioneer13 and a fast-growing plant14 , is a common species in the agricultural landscape, used as a
(Picea abies), Scots pine (Pinus sylvestris) and birch (Betula spp.) windbreak and hedge28 , and its foliage is used as fodder for cattle
tolerating a wide variety of edaphic and climatic conditions1 . It forests, in admixture with other deciduous species such as aspen
shows a high adaptability to different habitat conditions15 , such and goats29, 30 . In traditional medicine, goat willow extracts are
(Populus tremula) and rowan (Sorbus aucuparia)21, 22 , favoured used as a painkiller, astringent, antiseptic, eye tonic28 , or even to
as dry or wet sites16 , usually growing in mesic to moist stands4 . by disturbances which create open areas23 . In temperate forests
It prefers the calcareous sites, but it grows on almost all soil treat malaria, gout, neuralgia and intestinal diseases31 . As is also
dominated by oaks (Quercus spp.), European beech (Fagus the case with other willows, a good charcoal can be obtained from
its burnt wood, which in the past was used for gunpowder and
for drawing pencils32, 33 . The goat willow is particularly valuable
for its high biomass production in short rotation plantations and
its role in landscape restoration34, as it is able to tolerate even
Uncertain, no-data
polluted land35. Worldwide, this willow is also used for its ability
Marginal/no presence < 5% to extract heavy metals, such as Cd and Zn, (i.e. phyto-extraction
Low presence 5% - 10% strategy) from polluted sites34-36 . During spring holidays, in
several places across Europe, its flowering branches are in high
Mid-low presence 10% - 30%
demand37. It is also appreciated as a melliferous plant species38,
Medium presence 30% - 50% 39
, supplying a honey production of 150-200 kg/ha40 . The goat
Mid-high presence 50% - 70% willow distribution range overlaps with many areas in Europe with
high erosion rates, including European boreal areas and moist
High presence 70% - 90%
slopes with high drainage-area within the European mountain
Very-high presence > 90% systems41 . In these critically erosion-prone areas, it contributes
to key ecosystem services such as watershed protection and soil
stabilisation42 . As for other willows and poplars, it is also useful
for ecosystem restoration and phytoremediation 43,44 .
Finally, the goat willow plays an important role in maintaining
species diversity, by being the host of several lichen species4, 37.
Annual precipitation (mm)
Map 2: High resolution distribution map estimating the relative probability of presence. Male catkins turn yellow when the pollen is ready for release.
(Copyright AnRo0002, commons.wikimedia.org: CC0)
are in opposite pairs with 3-(5-7)-9 leaflets, which are usually > 75%
Chorology
ovate, acuminate and with serrate margins1, 2 . The leaves are Native
Distribution
The elder is common in western and central Europe. Its
Map 1: Plot distribution and simplified chorology map for Sambucus nigra.
northern limit is Scotland and southern Scandinavia, and it can Frequency of Sambucus nigra occurrences within the field observations as Pedunculate inflorescence with white flowers arranged in umbel.
be found as far south as Sicily and mainland Greece. It can also reported by the National Forest Inventories. The chorology of the native (Copyright Tracy Houston Durrant: CC-BY)
be found in Asia and North Africa (probably through cultivation), spatial range for S. nigra is derived after Meusel andJger11 .
events between taxa and populations. Within the subgenus Aria, forests in Croatia6 .
the section Aria, i.e. the Sorbus aria sensu lato (s.l.), includes Importance and Usage
primary diploid species, its subspecies, their apomictic triploid Habitat and Ecology
and tetraploid forms and their hybrids3, 4 . This species complex Whitebeam is a mesophile species that can grow in different Whitebeam is quite a popular ornamental tree but its
occurs in a wide range across western, central and southern soil types. However, it occurs relatively rarely in very acid and very practical use is comparatively low. There are many garden
Europe, in North-western Africa in mountains of Morocco, Algeria alkaline soils or in habitats with light and sandy soils, as well as in forms and the wood was used for furniture and various items of
and Tunisia, eastwards to Turkey, Lebanon, Caucasus and the heavy clay soils. Usually it prefers moist but well-drained, neutral household equipment in history. The frozen or cooked fruits are
edible but not tasty. In former times the fruits, leaves and twigs
were used as livestock-fodder and the mealy fruit substance was
processed into a type of meal or flour, which was used to extend
regularbreadflour (German name Mehlbeere = Flour Berry)1 .
Planting material can be propagated by seed and inoculation.
Whitebeam is a slow-growing species that may have a shrub form. Bark is smooth in young plants.
(Copyright Stefano Zerauschek, www.flickr.com: AP) (Copyright Silvano Radivo, www.actaplantarum.org: AP)
Uncertain, no-data
Tundra, cold desert
Negligible survivability
Low survivability
Mid-low survivability
Medium survivability
Mid-high survivability
High survivability
Fruits are orange to scarlet and are eaten and dispersed mostly by birds.
(Copyright Stefano Zerauschek, www.flickr.com: AP)
leaves are odd pinnate, 10-25cm, consisting of 9-19 (usually Scarlet fruits of 1cm in diameter; these pomes stay
attached to the tree during the winter.
15) pairs of 2-6cm leaflets, which are oblong, sessile, with (Copyright Aldo De Bastiani, www.actaplantarum.org: AP)
acute apex, serrate margins, dark green in colour becoming
orange in autumn5, 6 . This species is monoecious with numerous hardy and frost tolerant1, 3 . It is absent from wetlands, bare soil
hermaphrodite white flowers arranged in dense and woolly sites, clays and soft limestones1, 3 . Otherwise, it is one of the least
inflorescences1, 2 , flowering from May to June3 , or as late as July site-demanding trees3 . Rowan and birch (Betula spp.) have quite
in northern Europe2 . It is insect pollinated3 and starts to produce similar site requirements for growth with respect to acidity (up to
seed at an age of 15 years1 . The fruits are scarlet round pomes pH 7.0) and non-waterlogged conditions, but rowan is more shade
of about 1cm6, 7, and its yield is usually good with alternating tolerant and can persist at higher elevations14, 15 . It is widespread as
better and poorer years1 . The fruits stay attached to the tree a minor component of several vegetation communities, occurring
during the winter and therefore offer food for birds7, which from the lowlands to rocky mountain slopes and cliffs. Typical sites
disperse the seeds1, 3 . The tree is usually not expected to reach for rowan are forest edges, glades, rocky and stony sites, riverbank
an age of 100 years7. bluffs and undergrowth with several forest types and tree species1,
Map 1: Plot distribution and simplified chorology map for Sorbus aucuparia.
Frequency of Sorbus aucuparia occurrences within the field observations
2
. Often it occurs as a pioneer tree in open habitants and also in
Distribution as reported by the National Forest Inventories. The chorology of the native forests, as it is able to geminate and establish successfully on thick
The genus Sorbus has a complex taxonomy, as their species spatial range for S. aucuparia is derived after Meusel and Jager11 .
layers of raw humus in dense forest stands of spruce (Picea abies)
can create not only hybrids, but also duplicate their genome particularly, and tolerates competition from herbs16 . Dominant
resulting in different morpho-ecological characters that are Tyrol1, 11, 12 . It has also been introduced in the United States and well developed trees regenerate principally by seeds, while pioneer
classified as different species. The diploid form of Sorbus Canada as an ornamental tree and has become naturalised in trees spread laterally, developing clones by roots1, 14-16 . It has been
aucuparia belongs to subgenus Sorbus; furthermore apomictic some northern regions13 . shown that the genetic variation has remained relatively high even
triploid and tetraploid species, as a result of hybridisation with in quite fragmented populations, and there are no clear indications
the members of subgenus Aria, are grouped in the subgenus Habitat and Ecology of inbreeding, which is a good sign for forest restoration17.
Soraria8-10 . This tree is widespread across Europe, from extreme Rowan is a tree species adapted for a short growing season
northern regions (Iceland, Fennoscandia, Russia) to southern and it can tolerate high summer temperatures, if the temperature is
Europe (Spain, Italy, Balkans), where it is found only at higher not accompanied by high water stress. In other words, its spread is
elevations, and eastwards the species extends into Asia Minor limited rather by a combination of poor drought tolerance, adaption
(Caucasus) and Northern Asia (Siberia, North China). It can to a short growing season and a cold requirement for bud burst
be found from sea-level in northern latitudes, up to 2400m in than by high temperature1 . It is a stress-tolerant competitor14 , cold
Uncertain, no-data
Marginal/no presence < 5%
Low presence 5% - 10%
Mid-low presence 10% - 30%
Medium presence 30% - 50%
Mid-high presence 50% - 70%
High presence 70% - 90%
Very-high presence > 90% Hermaphrodite white flowers arranged in corymbs
formed of numerous individuals.
(Forestry Commission, www.forestry.gov.uk: Crown Copyright)
References
[1] O.Rasp, C.Findlay, A.-L. Jacquemart, [17] C.F.E. Bacles, A.J. Lowe, R.A. Ennos,
Journal of Ecology 88, 910 (2000). Molecular Ecology 13, 573 (2004).
[2] V.L. Komarov, etal., Flora of the USSR [18] C.Bosco, D.deRigo, O.Dewitte, J.Poesen,
- Volume IX: Rosales and Sarraceniales, P.Panagos, Natural Hazards and Earth
vol.9 of Flora of the USSR (Keter Press, System Science 15, 225 (2015).
Jerusalem, 1970). [19] F.Florineth, H.P. Rauch, H.Staffler,
[3] P.S. Savill, The silviculture of trees used in Proceedings of the International Congress
British forestry (CABI, 2013). INTERPRAEVENT 2002 in the Pacific Rim
[4] O.Mauer, E.Paltov, Journal of Forest (2002), vol.2, pp. 827837.
Science 48, 342 (2002). [20] J.E. Norris, A.DiIorio, A.Stokes, B.C.
[5] A.F. Mitchell, P.Dahlstrom, E.Sunesen, Nicoll, A.Achim, Slope Stability and
C.Darter, A field guide to the trees of Erosion Control: Ecotechnological
Britain and northern Europe (Collins, Solutions, J.E. Norris, etal., eds. (Springer
1974). Netherlands, 2008), pp. 167210.
[6] O.Johnson, D.More, Collins tree guide [21] A. Gil-Izquierdo, A. Mellenthin, European
(Collins, 2006). Food Research and Technology 213, 12
(2001).
[7] A.Anderberg, A.L. Anderberg, Den
virtuella floran (Elektronisk publikation, [22] K.Haffner, S.F. Remberg, Chronica
Naturhistoriska riksmuseet, Stockholm, Horticulturae 46, 19 (2006).
1996). Http://linnaeus.nrm.se/flora. [23] J.E. Milner, The tree book : the
[8] E. Nelson-Jones, D. Briggs, A. Smith, indispensable guide to tree facts, crafts
Theoretical and Applied Genetics 105, and lore (Collins & Brown, 1992).
953 (2002). [24] S.D. Wratten, P.Goddard, P.J. Edwards,
[9] A.Robertson, etal., Molecular Ecology 19, The American Naturalist 118, 916 (1981).
1675 (2010). [25] G.Jaastad, etal., Agricultural and Forest
[10] J.J. Aldasoro, C.Aedo, C.Navarro, F.M. Entomology 4, 233 (2002).
Garmendia, Systematic Botany 23, 189 [26] S.Kobro, etal., Population Ecology 45,
(1998). 25 (2003).
[11] H.Meusel, E.Jager, S.Rauschert, [27] Z.Polak, J.Zieglerova, Zeitschrift fur
E.Weinert, Vergleichende Chorologie der Pflanzenkrankheiten und Pflanzenschutz
Zentraleuropischen Flora (Gustav Fischer 103, 432 (1996).
Verlag Jena, 1978). [28] N.Mielke, H.-P. Muehlbach, Journal of
[12] E.Hultn, M.Fries, Atlas of North European General Virology 88, 1337 (2007).
vascular plants (North of the Tropic of [29] J.Grace, F.Berninger, L.Nagy, Annals of
Cancer), Vols. I-III. (Koeltz scientific books, Botany 90, 537 (2002).
Rowan tree grown in a dwarf mountain pine thicket (Pinus mugo) in the Adamello-Brenta Natural Park (Trentio-Alto-Adige, North Italy). 1986).
[30] P.A. Harrison, P.M. Berry, N.Butt, M.New,
(Copyright Giallopolenta, commons.wikimedia.org: PD) [13] C.S. Sargent, Manual of the Trees of Environmental Science & Policy 9, 116
North America (exclusive of Mexico), vol.2 (2006).
(Dover Publications, New York, 1961),
Autoecology diagrams based on harmonised second edn. [31] D.deRigo, etal., Scientific Topics Focus 2,
Field data in Europe (including absences) Observed presences in Europe field observations from forest plots. [14] J. P. Grime, J. G. Hodgson, R. Hunt,
mri10a15+ (2016).
Comparative Plant Ecology: A Functional [32] CABI, Lymantria dispar (gypsy moth)
Approach to common British Species (2015). Invasive Species Compendium.
(Castlepoint Press, 2007). http://www.cabi.org
[33] CABI, Lymantria monacha (nun moth)
Average temperature of the coldest month (C)
Distribution
Service tree is distributed through central and southern
Europe in particular in the Balkan peninsula, Italy and southern
France, and rarely in parts of North Africa and western Asia1 . At its
northern limit there are a few scattered individuals in Wales and
central England, although it is thought that these are probably
descended from cultivated trees4 . Nowadays, it is unclear how
much of its distribution range is natural, since this species has Map 1: Plot distribution and simplified chorology map for Sorbus domestica.
been spread through cultivation since Roman times1 . However, Frequency of Sorbus domestica occurrences within the field observations
as reported by the National Forest Inventories. The chorology of the native
it is rare throughout its range5 and endangered in many parts3 . spatial range for S. domestica is derived after EUFORGEN12 .
moderately acid or neutral soils6 . It can also be found in coppice and can command a high price; it is often used for making small
and former coppice forests as a result of historic cultivation6 . It objects such as screws, gear teeth, bearings and rifle butts1, 3 .
is a very light demanding species that can tolerate shade only
in its first few years. This makes it a weak competitor, unable Threats and Diseases
to dominate when crown closure occurs6 . Under conditions of a Diseases of apple, such as European canker of apple (Nectria
warming climate it may become more favoured in some sites as galligen) and apple scab (Venturia inaequalis) can affect the
a result of its drought tolerance, but its scarcity is likely to limit service tree and cause premature leaf fall8 . Across Europe, the
any range movement8 . species is very rare and its genetic diversity is threatened mainly
due to the reduction in number of individuals and disturbance of
the natural populations by human activities6, 11 .
References
[1] P.Rotach, EUFORGEN Technical guidelines [6] V.Paganov, Journal of Forest Science 54,
for genetic conservation and use for 216 (2008).
Service tree Sorbus domestica (2003). [7] P.Gonin, etal., Autoecology of
[2] A.F. Mitchell, P.Dahlstrom, E.Sunesen, broadleaved species (2013).
C.Darter, A field guide to the trees of [8] G.E. Hemery, etal., Forestry 83, 65
Britain and northern Europe (Collins, (2010).
1974).
[9] O.Vyviurska, S.Pysarevska, N.Jnokov,
[3] C.Bignami, Conservation, evaluation, I.pnik, Open Chemistry 13 (2015).
exploitation and collection of minor fruit
tree species: EC Project GENRES 29, [10] A.Termentzi, P.Alexiou, V.J. Demopoulos,
E.Bellini, E.Giordani, eds. (Horticulture E.Kokkalou, Pharmazie 63, 693 (2008).
Department, University of Florence, 1999). [11] J.Mllerov, R.Hdl, P.Szab, Forest
[4] M.Hampton, Watsonia 20, 379 (1995). Ecology and Management 343, 88 (2015).
[5] B.Demesure, Noble Hardwoods Network: [12] EUFORGEN, Distribution map of service
Report of the Second Meeting, 22-25 tree (Sorbus domestica) (2014).
March 1997, Lourizan, Spain, J.Turok, www.euforgen.org.
etal., eds. (Rome, Italy, 1997), pp. 4850.
10x7cm and have five to nine spreading, acute lobes. This Native
is given as 300-400 years1-3 . Individuals start fruiting typically Due to efficient seed dispersal by birds and small mammals, and
at 15-20 years of age, abundantly and frequently, with seed the ability to spread vegetatively by root suckers, the species can
years every 2-3 years4 . The fruits are small pomes, 10-15mm in easily colonise appropriate sites, such as larger gaps, clearings,
diameter, spotted with green to russet or brown colour, and are low-density forests and abandoned agricultural land surrounded
dispersed by birds or frugivorous mammals1-3 . Seeds are light by forests, thus remaining permanently present in the landscape8 .
brown, of ovate shape with an approximate size of 2-6mm1, 5, 6 . Regarding soil characteristics it is a very tolerant species that can
grow on both acid and basic soils (from clay to limestone), with
Distribution
a pH ranging from 3.5 to 8 and humus types from dismoder to
The wild service tree is a sub-Mediterranean species and
carbonate mull2 . It avoids both dry sandy soils and wet or marshy
occurs across western, central and southern Europe, but also Map 1: Plot distribution and simplified chorology map for Sorbus torminalis. soils and reaches its best growth on warm-dry limestone soils1.2 .
in mountains of north-western Africa and south-western Asia. Frequency of Sorbus torminalis occurrences within the field observations
Current abundance centres seem to be located in France and in as reported by the National Forest Inventories. The chorology of the native
spatial range for S. torminalis is derived after EUFORGEN13 .
the Balkan region2 . Upper limits are located at 500m in Germany
and 700m in the Swiss Mittelland3 . On south-facing slopes of species is getting increasingly scarce in mountain areas towards
the Jura Mountains and in valleys in the Alp the species occurs up the north of the distribution range, and is rather uncommon in
to 900m. In the Atlas Mountains it has its centre of distribution cool habitats like north-facing slopes or cold valley bottoms7.
between 1300-1800m. The highest occurrence in elevation is The same lack of summer warmth limits the general vertical
reported from Anatolia (Lake Van) at 2200m1 . distribution. The optimum mean annual temperature is between
10and 17C; minimum annual rainfall is about 500mm,
Habitat and Ecology while the optimum lies between 700 and 1500mm3 . The wild
This species tolerates harsh winter conditions and is quite service tree is a shade intolerant, post-pioneer (nomadic) species,
insensitive to late frosts: it withstands temperatures down to -5C requiring free-growth for optimal crown development. It is a minor
in April7. In increasingly drier regions like in southern Europe, the component of woodlands-types dominated by various oaks and
wild service tree is a mountain forest species. A certain amount less frequently pine or beech, where it is found as single trees or
of warmth is necessary during the growing season; therefore the in small groups, and population densities are generally very low2 .
Flowers are arranged in corymbs of 20-30 individual blossoms. Scented white hermaphrodite flowers: they are insect pollinated.
(Copyright Emma Silviana Mauri: CC-BY) (Copyright Sergio Piccolo, www.actaplantarum.org: AP)
Apples affected by the disease Venturia inaequalis; this ungulates. Among invertebrate consumers, only generalist
Ascomycete fungus can affect also wild service tree.
herbivore arthropods are known. On the leaves of the wild
References
(Copyright Jan Homann, commons.wikimedia.org: PD)
[1] U.Pietzarka, M.Lehmann, A.Roloff, [6] L.Bednorz, Dendrobiology 57, 3 (2007).
service tree three damaging factors are frequently recorded: leaf Enzyklopdie der Holzgewchse: [7] P.Gonin, etal., Autecology of broadleaved
Handbuch und Atlas der Dendrologie,
scab (Venturia inaequalis), leaf aphids and gall mites (Eriophyes A.Roloff, H.Weisgerber, U.M. Lang,
species (Institut pour le Dveloppement
Threats and Diseases spp.)3 . The tree is a wild host for the apple aphid (Aphis pomi)10, 11 . B.Stimm, P.Schtt, eds. (Wiley-Vch Verlag,
Forestier, Paris, 2013).
Weinheim, 2008), vol.3. [8] V.Kuerov, M.Honec, L.Paule, P.Zhelev,
Older trees of the wild service tree can reach a root-depth A serious threat to older stems is the honey fungus (Armillaria [2] V.N. Nicolescu, E.Hochbichler,
D.Gmry, Biologia 65, 817 (2010).
J.CoelloGomez, S.Ravagni, V.Giulietti, [9] L.Kutschera, E.Lichtenegger, Wurzelatlas
of 1-2 m, rendering it quite resistant to wind-throw9 . Young spp.), whose parasitic infestation can lead to lethal damage12 . Die Bodenkultur 60, 35 (2009). mitteleuropischer Waldbume und
trees are especially susceptible to browsing by deer and other [3] H.Kutzelnigg, Gustav Hegi - Illustrierte
Strucher (L. Stocker Verlag, Graz, 2002).
Flora von Mitteleuropa, Band 4, Teil [10] CABI, Aphis pomi (apple aphid) (2015).
2B: Rosaceae, H.Scholz, ed. (Blackwell Invasive Species Compendium.
Wissenschafts-Verlag, Berlin, 1995), pp. http://www.cabi.org
328385. [11] T.Gotlinuljak, D.Grubii, A.Mei,
[4] E.Favredanne, LAlisier torminal I.Juran, Natura Croatica 21, 191 (2012).
(Sorbus torminalis Crantz.): synthse [12] G.Hartmann, Corminaria 17, 24 (2002).
bibliographique, recherche sur la densit
du bois (Ecole Nationale du Gnie Rural, [13] EUFORGEN, Distribution map of wild
des Eaux et des Forts (ENGREF), Nancy, service tree (Sorbus torminalis) (2014).
France, 1990). www.euforgen.org.
[5] M.Orani, D.Drvodeli, T.Jemri, I.Ani,
S.Mikac, Periodicum biologorum 111,
495 (2009).
The genus Tamarix occurs naturally from Western Europe and Threats and Diseases
the Mediterranean to North Africa, northeastern China, Mongolia, Three species of fungi, Botryosphaeria tamaricis, Diplodia
India and Japan1, 6 . In Europe, 14 species are reported. African tamarascina and Leptosphaeria tamaricis, can form cankers
tamarisk (Tamarix africana), Bovs tamarisk (Tamarix boveana), and cause branch dieback on tamarisks. The latania scale
Canary Islands tamarisk (Tamarix canariensis), Dalmatian (Hemiberlesia lataniae) and oystershell scales (Lepidosaphes
tamarisk (Tamarix dalmatica), French tamarisk (Tamarix gallica), ulmi) are two insects which frequently infest these species20 .
Hampes tamarisk (Tamarix hampeana), small-flower tamarisk Map 1: Plot distribution and simplified chorology map for Tamarix spp.
Some species, principally salt cedar (Tamarix ramosissima)
(Tamarix parviflora), Smyrna tamarisk (Tamarix smyrnensis) Chorology of the native spatial range for Tamarix spp., derived after Meusel
and Jger22 . Chinese tamarisk (Tamarix chinensis), small-flower tamarisk,
and four-stamen tamarisk (Tamarix tetranda) are native to the French tamarisk and their hybrids, are considered as invasive
Mediterranean area, mainly France, Spain, Portugal, Italy and waterways and on sandy floodplains, especially where their roots weeds in the United States, and have been the target of many
Greece and Turkey7, 8 . The species Tamarix ramosissima, Tamarix can access underground water9 . They grow best in alkaline soils, control programmes since the 1960s21 .
laxa, Tamarix hispida, Tamarix gracilis and Tamarix meyeri are but also tolerate acidity12 . These plants are found on non-rocky
native to Eastern Europe, Moldova, Russia and Ukraine7. Some silt loams and clay loams of high organic matter along streams,
species have become naturalised in other countries, especially bottomlands, pond margins, banks of drainages and washes
America, where they were introduced as landscape ornamentals and other wet environments in arid and semiarid regions13, 14 .
and have escaped cultivation. Tamarisks are not shade-tolerant14 .
are almost always hollow2 making tree ring-based age estimation > 75%
Chorology
impractical, opinion is divided about the exact age of the oldest Native
Male yellow globose flowers at the base of the leaves.
specimens3 . Yew is often multi-stemmed and its height rarely (Copyright Michael Wunderli, www.flickr.com: CC-BY)
exceeds 20 m, but its longevity means that trunks can be very
large: up to 4m diameter. Root systems are shallow with extensive Threats and Diseases
horizontal roots3 . A non-resinous conifer, the needles are very dark Yew has experienced one of the sharpest declines of all
green above and yellow to matt green below; they are 2-3cm long European tree species. Historically it was felled to provide wood for
and 3mm wide and linear in shape with a sharp point. Needles are longbows and destroyed to prevent the poisoning of livestock13;
set spirally around the shoot in two ranks and are not whorled2 . yew is now endangered in many parts of its range where intensive
Yew is mainly dioecious; although examples of monoecious trees land-use has caused a decline in numbers5, 15, 16 and regeneration
exist this is rare and usually consists of separate sexed branches. rates are too slow to replace the existing populations5 . Additional
Male flowers are small green globules along the underside of last pressure is now being put on the species from harvesting for the
years shoots, whilst the female flowers are minute green flowers extraction of taxanes for pharmaceutical use17. Despite being
borne in the leaf axils of the previous years growth. After wind poisonous, it is also frequently damaged by browsing and bark
pollination scarlet berries approximately 7mm in diameter are stripping by wild and domestic animals. The thin bark means it is
Map 1: Plot distribution and simplified chorology map for Taxus baccata.
produced with a red fleshy aril (the only non-toxic part of the tree4) intolerant to fire although it has a lower combustibility than other
Frequency of Taxus baccata occurrences within the field observations as
surrounding the dark central seed. Seeds are dispersed by birds reported by the National Forest Inventories. The chorology of the native conifers. Relatively few insects attack it compared to other tree
spatial range for T. baccata is derived afterMeusel andJger, and Jalas species because of its toxicity18 , but seedlings can be killed by
which feed on the red flesh and pass the seeds. The yew is also and Suominen21, 22 .
capable of vegetative reproduction. pathogenic fungi17, stem canker has been seen in Switzerland19
and has been identified as the cause of death of some yews
Distribution Habitat and Ecology in Great Britain20 . Yew mite (Cedidophyopsis psilaspis), whose
The yew is native to most of Europe, the Atlas Mountains and The yew normally appears as individuals or small groups attacks cause bud mortality7, is considered a serious pest of yew
Asia Minor. In Europe, yew woodland can be found over most areas, of trees within the understorey, but natural stands can be found in northern and central Europe3 .
but it grows best in oceanic climates with moderate temperatures3 . across its entire range. It normally occupies the mid to lower tier
Its distribution is limited in northern Europe beyond Britain, Ireland of a forest3 . Yew can grow on almost all soil types with adequate
and southern Scandinavia by low temperatures and waterlogging drainage, typically on humus and base-rich soils, but also on dry
and in the south by drought and high temperature5, 6 . Elevation rendzina and sandy soils with adequate moisture7. The yew is
increases from north to south with moisture demand for this intolerant of prolonged frost and cold although its tolerance varies
reason and in the Mediterranean area, yew is usually restricted by region and season. They are moderately drought tolerant and
to higher elevations on northern slopes3, 7. It is thought to be the can cope with temporary flooding but are susceptible to long-
oldest tree genus in Europe; the oldest fossil record for Taxus in term poor drainage. The temperature range for photosynthesis is
Europe dates to the Lower Miocene8 . greater than any other European tree species, allowing the yew to
photosynthesise in winter when deciduous trees are bare of leaves;
this contributes towards the yews extreme shade tolerance9,
which is almost comparable to that of silver fir (Abies alba) and
beech (Fagus sylvatica), the two most shade tolerant tree species
of Europe10 . The yews root system is capable of penetrating the
most compressed soils meaning they are able to survive under
Red fleshy berry-like fruits surrounding the dark central seed.
extreme conditions such as rocky terrain and vertical cliff faces11 . (Copyright Roberto Verzo, www.flickr.com: CC-BY)
[10] H.H. Ellenberg, Vegetation Ecology of [22] J.Jalas, J.Suominen, Atlas Florae
Central Europe (Cambridge University Europaeae: distribution of vascular plants
Press, 2009), fourth edn. in Europe Vol. 2 Gymnospermae (Pinaceae
Annual precipitation (mm)
Tilia cordata, Tilia platyphyllos and other limes in Europe: distribution, habitat, usage and threats
E. Eaton, G. Caudullo, D. de Rigo 16m in diameter and may be up to 2000 years old, although
precise dating is difficult as the heartwood may have rotted
Tilia cordata Mill., known as small-leaved lime, and Tilia platyphyllos Scop., known as large-leaved lime, are very similar away long ago13 . Whilst neither T. cordata nor T. platyphyllos
trees, both native to Europe and preferring warmer climates. Growing into large trees, they occur from southern Finland is as susceptible to aphid infestation as Tilia x europaea, it has
to southern Italy, and from the Caucasus to north-west Spain and Wales. Limes prefer lowlands to higher elevations, and been said that the soils underneath lime trees may receive up to
have been a component of European woodlands for six millennia. Coppicing has been a common form of management 1 kg per square metre of sugars from honeydew. This nutrient
for limes, as they produce long straight poles and can be very long-lived (>2000 years) in this form. Lime wood is much input is thought to stimulate nitrogen-fixing bacteria in the soils,
valued for carving, as it is soft and resistant to splitting. enriching them with nitrogen and phosphorus14 .
Small-leaved lime (Tilia cordata Mill.) and large-leaved lime
(Tilia platyphyllos Scop.) are large-sized deciduous broad-leaved Importance and Usage
trees. They are long-lived, able to survive more than 1000 years Frequency Both the main lime species in Europe produce a wood that is
< 25%
even if coppiced1, 2 . T. cordata is the more common species in 25% - 50% light in colour and soft enough to carve, but resistant to splitting4 .
Europe, whilst T. platyphyllos extends farther south. Both species
50% - 75%
> 75%
Some of the earliest uses of lime wood includes bows and shields,
can reach 30-40m in height with straight trunks up to around Chorology as well as bast, which is a tough fibrous material derived from
Native
1m in diameter which are largely free of suckers and epicormic the inner bark and used for rope and clothing. Coppicing of Tilia
growth, unlike their hybrid Tilia x europaea (common lime). Their has long been practiced, as the trees are capable of producing
crowns tend to be quite neat and narrow, becoming untidier as long, straight poles4 . As the wood of both T. cordata and T.
they age, although the high crown can allow a branch-free bole platyphyllos can be worked easily, it has been a highly favoured
of 10-15m. Despite their common names, the leaves of these material for carving since the Middle Ages, as well as for musical
two species are very similar: both are often around 9cm long, instruments, clogs, beehives, and cuckoo clocks15, 16 . Honey from
with T. platyphyllos up to 15cm; pointed tips to the leaves are the flowers of lime trees is also much valued, and a tea made
common to both, as are a cordate base, which is more irregular from the flowers (Tilleul) has long been thought to have anti-
in form in T. platyphyllos, a finely-toothed leaf margin, and a dark inflammatory properties17. One of the common uses of lime trees
green shiny upper surface with the underside paler3 . T. cordata has been as a street tree in much of Europe, notably along Unter
has hairs in the vein axils on the lower surface of its leaves, den Linden in the centre of historic Berlin18 .
whereas T. platyphyllos is only sometimes hairy on its underside.
Both species flower profusely in June and July. The white or
pale flowers, which are insect-pollinated, are fragrant and occur Map 1-A: Plot distribution and simplified chorology map for Tilia cordata.
Frequency of Tilia cordata occurrences within the field observations as
in clusters of 4 to 5. Seeds are first produced around 30 to 40 reported by the National Forest Inventories. The chorology of the native
years of age, and every 2-3 years trees produce a reasonable spatial range for T. cordata is derived after EUFORGEN and Afonin et al.26, 27.
crop of seeds. The seeds of T. cordata are smaller than those of T.
platyphyllos: there are 7500 T. platyphyllos seeds per kilogram, and the Alps lime stands and forests were much more abundant
compared to 29000 T. cordata seeds per kilogram4 . before the expansion and intensification of agriculture 7000-
5000 years ago10 . T. cordata can grow on calcareous soils, podzols,
and brown earths, and can compete with oaks on stagno-gley
soils, whilst T. platyphyllos is more usually found on rendzinas
formed from limestone or basic igneous rocks. If the mean
annual precipitation is greater than 850mm, T. cordata can also
move onto more lime-rich soils4 , but it is quite drought tolerant11 .
Neither species is much affected by spring nor autumn frosts,
as flushing is relatively late and buds set early11 . However, both
species require some warmth, being limited in the north of their
ranges by temperature. This is particularly true for the production
of fertile seed, as in colder regions (such as northern Britain) it is
often too cold for the pollen tube to grow following pollination12 .
As such, opportunities for limes to spread in such areas only occur
following particularly warm summers. Given its relative drought-
tolerance and its preference for warmer temperatures, the range
of this species may increase in a warming climate11 . T. cordata
and T. platyphyllos are both tolerant of shade and tend to grow in
close proximity to other species in dense woodlands. Both species Isolated small-leaved lime (Tilia cordata) in Leskova Dolina (South Slovenia).
(Copyright Stefano Zerauschek, www.flickr.com: AP)
show substantial regenerative abilities and have been grown as
coppice for millennia. Individual coppice stools may form rings
Threats and Diseases
Inflorescences of white-yellowish fragrant flowers arranged in clusters of 4-5. T. cordata and T. platyphyllos are generally quite disease
(Copyright Giovanni Caudullo: CC-BY)
resistant. Bleeding stem cankers caused by Phytophthora
cactorum and Phytophthora citricola have been recorded on
Distribution limes. T. cordata is sensitive to Phytophthora plurivora19, 20 .
T. cordata and T. platyphyllos are native to much of Europe, Aphids can be a problem, but to a much lesser extent with T.
with their ranges extending from southern Finland to southern cordata and T. platyphyllos than with common lime: in severe
Italy and the Caucasus. T. cordata is the more abundant of the infestations the honeydew dripping onto lower leaves allows sooty
two species and its core region is central and eastern Europe. It moulds to grow, blocking light from reaching the leaf surface. In
can be found as far north as southern Norway and Finland and at common with several woody plants, limes are susceptible to be
elevations up to 1500m in the central Alps5 . T. platyphyllos has attacked by the gypsy moth (Lymantria dispar) and by the nun
a smaller range, reaching slightly farther south but only reaching moth (Lymantria monacha)21-24 . In particular, T. cordata is highly
southern Sweden at its northern extent and having a much more
patchy occurrence in northern central Europe. Neither species
is present in the far west of Europe, with the western extent in
North-West Spain and Wales6 . In Europe, two other species of
lime occur naturally: the silver lime (Tilia tomentosa Moench.)
and the Caucasian lime (Tilia dasystyla Stev.) with two noticeable
subspecies caucasica and dasystyla. T. tomentosa especially
occurs in the Balkans and Hungary, while T. dasystyla is peculiar
Annual precipitation (mm)
Frequency
< 25%
25% - 50%
vulnerable to the gypsy moth and T. platyphyllos is susceptible 50% - 75%
> 75%
to the nun moth. Invertebrates to which limes play host include Chorology
References
[1] I.ColinPrentice, H.Helmisaari, Forest [16] A.Roloff, H.Weisgerber, U.M. Lang,
Ecology and Management 42, 79 (1991). B.Stimm, Bume Mitteleuropas: Von Aspe
[2] J.E. Milner, The tree book : the bis Zirbelkiefer. Mit den Portrts aller
indispensable guide to tree facts, crafts Bume des Jahres von 1989 bis 2010.
and lore (Collins & Brown, 1992). (Wiley-VCH, 2010).
[3] A.F. Mitchell, P.Dahlstrom, E.Sunesen, [17] M.Blumenthal, American Botanical
C.Darter, A field guide to the trees of Council, Bundesinstitut fr Arzneimittel
Britain and northern Europe (Collins, und Medizinprodukte, Therapeutic Guide to
1974). Herbal Medicines, The Complete German
Commission E monographs (American
[4] P.S. Savill, The silviculture of trees used in Botanical Council, 1998).
British forestry (CABI, 2013).
[18] M.Forrest, C.Konijnendijk, Urban Forests
[5] J.Jensen, S.Canger, Noble Hardwoods and Trees, C.Konijnendijk, K.Nilsson,
Network: Report of the Third Meeting, 13- T.Randrup, J.Schipperijn, eds. (Springer
16 June 1998, Sagadi, Estonia, J.Turok, Berlin Heidelberg, 2005), pp. 2348.
etal., eds. (Bioversity International, 1998),
pp. 2838. [19] M.Mrazkova, etal., Plant Disease 94, 272
(2010).
[6] J.SvejgaardJensen, EUFORGEN Technical
guidelines for genetic conservation and [20] M.Mrzkov, etal., Plant Protection
use for Lime (Tilia spp.) (2003). Science 49, 155 (2013).
[7] K.Radoglou, D.Dobrowolska, G.Spyroglou, [21] D.deRigo, etal., Scientific Topics Focus 2,
V.N. Nicolescu, Die Bodenkultur 60, 9 mri10a15+ (2016).
(2009). [22] CABI, Lymantria dispar (gypsy moth)
[8] C.D. Pigott, B.Francis, Edinburgh Journal (2015). Invasive Species Compendium.
of Botany 56, 161 (1999). http://www.cabi.org
[9] G.Lang, Quartre Vegetationsgeschichte [23] CABI, Lymantria monacha (nun moth)
Leaves of European lime are very similar, with toothed margin, Trunk of small-leaved lime (Tilia cordata), which generally reaches Europas: Methoden und Ergebnisse (G. (2015). Invasive Species Compendium.
cordate base and pointed tips. lower heights than large-leaved lime (Tilia platyphyllos). Fisher, Jena, 1994). http://www.cabi.org
(Copyright Graham Calow, www.naturespot.ork.uk: AP) (Copyright AnRo0002, commons.wikimedia.org: CC0) [10] W.Tinner, B.Ammann, Global Change and [24] I.Skrzecz, Folia Forestalia Polonica 42,
Mountain Regions, U.Huber, H.Bugmann, 41 (2000).
M.Reasoner, eds. (Springer Netherlands, [25] C.D. Pigott, J.P. Huntley, New Phytologist
2005), vol.23 of Advances in Global 84, 145 (1980).
Change Research, pp. 133143. [26] EUFORGEN, Distribution map of lime (Tilia
Frequency [11] G.E. Hemery, etal., Forestry 83, 65 cordata) (2008). www.euforgen.org.
< 25% (2010). [27] A.N. Afonin, S.L. Greene, N.I. Dzyubenko,
25% - 50%
[12] C.D. Pigott, Journal of Ecology 79, 1147 A.N. Frolov, eds., Interactive Agricultural
50% - 75% (1991). Ecological Atlas of Russia and Neighboring
> 75% Countries: Economic Plants and their
[13] C.D. Pigott, New Phytologist 112, 117
Chorology
(1989). Diseases, Pests and Weeds [Online] (2008).
Native http://www.agroatlas.ru.
[14] D.J. Mabberley, The Plant-book: A Portable
Dictionary of the Vascular Plants Second [28] EUFORGEN, Distribution map of lime (Tilia
Edition (Cambridge University Press, platyphyllos) (2008). www.euforgen.org.
1997). [29] H.Meusel, E.Jger, eds., Vergleichende
[15] A.Praciak, etal., The CABI encyclopedia of Chorologie der Zentraleuropischen Flora
forest trees (CABI, Oxfordshire, UK, 2013). - Band I, II, III (Gustav Fischer Verlag,
Jena, 1998).
Map 1-B: Plot distribution and simplified chorology map for Tilia platyphyllos.
Frequency of Tilia platyphyllos occurrences within the field observations
Bark of small-leaved lime (Tilia cordata) forming longitudinal fissures
as reported by the National Forest Inventories. The chorology of the native
at the base of the trunk.
spatial range for T. platyphyllos is derived after EUFORGEN28 .
(Copyright Stefano Zerauschek, www.flickr.com: AP)
Hairs in the vein axils on the leaf lower face of small-leaved lime (Tilia cordata).
(Copyright David Nicholls, www.naturespot.ork.uk: AP)
Frequency
< 25%
25% - 50%
50% - 75%
> 75%
Chorology
Native
White elm (U. laevis) is a riparian tree growing principally along rivers.
(Copyright Stefano Zerauschek, www.flickr.com: AP)
water decay, so that timbers are used as underwater piles, in > 75%
Chorology
shipbuilding and for water pipes28, 29 . U. glabra grows at relatively Native
Map 1-C: Plot distribution and simplified chorology map for Ulmus minor.
Frequency of Ulmus minor occurrences within the field observations as
Threats and Diseases
reported by the National Forest Inventories. The chorology of the native The story of the elms is marked by the two destructive
spatial range for U. minor is derived after several sources15, 16, 47, 48, 51 .
epidemics, caused by the introduction of hyper-virulent fungal
living support for grapevines, also providing fodder in summer pathogens, Ophiostoma ulmi in the 1920s and Ophiostoma
and small-wood and poles when coppiced. This practice was then novo-ulmi in 1970s, usually called Dutch elm disease (DED). It
exported by the Romans to the whole of Europe35 . In fact, thanks is considered one of the most significant tree diseases known
to its fast-growing, re-sprouting ability and easy propagation by in the world, having devastated the elm populations throughout
suckering, U. minor has been widely propagated and planted in Europe and North America37. DED is very aggressive and is
the countryside and more recently on roadsides and in urban transmitted by the bark beetles of genus Scolytus or throughout
areas6 . However, the pandemics of the vascular Dutch elm disease root connections, attacking principally mature U. minor leading to
starting in the last century devastated the elm populations in the death in 2-3 years38 . U. glabra is susceptible to the fungal agent,
whole of Europe, making it necessary to substitute it with other but remains unaffected at higher latitudes, where the insect
tree species or with other Dutch elm disease-resistant elms, such vector cannot survive. U. laevis is susceptible, but less attractive
as Siberian elm and its hybrids36 . for the bark beetles, so its population was not severely damaged
by the DED, except in the western ranges6 . Several strategies
have been adopted in an attempt to prevent infection and
Taxonomy of elms reduce the impacts of DED. The chemical and biological controls
The genus Ulmus includes 20-40 species prevalently in the temperate against the pathogen or the insect vector encountered many
regions of the northern hemisphere and concentrated mostly in Eurasia1 . limits, due to poor results or high costs when utilized at large
The ambiguity in number of species results from the difficulty in their scales39 . Two breeding programs arose in Netherlands and in
classification, which is very complex and is still under debate. In fact elms Italy after the pandemics, having a success for selecting cultivars
have a strong intra-specific variability in morphological traits, not only resistant to DED with the use of Asian parental materials. Most
Seed germination of the field elm (U. minor). geographically, but even in the same tree during growing stages as a of those hybrids were planted in substitution of garden elms38, 40 .
(Copyright Franco Giordana, www.actaplantarum.org: AP)
response to environmental conditions. Moreover, they have an extensive However, the widespread use of cultivars has increased the risk
natural hybridisation capacity between them2 . With the use of chloroplast of genetic pollution, mainly when planted in the countryside
DNA, a simplified classification of European elms has been proposed. It near to natural forests6 . Despite their drastic reduction during the
Frequency
counts three large species belonging to two sections, which comprise last century, the European elm species (in a narrow sense) can
< 25%
25% - 50% different varieties and subspecies historically described3, 4: Ulmus laevis be considered not actually in danger. The main losses regarded
50% - 75%
Pall. (European white elm) belonging to the Blepharocarpus section, Ulmus introduced and cultivated plants in urban areas and countryside
> 75%
Chorology glabra Huds. (wych elm) and Ulmus minor Mill. (field elm) belonging to with relatively poor genetic variability6 . Concerning elms in forest
Native
the Ulmus section. U. minor can be defined as a complex of species, habitats, although larger trees are heavily affected by DED, in the
sometimes described on the basis of morphological traits and geographical United Kingdom and southern Sweden there has been reported
distribution as separated species: i.e. Plots Elm (Ulmus plotii), the English significant resilience of a fast turnover population of poles in the
elm (Ulmus minor var. vulgaris syn. Ulmus procera) and the grey elm understorey41, 42 . Particular attention in conservation has been
(Ulmus minor subsp. canescens syn. Ulmus canescens)5, 6 . U. glabra has also given to small and marginal populations at the borders of
a strong latitudinal variations and two subspecies have been proposed: U. their natural distributions, with the aim of minimising the risk
glabra subsp. glabra in the southern range and U. glabra subsp. montana of genetic loss. Among the European elms, U. laevis is the less
in the northern range. A variety is also recognised in Caucasus (U. glabra susceptible to DED. However, its populations are often in small
var. trautvetteri), which is sometimes treated as a separate species (Ulmus and fragmented stands as a result of the reduction of the
elliptica)7, 8 . The variety Ulmus laevis var. celtidea syn. Ulmus celtidea is riparian forests, caused by water-basin regulation and demand
an endemic elm native of northern Ukraine, which differs morphologically for agricultural lands. Potentially, genetic drift as a result of
from U. laevis6, 9, 10 . Finally U. glabra and U. minor can naturally hybridise
in intermediate forms, commonly referred to as Ulmus x holandica (Dutch
Map 1-B: Plot distribution and simplified chorology map for Ulmus laevis. elm), and both hybridise with Ulmus pumila (Siberian elm), recently
Frequency of Ulmus laevis occurrences within the field observations as
reported by the National Forest Inventories. The chorology of the native
introduced from Asia.
spatial range for U. laevis is derived after several sources16, 46, 50 .
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[4] R. H. Richens, Elm (Cambridge University forest trees (CABI, Oxfordshire, UK, 2013).
Press, 1983). [30] H. Dcamps, M. Fortun, F. Gazelle, G.
[5] E. Collin, I. Bilger, G. Eriksson, J. Turok, The Pautou, Landscape Ecology 1, 163 (1988).
Seedling of field elm (U. minor). Field elm (U. minor) isolated tree in rural area. Elms: Breeding,Conservation, and Disease [31] A. Schnitzler, The Botanical Review 63,
(Copyright Franco Giordana, www.actaplantarum.org: AP) This elm rarely exceeds 20m in height. Management, C. P. Dunn, ed. (Springer US, 40 (1997).
(Copyright Ptelea, commons.wikimedia.org: PD)
2000), pp. 281293.
[32] G. F. Borlea, Investigacin Agraria:
[6] E. Collin, Noble Hardwoods Network: Sistemas y Recursos Forestales 13, 29
Report of the Fourth Meeting, 4-6 (2004).
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the Fifth Meeting, 17-19 May 2001, [33] C. Bosco, D. de Rigo, O. Dewitte, J. Poesen,
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[7] P. Uotila, Sorbifolia 28, 5 (1997). Nicoll, A. Achim, Slope Stability and
Erosion Control: Ecotechnological
[8] A. Buchel, The Elms - Breeding, Solutions, J. E. Norris, et al., eds. (Springer
Conservation, and Disease Management, Netherlands, 2008), pp. 167210.
C. P. Dunn, ed. (Springer, 2000), pp.
351358. [35] H. M. Heybroek, Mitteilungen der
Deutschen Dendrologischen Gesellschaft
[9] V. L. Komarov, et al., Flora of the USSR - nr. 88 (2003), pp. 117119.
Volume V (Keter Press, Jerusalem, 1970).
[36] A. Solla, et al., Forest Science 51, 134
[10] H. J. Elwes, A. Henry, The Trees of Great (2005).
Britain and Ireland Vol. 7 (Privately
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(1991).
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of Britain and northern Europe (Collins, [38] L. Mittempergher, A. Santini, Investigacin
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13, 161 (2004).
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caused by biotic agents - final report,
[13] M. Goldstein, G. Simonetti, M. Watschinger, Tech. Rep. KH-32-13-151-EN-N (2011).
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[14] R. Petrokas, Baltic Forestry 14, 204 Commission (DG ENV).
(2008). [40] A. Santini, A. Fagnani, F. Ferrini, L.
[15] E. Hultn, M. Fries, Atlas of North European Ghelardini, L. Mittempergher, HortScience
vascular plants (North of the Tropic of 42, 712 (2007).
Cancer), Vols. I-III. (Koeltz scientific books, [41] G. F. Peterken, E. P. Mountford, Journal of
1986). Ecology 86, 205 (1998).
[16] J. Jalas, J. Suominen, Atlas Florae [42] G. von Oheimb, J. Brunet, Acta Oecologica
Europaeae: distribution of vascular 31, 229 (2007).
plants in Europe Vol. 3 Salicaceae to
Balanophoraceae (Committee for Mapping [43] D. de Rigo, et al., Scientific Topics Focus 2,
the Flora of Europe and Societas Biologica mri10a15+ (2016).
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[17] H. Meusel, E. Jger, eds., Vergleichende Hajek, Agricultural and Forest Entomology
Chorologie der Zentraleuropischen Flora 11, 359 (2009).
- Band I, II, III (Gustav Fischer Verlag, [45] N. A. Straw, N. J. Fielding, C. Tilbury, D.
Jena, 1998). T. Williams, D. Inward, Forestry 88, 84
[18] J. C. Lpez-Almansa, Investigacin Agraria: (2015).
Sistemas y Recursos Forestales 31, 17 [46] A. N. Afonin, S. L. Greene, N. I. Dzyubenko,
(2004). A. N. Frolov, eds., Interactive Agricultural
[19] R. Petrokas, V. Baliuckas, Baltic Forestry Ecological Atlas of Russia and Neighboring
20, 238 (2014). Countries: Economic Plants and their
Diseases, Pests and Weeds [Online]
[20] H. Hytteborn, A. A. Maslov, D. I. Nazimova, (2008). http://www.agroatlas.ru.
L. P. Rysin, Ecosystems of the World, Vol.
6: Coniferous Forests, F. A. Andersson, ed. [47] B. Jonsell, ed., Flora Nordica, Vol 1:
(Elsevier, 2005), pp. 2399. Lycopodiaceae - polygonaceae (The Royal
Swedish Academy of Sciences, Stockholm,
[21] G. F. Peterken, Woodland Conservation 2000).
and Management (Springer Science &
Flowers of white elm (U. laevis) are clustered on long stems. Business Media, New York, 1993). [48] R. Ala Miranda, et al., Regiones de
(Copyright Velella, commons.wikimedia.org: PD) procedencia de especies forestales en
[22] H. H. Ellenberg, Vegetation Ecology of Espaa (Organismo Autnomo Parques
Central Europe (Cambridge University Nacionales, Madrid, 2009).
Press, 2009), fourth edn.
Autoecology diagrams based on harmonised field [49] Sociedade Portuguesa de Botnica, Flora-
[23] L. Saarse, S. Veski, Proceedings of the
Field data in Europe (including absences) Observed presences in Europe observations from forest plots for Ulmus minor. Estonian Academy of Sciences, Geology
On: Flora de portugal interactiva (2014).
http://www.flora-on.pt.
50, 51 (2001).
[50] EUFORGEN, Distribution map of European
[24] U. Bohn, et al., Karte der natrlichen white elm (Ulmus laevis) (2008).
Vegetation Europas; Map of the www.euforgen.org.
Natural Vegetation of Europe
Average temperature of the coldest month (C)
Achene: a simple dry fruit containing a single seed. [see Fig. 3a overleaf]. (for example, the frequency of field observations concerning a given tree scrub species associated principally with calcareous rocky soils and hot
Acuminate: gradually tapering to a point [see Fig. 4f overleaf]. species) which considers multiple spatial scales to account for the different arid climates. This formation often develops as consequence of a degraded
local density of data. Furthermore, C-SMFA is constrained to be consistent maquis vegetation after fires or intense exploitations.
Adventitious: a structure arising from an unusual place; e.g. roots growing with an additional available information to improve the overall quality of the
from the shoot system. Genotype: the genetic makeup of an organism or group of organisms with
frequency analysis. In this Atlas, S-SMFA is exploited to estimate the relative reference to a single trait or set of traits, as distinguished from the physical
Aeolian soil: soil generated with sediments eroded, transported and probability of presence of forest tree species. appearance.
deposited by the action of the wind. Constraint: for the meaning in computational modelling, see semantic Geospatial: in computational science, it refers to data or information
Agroforestry: a land-use system with two or more interacting plant species, constraint. which is geographically distributed and covers significantly broad spatial
where one of them is a tree or shrub. Cordate: having a heart-shaped outline. [see Fig. 4b overleaf] . extents. Under these circumstances, for example the simple approximation of
Alkaloid: a group of natural chemical compounds containing basic nitrogen Coriaceous: leathery. the portion of Earths surface covered by the spatial extent as a geometrical
atoms produced by a large variety of organisms (bacteria, fungi, plants, plane is no more valid.
animals). They can be extracted and used in traditional and modern medicine, Corolla: the inner circle of petals of a flower, usually of a colour other than
green. Glabrous: smooth; without hair or scales.
interacting with the metabolic systems of humans and animals.
Corvid: a bird belonging to the crow family Corvidae, e.g. jays, magpies, Glacial-interglacial cycles: Continental ice-sheets in the Northern
Allelopathy (adj. allelopathic): negative effects created by a plant on the Hemisphere have grown and retreated many times in the past. Times with
growth or development of other nearby plants by releasing toxic substances. choughs.
large ice-sheets are known as glacial (cold) periods that were separated by
Anamorph (adj. anamorphic): in mycology, an asexual reproductive stage in Corymb: a flat-topped flower cluster in which the individual flower stalks (warm) interglacial periods. We are currently in an interglacial period called
a fungus life-cycle, which also includes a teleomorph, the sexual reproductive grow upward from various points on the main stem to approximately the the Holocene that initiated 11700 years BP.
stage. In the past some fungi had a double nomenclature related to their same overall height. [see Fig. 1b overleaf] .
Glaucous: having a pale greyish or bluish green colour, often with a whitish
different stages, recently declared scientifically invalid. Cotyledon: an embryo tissue within the seed which upon germination forms bloom which may be rubbed off.
Androdioecious: species having separate plants with staminate flowers and the embryonic leaf/leaves of the plant.
Globose: ball-shaped.
plants with perfect fully-functional flowers [see Fig. 2b overleaf]. Coumarin: an organic compound naturally present in many plants. It has
a sweet odour, recognised as the scent of new-mown hay. It is used in Gynodioecious: species having separate plants with pistillate flowers and
Androecious: the plant having the male flowers in a dioecious population. plants with perfect fully-functional flowers [see Fig. 2b overleaf].
perfumes, for giving aroma to drinks and tobaccos, and more recently for
Androgynomonoecious: species having in the same plant perfect fully- Gynoecious: the plant having the female flowers in a dioecious population.
clinical medical purposes.
functional flowers, pistillate and staminate flowers. [see Fig. 2b overleaf]
Cryptic: see refugia Gynomonoecious: species having in the same plant perfect fully-functional
Andromonoecious: species having in the same plant perfect fully-functional flowers and pistillate flowers [see Fig. 2b overleaf]
flowers and staminate flowers. [see Fig. 2b overleaf] Cuneate: having the shape of a wedge, triangular at the base and tapering to
a point. [see Fig. 4g overleaf]. Gynosterility (adj. gynosterile): see stamen
Androsterility (adj. androsterile): see pistil.
Cyme: A usually flat-topped or convex flower cluster in which the main stem Habitat suitability: (acronym: HS) potential suitability for a certain
Anthropogenic: caused or produced by humans. organism (e.g. a tree species) to live in a given local habitat. It is generally
and each branch end in a flower.
Apomixis (adj. apomictic): reproduction of plants where the embryo develops a quantity varying from 0 (0%, unsuitable habitat) to 1 (100%, potentially
without fertilisation occurring. Decumbent: spreading horizontally with the ends growing upwards
very suitable habitat). Computational science may be required for HS to be
Apophysis: A natural outgrowth, swelling or enlargement. Dehesa: a Spanish word which refers to the savannah-like open woodland quantitatively estimated. Depending on the objective to be investigated, HS
traditionally managed as a multi-purpose agricultural system in the Iberian may consider interspecific competition and other ecological aspects, or may
Arborescent: having the shape of a tree, tree-like. Peninsula. In Portugal it is named montado. This agro-sylvo-pastoral land instead just consider bioclimatic limitations. In the latter case, the estimated
Arcto-Tertiary geoflora: floral assemblage that once covered the Northern use provides firewood, shading and secondary products (i.e. acorn, olive, nuts, HS constitutes a potential limit which may be altered by the local ecology
Hemisphere, from roughly the late Mesozoic to mid Cenozoic Eras. cork, manna, etc.) from trees and pasture for livestock from grass land. Main (typically, the bioclimatic-related HS of a certain taxon may be higher than
Aril (syn. arillus): a fleshy fruit that covers or is attached to the seed, often tree species managed as dehesa/montado are principally Mediterranean the actual one due to competing taxa which are more fit in a given habitat).
edible to encourage dispersion by animals. evergreen oaks, holm oak (Quercus ilex), cork oak (Quercus suber), Lusitanian
oak (Quercus faginea) and Pyrenean oak (Quercus pirenaica), but also ash Heartwood: the older, non-living central wood of a tree, consisting of non-
Array: of data (including spatial data), information or numerical models. In (Fraxinus angustifolia) and stone pine (Pinus pinea) with other accompanying functioning xylem tissue that has become blocked with resins, tannins and
computational science, collection of homogeneous or heterogeneous layers tree such as olive (Olea europaea) and junipers (Juniperus oxycedrus and J. oils and providing mechanical support. Heartwood is usually darker and
(for example, field observations of forest tree species; land cover maps on thurifera). harder than the younger sapwood.
the distribution of broadleaved/coniferous taxa; maps on bioclimatic and Hectare: one hectare is 100 100 metres.
geographic information) which may be processed in a compact way (array Dendrochronology: the science of dating based on the analysis of tree ring
programming) as a sequence within numerical models. patterns. Heliophilous: organism that needs, or tolerates, a high level of direct sunlight.
Arthropod: an invertebrate animal with an external skeleton (exoskeleton), Dioecious: species that has male and female flowers (or reproductive Hermaphrodite: organism or reproductive organs that have both male and
a segmented body, and jointed appendages. Arthropods include the insects, structures) in separate plants. female structures. Relating to flowers, hermaphrodite flowers can be called
arachnids, myriapods, and crustaceans. also bisexual or perfect. [see Fig. 2b overleaf].
Diploid: organism which contains two sets of chromosomes, one set inherited
Autoecology: the term generally refers to diagrams also known as climate- from each parent. Hilium: a scar or a mark remaining on a seed or spore at its point the site of
space diagrams. For a given taxon, an empirical autoecology diagram is its former attachment.
Disking: Agricultural term to describe the process of using a disk, or harrow,
based on the available field observations to illustrate the pattern of selected to disturb the soil, setting back natural succession by cutting up grassy Holocene: The Holocene is the geological epoch that began approximately
bioclimatic quantities (for example, the average annual temperature versus vegetation, thus preventing an area from maturing into briars, shrubs, and 11700 years BP and continues to the present.
the total annual precipitation) associated with the locations where the taxon trees. Hygrophylous: adapted for growth in a wet or damp environment.
has been observed. When available (as is the case in this Atlas), autoecology Dormant (adj. dormancy): an induced and hormone-regulated stage of Hypoglycin A: a natural amino acid which occurs in several plants; this
diagrams also illustrate the empirical climate space occupied by all the taxa inactive state in which growth stops and metabolism is slowed. compound is toxic if ingested, causing profound hypoglycaemia.
(presence/absence autoecology).
Drupe: a fruit with the seed inside a woody shell (endocarp) surrounded by a Imbricate: Having regular overlapping edges
Axil: the upper angle between one part of a plant and another part. fleshy part (mesocarp) and skin (exocarp). [see Fig. 3b overleaf].
Auriculate: having an auricle, i.e. an ear-shaped lobe at the base of a leaf. Inflorescence: a grouping of flowers on a stem or in a leaf axil.
Ecotone: the transition zone between two different plant communities, e.g. Interstadial: Time period corresponding to a temporary period of thaw in the
Big data: in computational science, collection of data whose appropriate between forest and prairie.
analysis and computational modelling requires specific techniques in order middle of an ice age.
Ectomycorrhiza: a fungus that forms a symbiotic relationship with the roots Involucre: a ring of small leaves, or bracts, at the base of a flower, flower
for the challenging amount of information to be managed. Usual standard
of various plant species. Unlike other mycorrhizae, they do not penetrate their cluster, or fruit.
approaches of numerical processing are generally inadequate with big data
hosts cell walls, developing in the intercellular spaces. They form a dense
due to their dimension or complexity/heterogeneity. The qualification of data Iron Age: in archaeology, the Iron Age (in Europe between about 1200BC
hyphal sheath surrounding the root surface, helping the host plant to take up
as big data highly depends on the typical computational resources, software 1BC) refers to the advent of ferrous metallurgy to produce hard metal
water and nutrients and to access carbohydrate supplies.
and knowledge available in given historic time. tools. The adoption of iron coincided with other changes in past cultures, often
Edaphic: related to the soil. including more sophisticated agricultural practices and artistic techniques.
Bioclimatic: referring to climatic aspects relevant for the biology of
the organism (for example, referring to temperature, precipitation, Emarginate: having a broad shallow notch at the apex. [see Fig. 4f overleaf] Karst (adj. karstic]: terrain or landscape characterised by largely bare, rocky
evapotranspiration, solar irradiation, etc.). Empyreuma: the smell and taste associated with burning vegetable and surface and irregular limestone in which erosion has produced fissures,
Biogenic volatile organic compounds: volatile organic compounds produced animal matter. Empyreumatic oils are obtained by distilling various organic sinkholes, underground streams and caverns and underneath which rivers flow.
by plants. Involved in plant growth, development, reproduction and defence. substances at high temperatures. Lanceolate: having a narrow shape and tapering to a point at each end, with
While contributing to tropospheric ozone and secondary particle formation, they Endemic (adj. endemism): a species that lives in a defined and restricted the base slightly wider. [see Fig. 4b overleaf].
also have indirect effects on cloud formation and climate change. geographical zone or habitat type. Lenticel: small raised corky spot or line through which gaseous exchange
Bisexual: see monoecious Endocarp: see drupe occurs. They typically appear on bark but also on fruits.
BP: acronym standing for Before Present with Present being the 1st January Entomophilous: form of pollination whereby pollen or spores are distributed Lignicolous: organism which grows or lives on or in wood.
1950, reflecting the time when radiocarbon dating became practicable. by insects. Maquis (syn. matorral): this is the main vegetation along the Mediterranean
Brachyblast: branches with shortened internodes (brachyblasts) as opposed Epicormic: growing from the bark of the trunk, stem or branch. coasts, where summers are hot and dry, and rain falls abundantly in winter.
to branches with long internodes. It contains densely packed bushes, shrubs and evergreen trees with generally
Epiphyte: a plant that grows harmlessly upon another plant, deriving its
Bronze Age: the term Bronze Age refers to a period in human cultural moisture and nutrients from air and rain. In the temperate zone they are coriaceous leaves, occasionally deciduous during the summer.
development when the metalworking included techniques for smelting copper principally mosses, lichens, algae and liverworts. Marcescence (adj. marcescent): deciduous tree that retains its leaves
and tin from naturally occurring outcrops of ores, and then combining them through the winter, shedding them only when new leaves emerge in spring.
Eutrophic: characterised by an abundant accumulation of nutrients that
to cast bronze to produce sophisticated tools such as sickles or knives. This is
support a dense plant life. Marl: a crumbly mixture of clays, calcium and magnesium carbonates, and
the earliest period in which some civilisations introduced writing.
Fascicle: A bundle or cluster of stems, flowers or leaves. remnants of shells that is sometimes found under desert sands and used as
BVOC: See biogenic volatile organic compounds. fertiliser for lime deficient soils.
Calyx: the outer floral envelope that protects the developing flower bud, Fastigate: having erect branches, often appearing to form a single column
with the stem. Matorral: see maquis
consisting of the sepals.
Fennoscandia: a region comprising the Scandinavian Peninsula, Finland, Maximum habitat suitability: (acronym: MHS) referring to bioclimatic-
Cambium: the tissue in the stems and roots of plants, consisting of cells that based habitat suitability, it represents the maximum spatial extent where
divide rapidly to form new layers of tissue. In woody plants it lies between Karelia and the Kola Peninsula.
a given taxon is suitable to live. It may also be referred to as survivability.
the bark and wood of the stem; on its outer surface the vascular cambium Field observations: data referring to measurements performed in the Complementary to the average habitat suitability (AHS), which only focuses
forms new layers of phloem, and on its inner surface, new layers of xylem. environment where a certain organism or association of organisms lives on the optimal ecological niche of a given taxon. MHS is greater or equal to
Chasmophyte (adj. chasmophytic): plant growing in rock crevices. (for example, measurements of whether a certain tree species is found in a AHS. Low values of MHS may be exploited to identify the less suitable areas
certain local environment). Complementary to laboratory measurements and for a given taxon to survive, irrespective of other ecological aspects such as
Chorology: study of the spatial distribution of organisms. computational modelling estimations. interspecific competition. Conversely, correspondingly low values of AHS may
Climate-space diagrams: see autoecology. Flexuose: bent from side to side in one plane in zigzag form. only identify non-optimal areas. In these areas, the taxon is in a sub-optimal
Computational model: mathematical model in computational science Forest plot: local-scale field observation concerning forest resources. condition where it may or may not be able to survive, depending on other
requiring computational resources to analyse or estimate specific statistics Forest plots are often the basis of national forest inventories and of other factors.
and information on the behaviour of a natural or artificial system (for thematic datasets on forests. Melliferous: producing or promoting the production of honey.
example, the relative probability of presence or the habitat suitability
of forest tree species). Frugivore: (adj. frugivorous) an organism that feeds on fruit. Mesic: characterised by or adapted to a moderately moist habitat.
Computational science: scientific and technical methods to analyse Galbulus (pl. galbuli): a fleshy berry-like seed cone of junipers and cypresses. Mesocarp: see drupe.
existing information and to estimate unavailable information with the help of Gall (syn. cecidium): an abnormal outgrowth of plant tissue, similar to Mesoclimate: the climate of a small area not representative of the general
mathematics and computational models. tumours, caused by parasites, principally insects, and used as a habitat for climate of the district
Constrained spatial multi-frequency analysis: (acronym: C-SMFA) in development and as a food source. Mesolithic: This period began at the end of the Pleistocene epoch, around
computational science, a spatial frequency analysis of a given quantity Garrigue: a discontinuous Mediterranean vegetation composed of low bushy 9200 BC, and ended with the introduction of agriculture, the date of which
Petal (Corolla)
Ovary Perianth
Sepal (Calyx)
Peduncle Receptacle flower with both male flower male flower female
and female structure structure structure
Androdioecious
Raceme Compound raceme Spike Catkin
Dioecious
Androecious Gyneocious
Gynodioecious
Fig. 3: Fruits
Fig 3a: Dry Fig 3b: Fleshy
Cordate Deltoid Elliptical Lanceolate Linear Oblanceolate Oblong Obovate Ovate Rhomboid Spathulate
- compound
Aristate Cuspidate Acuminate Spine-tipped Bristle-tipped Mucronate Attenuate Cuneate Rounded Truncate Oblique
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Trees, as all other forms of life, adapt to the conditions in which they live.
This adaptation is reflected in the shape of their leaves, their seeds, and their
strategy for dissemination and expansion. Each chapter of the Atlas presents,
in addition to the description of each tree species, high quality graphics and
photographs showing the climatic preferences and singularities of the different
species of trees.
ISBN 978-92-79-52833-0