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The European Forest Plant Species List (EuForPlant): Concept and
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Article in Journal of Vegetation Science · May 2022
DOI: 10.1111/jvs.13132
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Received: 7 May 2021
| Revised: 14 March 2022 | Accepted: 26 March 2022
DOI: 10.1111/jvs.13132
Journal of Vegetation Science
RESEARCH ARTICLE
The European Forest Plant Species List (EuForPlant): Concept
and applications
Thilo Heinken1
| Martin Diekmann2
| Jaan Liira3
| Anna Orczewska4
|
5
6
7
8
Marcus Schmidt
| Jörg Brunet
| Milan Chytrý
| Olivier Chabrerie
|
8
9
7
Guillaume Decocq
| Pieter De Frenne
| Pavel Dřevojan
|
10
11
12
Zbigniew Dzwonko
| Jörg Ewald
| Jon Feilberg
| Bente Jessen Graae13
John-­Arvid Grytnes14
| Martin Hermy15
| Wolf-­Ulrich Kriebitzsch16 |
Māris Laiviņš17 | Jonathan Lenoir8
| Sigrid Lindmo13
| Damien Marage18
|
Vitas Marozas19
| Thomas Niemeyer20
| Jaanus Paal3
Elle Roosaluste3 | Jiří Sádlo21
| Joop H.J. Schaminée23
Kris Verheyen9
| Monika Wulf25
| Thomas Vanneste9
| Petr Pyšek21,22
| Torbjörn Tyler24
|
|
|
1
General Botany, University of Potsdam, Potsdam, Germany
2
Vegetation Ecology and Conservation Biology, FB02, University of Bremen, Bremen, Germany
3
Institute of Ecology and Earth Sciences, University of Tartu, Tartu, Estonia
4
Institute of Biology, Biotechnology and Environmental Protection, University of Silesia, Katowice, Poland
5
Northwest German Forest Research Institute, Department E (Forest Nature Conservation), Hannoversch Münden, Germany
6
Southern Swedish Forest Research Centre, Swedish University of Agricultural Sciences, Uppsala, Sweden
7
Department of Botany and Zoology, Faculty of Science, Masaryk University, Brno, Czech Republic
8
Ecologie et Dynamique des Systèmes Anthropisés (EDYSAN, UMR CNRS 7058), Université de Picardie Jules Verne, Amiens, France
9
Forest & Nature Lab, Ghent University, Ghent, Belgium
10
Institute of Botany, Jagellonian University, Kraków, Poland
11
Botany & Vegetation Science, University of Applied Sciences, Freising, Germany
12
Kastrupvej 8, Haraldsted, Ringsted, Denmark
13
Department of Biology, Norwegian University of Science and Technology, Trondheim, Norway
14
Department of Biological Sciences, University of Bergen, Bergen, Norway
15
Division Forest, Nature and Landscape, University of Leuven, Leuven, Belgium
16
Institute for World Forestry, Johann Heinrich von Thünen Institute (vTI), Hamburg, Germany
17
Latvian State Forest Research Institute Silava, Salaspils, Latvia
18
University of Franche-­Comte, UMR TheMA-­CNRS, Besançon, France
19
Agriculture Academy, Vytautas Magnus University, Kaunas, Lithuania
20
Institute for Ecology, Leuphana University Lüneburg, Lüneburg, Germany
21
Czech Academy of Sciences, Institute of Botany, Department of Invasion Ecology, Průhonice, Czech Republic
22
Department of Ecology, Faculty of Science, Charles University, Prague, Czech Republic
23
Alterra Wageningen UR, Wageningen, the Netherlands
24
Department of Biology, Botanical Museum, Lund University, Lund, Sweden
25
Leibniz Centre for Agricultural Landscape Research (ZALF), Research Area 2, Müncheberg, Germany
Jon Feilberg and Wolf-­Ulrich Kriebitzsch authors are retired. © 2022 International Association for Vegetation Science
J Veg Sci. 2022;33:e13132.
https://doi.org/10.1111/jvs.13132

wileyonlinelibrary.com/journal/jvs
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Correspondence
Thilo Heinken, General Botany, University
of Potsdam, Potsdam, Germany.
Email: heinken@uni-potsdam.de
Funding information
We are grateful to the Research
Foundation –­Flanders (FWO) for
funding the scientific research network
FLEUR. PDF and TV were supported by
the European Research Council (ERC)
through the ERC Starting Grant FORMICA
(Grant/Award Number: 757833) and
KV though the ERC Consolidator Grant
PASTFORWARD (Grant/Award Number:
614839). JL was funded by the European
Regional Development Fund (the
Centre of Excellence EcolChange) and
the Estonian Research Council (Grant/
Award Number: PRG1223). MC and PD
received funding from the Czech Science
Foundation (Grant/Award Number:
19-­28491X), while PP was supported by
EXPRO grant (Czech Science Foundation,
Grant/Award Number: 19-­28807X) and
long-­term research development project
RVO (Czech Academy of Sciences, Grant/
Award Number: 67985939)
Co-ordinating Editor: Zoltán Botta-­Dukát
HEINKEN et al.
Abstract
Question: When evaluating forests in terms of their biodiversity, distinctiveness and
naturalness, the affinity of the constituent species to forests is a crucial parameter.
Here we ask to what extent are vascular plant species associated with forests, and
does species’ affinity to forests vary between European regions?
Location: Temperate and boreal forest biome of Northwestern and Central Europe.
Methods: We compiled EuForPlant, a new extensive list of forest vascular plant species in 24 regions spread across 13 European countries using vegetation databases
and expert knowledge. Species were region-­specifically classified into four categories
reflecting the degree of their affinity to forest habitats: 1.1, species of forest interiors;
1.2, species of forest edges and forest openings; 2.1, species that can be found in
forest as well as open vegetation; and 2.2, species that can be found partly in forest,
but mainly in open vegetation. An additional “O” category was distinguished, covering
species typical for non-­forest vegetation.
Results: EuForPlant comprises 1,726 species, including 1,437 herb-­layer species, 159
shrubs, 107 trees, 19 lianas and 4 epiphytic parasites. Across regions, generalist forest
species (with 450 and 777 species classified as 2.1 and 2.2, respectively) significantly
outnumbered specialist forest species (with 250 and 137 species classified as 1.1 and
1.2, respectively). Even though the degree of shifting between the categories of forest affinity among regions was relatively low (on average, 17.5%), about one-­third of
the forest species (especially 1.2 and 2.2) swapped categories in at least one of the
study regions.
Conclusions: The proposed list can be used widely in vegetation science and global
change ecology related to forest biodiversity and community dynamics. Shifting of
forest affinity among regions emphasizes the importance of a continental-­scale forest
plant species list with regional specificity.
KEYWORDS
biogeographical regions, boreal zone, expert knowledge, forest affinity, forest plant species,
habitat shift, nemoral zone, species diversity, vascular flora, woodland
1
|
I NTRO D U C TI O N
existed for a long time with different aims. Classical phytosociology
is, among other goals, concerned with identifying diagnostic (i.e. dif-
When forests are evaluated in terms of their biodiversity, distinc-
ferential and character) species for specific vegetation types. Braun-­
tiveness and naturalness, the question of the forest affinity of each
Blanquet (1918, 1964) introduced the term “fidelity” as a measure of
plant species venturing into forest systems is crucial (Hermy et al.,
the association of a species and a certain vegetation type. Five levels
1999; Verheyen et al., 2003; Jaroszewicz et al., 2019; Schneider
of fidelity were distinguished, ranging from species almost exclusively
et al., 2021). For various ecological research questions related to
associated with a given plant community to accidental species (e.g.,
forest ecosystems (e.g., the influence of alien tree species, global
rare, more or less random sprinklings or relic occurrences from pre-
changes and land-­use legacies on forest biodiversity; Ammer et al.,
ceding communities). These fidelity levels were modified and further
2018), it is important to separate forest-­specific species from habitat
developed by various authors (Barkman, 1989; Bergmeier et al., 1990).
generalists and open-­land species. This is only possible in a com-
Fidelity is also a key concept in up-­to-­date numerical approaches to
prehensible way using species lists that are derived from a broad
define diagnostic species or discriminate species groups in phytoso-
consensus and on a continental scale as a reference.
ciology (Bruelheide, 2000; Chytrý et al., 2002; Tichý et al., 2019).
Approaches to define the affinity of plant species to a given habitat
Although phytosociological character species (i.e. species that
type (e.g., forest, grassland, scrub) or a specific plant community have
preferably occur in a single vegetation type and thus have high
HEINKEN et al.
Journal of Vegetation Science
| 3 of 16
fidelity to it) have often been used to analyse the properties of for-
evaluation of species that are widespread in both forest and open
est species (Bossuyt & Hermy, 2001; Ewald, 2003) and to quantify
vegetation as typical forest plants. Moreover, habitat shifts of spe-
forest biodiversity (Dupré, 2000; Willner et al., 2009; Slabejová
cies were also considered through regionalization. The list was sub-
et al., 2019), this approach has some serious shortcomings in this
sequently improved and supplemented with lists of forest-­dwelling
context. First, a plant species can have a different amplitude towards
bryophyte and lichen species by Schmidt et al. (2011). More recently,
certain ecological factors in different vegetation formations (forests,
a list of selected groups of forest animals has also been published for
scrub, open land). However, assigning a given plant species to a sin-
Germany, using a slightly modified approach (Dorow et al., 2019, see
gle formation in which it chiefly occurs, as for instance in the classi-
also Schneider et al., 2021).
fication of sociological behaviour in Ellenberg et al. (1992), does not
To date, the German forest species list has been applied in ca.
mirror its ecological behaviour. For example, Ellenberg et al. (1992)
200 scientific publications, including analyses of the effects of man-
assigned Caltha palustris exclusively to wet meadows (Calthion), al-
agement (Heinrichs et al., 2019) and land-­use history (Naaf & Kolk,
though the species also occurs with high frequency in moist forests.
2015) on forest plant diversity and species composition, analyses of
The same applies to heliophilous and drought-­tolerant plant species
long-­term vegetation changes (Naaf & Kolk, 2016), studies of plant
such as Brachypodium pinnatum and Polygonatum odoratum, which
dispersal (Brunet, 2007), analyses of the effects of habitat fragmen-
were classified as dry grassland (Festuco-­Brometea) or forest fringe
tation (Vanneste et al., 2019; Paal et al., 2020), range size and niche
(Trifolio-­Geranietea) species, respectively. Relying solely on such
breadth analyses (Michaelis et al., 2016), and conservation network
classification, these species would be excluded from a list of typical
planning (Culmsee et al., 2014). The wide range of possible applica-
forest plants. Contemporary phytosociology partly overcomes this
tions and the frequent use of this list, as well as the potential changes
issue by identifying such species as diagnostic for both forest and
in habitat affinity of many plant species, underpin that there is an
open-­land habitats (Mucina et al., 2016, Chytrý et al., 2020). Yet,
urgent need to expand the list towards larger biogeographical scales.
even this approach does not offer a direct way of classifying the
In this study, we aimed to:
affinity of plant species to forests as a whole. Second, the species
affinity to a given habitat may vary considerably at regional scales
1. Develop the first forest plant species list (EuForPlant) for tem-
due to differences in, for example, (micro-­)climate, land use and
perate and boreal Europe with a region-­
specific classification
land-­use history, topography, soil and biotic interactions. For exam-
of each species, based on large vegetation surveys and related
ple, Anemone nemorosa is characteristic of temperate broadleaf for-
vegetation databases, regional floras and expert knowledge;
ests, albeit under montane conditions, and in northern Europe it is
2. Analyse potential shifts in forest affinity of plant species between
also common in mountain meadows, roadside verges and grasslands
(Dierschke, 1997; Peppler-­Lisbach & Petersen, 2001; Chytrý, 2007).
the study regions;
3. Reflect on the applications and limitations of the list.
A potential explanation could be that this species finds more suitable
climatic conditions outside forests towards northern latitudes and
elevations (i.e. at the cold limit of its distribution range), whereas it
needs cooler and more humid growing conditions in the forest understorey of temperate and lowland forests (De Frenne et al., 2019;
Zellweger et al., 2020). Such changes in habitat affinity are difficult
2
|
M E TH O D S
2.1 | Reference area: countries and biogeographical
regions
to consider in the phytosociological system, underscoring the need
for a supra-­regional classification.
Forest regions for habitat affinity assessment of vascular plant spe-
Beyond phytosociology, a myriad of different approaches has
cies were delineated using a combination of country borders with:
been developed over the past decades to define the forest affinity
(a) the Natura 2000 Biogeographical Regions (status 2011, https://
of plant species and use this to assess forest biodiversity or inform
ec.europa.eu); and (b) country-­specific phytogeographical or geo-
conservation decision-­making (Peterken, 1974, 1977, Matlack, 1994,
graphical classifications of natural landscapes. This resulted in 24
Peterken, 1996, Hermy et al., 1999, Honnay et al., 1999, Peterken
regions from 13 countries, encompassing the nemoral, hemiboreal
& Francis, 1999). Such endeavours were, however, based on a lim-
and boreal vegetation zones as well as lowlands, uplands and high
ited number of single studies or local observations, limiting their ap-
mountains (Figure 1 and Appendix S1).
plicability at larger scales. By contrast, Schmidt et al. (2003, 2011)
developed the first nationwide and ecologically grounded reference
list of vascular plant species occurring in forests across Germany,
2.2 | Forest definition
based on expert knowledge as well as readily available resources of
plant diversity monitoring programmes and land-­cover data, thus
Prior to the species classification, we needed to define forest. We
avoiding misinterpretations related to traditional phytosociological
slightly modified the forest definition of the German forest species
classifications. In accordance with the principles of phytosociology,
lists (Schmidt et al., 2003, 2011) containing elements of the forest
the species listed in this supra-­regional forest species list were as-
definitions given by Mueller-­Dombois and Ellenberg (1974) and the
signed to different categories of forest affinity, which also allows the
German Federal Forest Inventory (http://www.bunde​swald​inven​tur.
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HEINKEN et al.
BOX 1 Forest definition
Forests are defined as tree-­covered areas with normally
at least 30% canopy cover (but including, for example,
pine forests on sand dunes and bogs, forests on dry rocky
slopes, pasture woods). In the case of completely closed
canopies, the area of a stand should be equal to at least a
circle with the radius of the maximal height of its tree layer.
In the case of an open canopy, the minimum area increases
proportionally with the decrease in overhead shading.
Forests in our definition include:
• Clear-­cuts or temporarily open areas such as forest gaps;
• Stands in the regeneration phase or with shoots from
stumps after coppicing;
• Forest edges, including edges of forest tracks.
Forests do not include:
• Scrub, i.e. stands in which the woody component is dominated by shrubs;
• Tree rows and hedgerows;
• Intensively managed parks with forest-­like structures;
• Paved forest tracks and forest roads;
• Permanently open areas within forests such as forest
aisles (e.g., for power lines, if not under coppice-­like
management) or forest division, fire break lines, grasslands, cultivated deer pastures and feeding places of
game animals;
• Orchards;
F I G U R E 1 Overview of the 24 forest regions for which
the European Forest Plant Species List (EuForPlant) has been
established: (1) France, Atlantic region; (2) France, continental
region; (3) France, mountains; (4) Belgium, lowlands; (5) Belgium,
uplands and mountains; (6) Netherlands; (7) Luxembourg; (8)
Germany, NW lowlands; (9) Germany, NE lowlands; (10) Germany,
uplands and mountains; (11) Germany, Alps; (12) Poland, lowlands;
(13) Poland, mountains; (14) Czech Republic, lowlands; (15) Czech
Republic, uplands and mountains; (16) Denmark, Atlantic region;
(17) Denmark, continental region; (18) Sweden, Nemoral zone;
(19) Norway, hemiboreal zone: Atlantic coastline; (20) Sweden,
hemiboreal zone; (21) Estonia; (22) Latvia; (23) Lithuania; (24)
Sweden, boreal zone (except alpine)
• Christmas tree and ornamental twig plantations;
• Short rotation coppices for bioenergy on former arable
land;
• Herbaceous fringe vegetation in the phytosociological sense (Galio-­Alliarietalia, Trifolio-­Geranietea) without
connection to forest stands.
1. All species classified by Ellenberg et al. (1992) as character
species of deciduous forests and related scrub (Alnetea glutinosae, Rhamno-­Prunetea, Quercetea robori-­petraeae, Querco-­
de). We emphasized the ecological context (e.g., stand structure and
Fagetea, Salicetea purpureae), coniferous forests and related
occurrence of forest species) and reduced the focus on legal defini-
heaths (Erico-­Pinetea, Pulsatillo-­Pinetea, Vaccinio-­Pinetea), as well
tions (Box 1).
as vegetation of perennial herbs and shrubs in the periphery
of forests (Betulo-­Adenostyletea, Epilobietea, Trifolio-­Geranietea);
2.3 | Reference list of species
2. Species with Ellenberg indicator values for light between 1 and 6
(“deep shade plant” to “between semi-­shade plant and semi-­light
plant”);
Based on the flora of Germany, Schmidt et al. (2002), Schmidt et al.
3. Species selected from an evaluation of forest vegetation surveys,
(2003) developed a regionalized list of potential forest vascular plant
mainly based on larger compilations or databases (for a detailed
taxa (mainly species but in some cases subspecies and a few aggre-
list see Appendix S1). All species with a relative frequency of at
gates; henceforth referred to as “species”) as a starting point for dis-
least 10% in at least one forest vegetation type (association or
cussion in an expert board. This list chiefly included:
unranked community) were considered.
HEINKEN et al.
Journal of Vegetation Science
| 5 of 16
This list was then sent to a board of 45 experts with a question-
information on rare species and regions where vegetation-­plot data
naire concerning the amendment and acceptance of the method-
were scarce was extracted from regional and national floras (see
ological approach, as well as the category definitions and tentative
Appendix S1 for a detailed list) supplemented with expert knowl-
assignment of the individual species to categories (Figure 2). Chairs
edge. Accordingly, the list was gradually expanded to include new
for vegetation science and the forest botany chairs at forest faculties
regions and species. In a few cases of striking differences in the
were asked to propose experts; further contacts (e.g. retired scien-
classification of single species between adjacent regions, the lead
tists) were appointed. The assignment was based chiefly on species
author (TH) made a further personal inquiry of the experts to clarify
frequency in vegetation surveys (forest communities according to
whether this resulted from an actual habitat shift or from misinter-
the forest definition in Box 1 vs open vegetation communities).
pretations of the basic definitions. Towards the end of this process,
Species were classified as forest species (category 1) when they have
the extended list was again sent to all experts for a final check (es-
more than double frequency in at least one forest community (asso-
pecially for classification of newly added species) and, if necessary,
ciation or community with a larger distribution area) compared with
correction of the classifications.
in any open vegetation communities. Rare species were classified
according to the best available knowledge (for details see Schmidt
et al., 2003, 2011).
2.4 | Taxonomy and additional species information
In an attempt to expand this list to other European regions, the
FLEUR network (www.fleur.ugent.be) of forest vegetation ecolo-
The species concept and nomenclature were based on the Euro+Med
gists, many of whom are trained originally in phytosociology, took
PlantBase (Euro+Med, 2006–­
). Exceptions are some groups of
the initiative to establish an international expert board across the 24
species (mainly composed of apomicts) that are difficult to distin-
European forest regions in Belgium, the Czech Republic, Denmark,
guish and/or lack a consistent taxonomy in floras across the coun-
Estonia, France, Germany, Latvia, Lithuania, Luxembourg, the
tries involved (Alchemilla spp., Callitriche palustris aggr., Hieracium
Netherlands, Norway, Poland and Sweden (Appendix S1). We re-
sect. Alpestria, Rubus fruticosus aggr.). By contrast, some clearly
stricted ourselves to the nemoral and boreal zones of Europe with
recognizable hybrids with specific ecology (Circaea × intermedia,
the exception of parts of the Alps, Eastern Europe and the Balkans
Polypodium × mantoniae, Populus × canadensis, Populus × canescens,
because we were lacking experts in these regions who belonged to
Rhododendron × intermedium, Salix × rubens, Spiraea × billardii) were
or were connected to FLEUR.
added to the list. In other cases, hybrids were included in one of the
The definitions and accompanying explanations (Box 1, Box 2;
parental species. Some subspecies were also distinguished if they
Figure 3; Appendix S2) together with the German forest species list
are treated as independent species in many floras and differ in their
from 2011 were then sent to all experts as a basis for the delineation
ecology from related subspecies (e.g., Aconitum lycoctonum subsp.
of regions and classification of species (Figure 2). In all 24 regions,
lasiostomum, subsp. lycoctonum and subsp. vulparia, Carex muricata
plant species were assigned to a category reflecting their affinity to
subsp. muricata and subsp. pairae, Dactylis glomerata subsp. glom-
forest habitats (see below) following the methodology of Schmidt
erata and subsp. lobata, Juniperus communis subsp. communis and
et al. (2011), that is, wherever possible based on frequency data of
subsp. nana, Lamium galeobdolon subsp. argentatum, subsp. flavidum,
species in plant communities in large vegetation surveys or related
subsp. galeobdolon, and subsp. montanum, Pinus mugo subsp. mugo
vegetation-­plot databases (for a detailed list see Appendix S1). In
and subsp. rotundata, Viscum album subsp. abietis, subsp. album and
the case of the Czech Republic, part of the species information was
subsp. austriacum). Angiosperm plant families follow the nomencla-
extracted from a national database of vascular plant species pools
ture of the APG IV system (Stevens, 2001), whereas gymnosperm
in different habitat types (Sádlo et al., 2007), and subsequently
and pteridophyte plant families are according to the Euro+Med
matched to our category definitions with some editing. Additional
PlantBase (Euro+Med, 2006–­).
F I G U R E 2 Workflow for the creation
of the European Forest Plant Species list
(EuForPlant)
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| Journal of Vegetation Science
HEINKEN et al.
F I G U R E 3 Explanation for species assignment to categories of forest species. Plant species may either occur in one habitat type
(1.1/1.2/O), or in different combinations of forest and open-­landscape habitats (2.1/2.2). If species restricted to forests grow in habitat
types 1.1 and 1.2, they belong to category 1.1. Category 1.1 includes species that can be found mainly in closed forests, whereas 1.2 species
occur predominantly along forest edges and in forest openings. Category 2.1 includes species that are part of at least one near-­natural (late-­
successional) forest type. The majority of 2.1 species probably have their primary occurrences in forests, surviving and regenerating in open
(cultural) landscape habitats, but also in remnant forests. Species are assigned to category 2.2 if they only enter (open) forests created by
afforestation or during succession of open vegetation (dwarf-­shrub heaths, calcareous and sand grasslands, wet meadows). All species that
do not meet the requirements for the four forest affinity categories are classified as “species of open landscape” (category O) (Box 2)
to vegetation layers:
2.5 | Classification of plant species according to
forest affinity
• Trees –­woody perennial plants, typically having a single stem
Following Schmidt et al. (2003, 2011), our forest species list distin-
or trunk contributing to the upper canopy layer(s) of a forest
guishes five categories of forest affinity referring to the forest defi-
(>7 m);
nition above. This region-­specific classification scheme was applied
We distinguished the following life forms and assigned species
• Epiphytic parasites –­plants that grow as hemiparasites in the
crown of trees;
to each species. The categories are nested within two larger groups,
i.e. specialist forest species and generalist forest species (Box 2).
• Lianas –­long-­stemmed, woody or rarely herbaceous vines that
are rooted in the soil and use trees or shrubs to climb up to the
canopy;
2.6 | Data analysis
• Shrubs –­medium-­sized perennial woody plants which are distinguished from trees by their multiple stems, shorter stature (≤7 m)
First, we calculated the number of native, regionally alien and alien
and regularly surmount the herb layer;
species for each life form (trees, epiphytic parasites, lianas, shrubs
• Herb-­layer species –­annual, biennial or perennial herbaceous
species as well as woody dwarf shrubs (chamaephytes).
and herb-­layer species) as well as for the different forest affinity
categories (1.1, 1.2, 2.1 and 2.2/O). Second, we computed the distribution of the forest affinity categories across the 24 study regions,
For all regions separately, we distinguished between native spe-
and determined the representation of plant families within each
cies including archaeophytes (i.e. species established before AD
category. Differences in the distribution of forest affinity catego-
1500) and alien species (i.e. neophytes, introduced after AD 1500),
ries among study regions were then tested using a Kruskal–­Wallis
based on regional and national floras or reference lists. Native spe-
non-­parametric analysis of variance (ANOVA) with the “kruskal.test”
cies and archaeophytes could not be clearly separated because the
function in the stats package, followed by Dunn–­Bonferroni post-­
status of archaeophytes is poorly known in some countries (see also
hoc tests using “dunnTest” function in the FSA package (version 0.8;
Wagner et al., 2017). We defined alien species that were native in at
R Core Team, R Foundation for Statistical Computing, Vienna, AT).
least one of the study regions as “regionally alien species” and spe-
Third, for all species, we calculated the probability of assignment to
cies that were not native in any of the study regions as “alien species”
each of the four forest affinity categories (1.1, 1.2, 2.1 or 2.2) or the
(mainly non-­European neophytes).
O category in the 24 study regions (e.g., if a species was assigned to
HEINKEN et al.
Journal of Vegetation Science
| 7 of 16
BOX 2 Categories of forest affinity
Group 1. Specialist forest species (species that occur only in forests)
• Category 1.1 –­Species that can be found mainly in the closed-­canopy forest;
• Category 1.2 –­Species that occur predominantly along forest edges and in forest openings, including: (a) species of forest edges
(both sunny and shaded, both outside and inside the forest); (b) species of windthrow, burned or clear-­cut areas; (c) species that
mainly occur on skid trails and unpaved forest paths; and (d) species that are restricted to open forests found under extreme site
conditions (shallow soils on slopes, rock outcrops, boulder and scree slopes, extremely wet forests).
Group 2. Generalist forest species (species that occur both in and outside forests)
• Category 2.1 –­Species that can be equally found in forest and open vegetation;
• Category 2.2 –­Species that can be found mainly in open vegetation, but also in forest.
The following explanations are important for the assignment of plant species to the four categories (Figure 3 and Appendix S2).
Group 1 vs 2. The species with the highest degree of forest affinity are classified into group 1, whereas those assigned to group 2 are
typical forest species that also occur regularly in one or several non-­forest, open-­landscape habitats. Some of the group 1 species
may also occur occasionally in woody habitats outside the forest such as hedgerows, coppices, mountain pine and green alder
scrub (krummholz). The abundance of species in forest vs open vegetation was considered irrelevant for assignment to the main
groups because this is highly dependent on the landscape configuration, the naturalness of forests in a region and other factors.
Category 1.1 vs 1.2. Many category 1.1 species may show a positive response to management, i.e. they flower and set fruit especially
on sites of 1.2 with less shady conditions, but are not included in this group if they regularly occur in closed-­canopy forests (e.g.,
Milium effusum). On the other hand, any species classified to category 1.2 may also occur in a closed-­canopy forest, but usually
only in small numbers and with lower vitality (often non-­flowering).
Category 2.1 vs 2.2.
Category 2.1 includes species that are part of at least one near-­natural (late-­successional) forest community. The majority of species
probably have their primary occurrences in forests, surviving and regenerating in open habitats of (cultural) landscape, but also
in remnant forests. Examples of this group include: several species of wet (spring) forests and wet grasslands such as Angelica sylvestris, Caltha palustris and Crepis paludosa; and several species of thermophytic forests and dry grasslands (e.g., Anthericum liliago,
Polygonatum odoratum, Sesleria caerulea). Even if there are very few remnant populations of these species in near-­natural forests
of a region, they were assigned to category 2.1.
Category 2.2 includes species that are not part of near-­natural (late-­successional) forest communities. Species are assigned to category 2.2 if they only enter (open) forests created by afforestation or during the succession of open vegetation (dwarf-­shrub
heaths, calcareous and sand grasslands, wet meadows). Also included are species of pastured forests, stands of alien trees (e.g.,
Robinia pseudoacacia) and severely disturbed forest stands such as many alluvial forests (Salicion albae). In general, most 2.2 species are characterized by high light requirements. The category considers that a significant part of European forest is recent, and
their flora still shows signs of former land uses.
Category O. All species that do not meet the requirements for the four forest species categories are classified as “species of open
landscape” (category O). These are only included in the forest species list if they are categorized as forest species in at least one
of the regions. Whether a species is classified as 2.1/2.2 or O depends on its relative abundance in certain types of forest vs open
vegetation, and not its general frequency in a region (see Group 1 vs 2). Accidental occurrences of a particular species in a forest are
thus not sufficient for assignment to categories 2.1 or 2.2. The same is true for the occurrence of open-­landscape species in forests.
category 1.1 in 23 regions and to 1.2 in one region, its probabilities of
statistical analyses were performed in R version 3.5.1 (R Core Team,
assignment to 1.1 and 1.2 were 96% and 4%, respectively). We also
R Foundation for Statistical Computing, Vienna, AT).
calculated the probability that each species was consistently assigned
to the same category across all study regions, or whether it shifted
to another category. For instance, if a species was assigned to 1.1
in five regions, to 1.2 in three regions and to 2.1 in two regions, the
probability of assignment to its main category 1.1 was 50%, whereas
3
|
R E S U LT S
3.1 | Overview
the probability of 1.1 → 1.2 or 1.1 → 2.1 shifts were 33% and 22%, respectively. Probability distributions of habitat consistency (i.e. assign-
EuForPlant comprises a total of 1,726 vascular plant species (Table 1),
ment to its main category across the study area) or shifts were then
most of them (1,621; 94%) native in at least some of the studied re-
computed across all 1,726 forest species, and visualized using the
gions. The majority of species (1,437; 83%) are part of the herb layer,
“geom_density” function in the ggplot2 package (Wickham, 2016). All
followed by tree and shrub species. Only a few species are lianas and
8 of 16
| Journal of Vegetation Science
HEINKEN et al.
epiphytic parasites. Life forms differ between native and alien forest
pattern was similar for all forest affinity categories except 1.2 spe-
species (Figure 4a); whereas nearly 90% of native and archaeophytic
cies, occurring significantly less frequently across the study regions
species are herb-­layer species, ca. 50% of the species classified as
than species of the other categories (median of 6 regions compared
alien in the whole study area are trees, lianas and shrubs.
with 17 for 1.1, 16 for 2.1 and 15 regions for 2.2/O species; Kruskal–­
Of the 1,726 species, 250 were exclusively (or in more than 50%
Wallis test, χ² = 79.6, df = 4, p < 0.001, with Dunn–­Bonferroni
of the regions) classified as species of closed-­canopy forest (category
post-­hoc tests; Figure 5). The number of forest species varied sub-
1.1; e.g., Elymus caninus, Fagus sylvatica, Gymnocarpium dryopteris,
stantially between regions; the highest number was encountered
Oxalis acetosella; see Appendix S3 for a complete list). Forest edge,
in the French mountain areas (1,220 species), whereas the lowest
open forest and clear-­cut species of category 1.2 were the smallest
number was found in the Netherlands (442). Focusing exclusively on
group with 137 species (e.g., Cardamine flexuosa, Digitalis purpurea,
specialist forest species (1.1 and 1.2), the most species-­rich region
Humulus lupulus). Note that many Trifolio-­Geranietea species such as
was the German uplands and mountains (300 species), whereas the
Peucedanum oreoselinum and Polygonatum odoratum are also found in
boreal zone of Sweden (103) appeared to be the most species-­poor
other habitats than forest edges, and are therefore usually not classi-
region (see also Appendix S4).
fied as category 1.2. Much larger than the groups of specialist forest
The majority of forest species were dicots (1,261) and monocots
species were the generalist forest and open-­landscape groups, with
(379), whereas gymnosperms (16) and pteridophytes (70) were of
450 species classified as 2.1 (e.g., Caltha palustris, Deschampsia cespi-
minor importance. Among the 117 plant families represented in the
tosa, Pteridium aquilinum, Quercus robur) and 777 species classified as
list, the largest were the Asteraceae (194 species), Rosaceae (144)
2.2/O (e.g., Aesculus hippocastanum, Festuca rubra, Silene flos-­cuculi).
and Poaceae (115). The 23 plant families with at least 20 forest spe-
Many typical widespread species that occur both in closed-­canopy
cies are listed in Table 2. There were remarkable differences in family
forest and in open landscape were shrubs and trees because they
representation between the forest species groups. Species only oc-
rejuvenate and establish themselves in open vegetation outside the
curring in closed-­canopy forests (1.1) were comparatively rare among
forest (Appendix S3).
Apiaceae, Asteraceae, Fabaceae and Rosaceae, whereas no Salicaceae
Prevailing habitat preference differed between native and
were part of this group. For example, families with disproportionally
regionally alien species, on the one hand, and other (mostly non-­
high species numbers among category 1.1 were Orchidaceae (e.g.,
European) alien species, on the other hand (Figure 4b). Whereas ca.
Cephalanthera spp., Epipactis helleborine, Goodyera repens, Neottia
15% of native and regionally alien species were almost exclusively
nidus-­avis), Juncaceae (Luzula spp.) and Violaceae (Viola spp.). This was
species of closed-­canopy forest (category 1.1), this was true only for
also the case for pteridophytes (e.g., Athyrium filix-­femina, Lycopodium
5% of the other alien species. The latter were mainly comprised of
annotinum, Phegopteris connectilis) and some spermatophyte families
the group of open-­landscape species (groups 2.2 and O).
such as Pinaceae (e.g., Abies alba, Larix decidua, Pseudotsuga menziesii)
Many vascular plant species (260) occurred in all 24 studied re-
and Papaveraceae (e.g., Corydalis spp). Conversely, only two 1.1 spe-
gions (e.g., Adoxa moschatellina, Carex elongata, Athyrium filix-­femina,
cies were present among Asteraceae (Aposeris foetida and Prenanthes
Ribes nigrum, Ulmus glabra; see Figure 5). Yet, many other species had
purpurea) and only one among Fabaceae (Lathyrus vernus). Apiaceae
limited distribution and occurred in only one or a few regions. This
(e.g., Chaerophyllum temulum, Torilis japonica) and Fabaceae (e.g.,
TA B L E 1 Overview of the total number of species and relative
frequency in the European Forest Species List (EuForPlant),
grouped by their native status and life form
egory 1.2. Among generalist forest species (2.1 and 2.2), Asteraceae
Number of
species
Percentage
1,726
100
1,339
77.6
Regionally alien species
282
16.3
Alien species (across all
study regions)
105
6.1
All forest species
Nativeness
Native species (incl.
archaeophytes)
107
6.2
Epiphytic parasites
4
0.2
Lianas
19
1.1
Shrubs
159
9.2
1,437
83.3
Herb-­layer species
(e.g., Centaurea scabiosa, Cirsium oleraceum, Hieracium murorum,
Tussilago farfara) outnumbered the other families.
3.2 | Habitat shifts
Despite the relatively low probability of shifting categories between
regions (on average there was a 17.5% ± 4.2% SE probability of an
among-­region shift compared with a 75.6% ± 6.8% SE probability of
no shift; Figure 6), only about one-­third of all species (594) had a constant habitat preference, which implies that they were consistently
assigned to the same category across all the study regions. Species
Life form
Trees
Lathyrus niger, Vicia sylvatica) were especially well represented in cat-
with a constant habitat preference mainly belonged to categories
1.1 and 2.1. The majority of species, however, showed a habitat
shift throughout their distribution range. The most common shifts
were those in which species swapped between two categories (776
species) or species either shifted from one habitat to another (e.g.,
1.1→1.2; 30 species) or occupied an additional habitat (e.g., 1.1→2.1;
HEINKEN et al.
Journal of Vegetation Science
| 9 of 16
regions. Among these, a habitat shift 2.1 → 2.2 → O (179 species) was
especially well represented, implying a different ability of those species to grow in forests with different degrees of canopy openness.
4
|
DISCUSSION
4.1 | A new approach to defining the forest affinity
of European plant species
EuForPlant comprises nearly 2,000 vascular plant species in 24
different regions, covering a large part of Europe's temperate and
boreal forest flora. The species were categorized according to their
affinity to forests, and this classification was applied by region. The
list goes significantly beyond existing local and supra-­regional lists
of forest species in providing an expert-­based approach using a
standard protocol. Where possible, vegetation databases were used
to assess the linkage of plant species to forests across large geographical scales.
EuForPlant encompasses a variety of life forms (chiefly understorey herb, shrub and tree species, but also lianas and epiphytic
parasites such as Viscum spp.), and covers 117 plant families.
Among all vascular plant species on this list, 386 were alien in at
least one of the study regions, with phanerophytes (trees, shrubs
F I G U R E 4 Proportion of different (a) life forms and (b) forest
affinity categories (1.1, species of closed-­canopy forests; 1.2,
species of forest edges and forest clearings; 2.1, species of forests
and open vegetation; 2.2/O, species of mainly open vegetation
(but sporadically occurring in forests) of the 1,726 forest species in
EuForPlant among the 1,339 native species (incl. archaeophytes),
282 regionally alien species and other 105 (mostly non-­European)
alien species. For each species, the most common category across
the 24 study regions was considered. If two or more categories
were equally frequent, the species was ranked as “no preference”.
When summarizing across all regions, we did not distinguish among
2.2 and O species because these are the only categories including
species that are rarely found in forests. Moreover, species classified
as O necessarily have a 2.2 habitat preference in at least one
region, otherwise they would not be part of this forest species list
and lianas) being the most species-­rich life form among aliens.
These numbers are in accordance with Wagner et al. (2017) for
woodlands across the entire European continent, albeit the representation of alien species in terms of relative frequency was
three times higher (23.7% of all species) in our data set compared
with Wagner et al. (2017) (7%). Interestingly, some plant families
were either over-­ or underrepresented in our forest species list
compared with the whole flora of the regions of the European
Forest Plant Species List. To illustrate, in Germany, in the centre
of the study area, plant families that were overrepresented on the
list, thus having strong affinity to forests, were: Campanulaceae,
Caprifoliaceae, Ericaceae, Orchidaceae, Ranunculaceae, Rosaceae,
Rubiaceae, Salicaceae and Violaceae. Of these families, Ericaceae
and Orchidaceae were particularly well represented among the 1.1
72 species). A remarkably common habitat shift was from 2.2 to O
species. This is not surprising given that many pyroloids (Ericaceae,
(451 species); in some regions, the species also occurred in open,
Pyroleae tribe) and orchids from temperate and boreal forests
early-­successional forests, whereas in other regions, they were re-
associate with fungi that form ectomycorrhizas on surrounding
stricted to open landscape (e.g., Artemisia campestris, Carex lasio-
trees (Bidartondo et al., 2004; Tedersoo et al., 2007). One of
carpa, Galium verum, Linum catharticum, Sanguisorba minor; Appendix
the ecological attributes of such plants with myco-­h eterotrophy
S3). Another common shift was from 2.1 to 2.2 (108 species); in some
(e.g., Corallorhiza trifida or Neottia nidus-­avis) or mixotrophy (e.g.,
regions, the species also occurred in (semi-­)natural, late-­successional
Cephalanthera spp. or Epipactis spp.) is that they are independent
forests, whereas in other regions, they were restricted to early-­
of solar radiation and are able to colonize deeply shaded forest
successional forests (e.g., Agrostis capillaris, Brachypodium pinnatum,
habitats (Zimmer et al., 2007). Moreover, many Ericaceae are
Carex nigra, C. pseudocyperus, Potentilla erecta; Appendix S3).
known to thrive on nutrient-­p oor soils with acid humus (mor), and
Finally, 1.1 → 2.1 shifts also occurred; although species were re-
are often a prominent feature of the vegetation in heathlands, bogs
stricted to closed forests in some regions, they also occurred in open
and moors, but also in forests (Schwery et al., 2015; Olleck et al.,
landscape in others (e.g., Anemone nemorosa, Convallaria majalis,
2020). Some other species-­r ich families were underrepresented or
Dryopteris spp., Vaccinium myrtillus). A considerable number of forest
hardly present among forest species: Amaranthaceae, Asteraceae,
species (356) were assigned to three or more categories over the 24
Brassicaceae, Caryophyllaceae Gentianaceae and Plantaginaceae.
10 of 16
| Journal of Vegetation Science
HEINKEN et al.
F I G U R E 5 Species range size distribution of the 1,726 vascular plant species over the 24 forest regions in temperate and boreal Europe.
The x-­axis indicates the number of regions that are occupied by the respective species. Forest species are divided according to the forest
affinity categories (1.1, species of closed-­canopy forests; 1.2, species of forest edges and forest clearings; 2.1, species of forests and open
vegetation; 2.2/O, species of mainly open vegetation, but sporadically in forests). For each species, the most common category across the 24
study regions was considered. If two or more categories were equally frequent, the species was ranked as “no preference”. See Figure 4 for
the reason of merging the categories 2.2 and O. Moreover, species classified as O necessarily have a 2.2 habitat preference in at least one
region, otherwise they would not be part of this forest species list
Amaranthaceae, for instance, are known to prefer warm, open
2013). This phenomenon is partly referred to the “law of relative site
habitats such as temperate grasslands, sand dunes and agricul-
constancy” (Walter & Walter, 1953), implying that a species’ indi-
tural habitats (Kadereit et al., 2003), and were therefore nearly
vidual physiological requirements can be met in different types of
absent from this list of forest species. However, some of these
vegetation under varying climatic conditions. Similar microclimatic
families, such as Asteraceae, Brassicaceae and Gentianaceae, are
conditions can, for instance, be found in highly different ambient
more strongly represented in forest vegetation in the southeast-
climates owing to the complex interplay between macroclimate and
ern part of Europe (Večeřa et al., 2021) that is not covered by the
vegetation (Lenoir, 2020). As a result, the observed shift of several
forest species list, probably reflecting the evolutionary history of
typical forest plant species (e.g., Anemone nemorosa and Convallaria
the families.
majalis) from 1.1 to 2.1 towards higher latitudes, mountainous regions and coastal areas may arise from the fact that local climatic
4.2 | Importance of the regionally specific
classification system
conditions outside forests in these northern, high-­
elevation and
Atlantic regions tend to be similar to microclimatic conditions below
the forest canopy in temperate, lowland and continental regions (De
Frenne et al., 2019; Lenoir, 2020). Occasionally, however, the assign-
Despite the relatively low proportion of between-­
region habitat
ment of species to forest affinity categories may simply depend on
shifts, a remarkable number of species (nearly 75%) swapped cat-
the occurrence of certain forest habitats in the respective regions. In
egories in at least one of the study regions. The most common shifts
sum, these habitat shifts illustrate that the affinity of vascular plant
were from category 2.1 to 2.2 or from 2.2 to O, although a shift
species to forests strongly depends on region. They underpin the
from 1.1 to 2.1 was also detected in 12% of category 1.1 species.
importance of applying the classification at a regional level across
Thus, although species may be restricted to early-­successional for-
broad spatial scales.
ests, afforestations or even to open vegetation in some regions, they
may also occur in (semi-­)natural, late-­successional forests in others. Although some bias due to differences in judgement between
experts cannot be ruled out, such changes in a species’ ecological
4.3 | Expected impact, limitations and
ways forward
niche over broad geographical areas can most likely be attributed
to large-­scale gradients in climatic conditions, land use and deposi-
The current list was made publicly available on figshare in 2019
tion of airborne pollutants, as well as local variation in topography,
(Heinken et al., 2019, https://doi.org/10.6084/m9.figsh​are.80952​
microclimate and biotic interactions (see, for example, Wasof et al.,
17.v1), and has since been applied in several scientific publications,
HEINKEN et al.
Journal of Vegetation Science
| 11 of 16
TA B L E 2 Family representation among vascular plant species in the European Forest Plant Species List (EuForPlant)
Occurrence mainly in category
All species
1.1
1.2
2.1
2.2/O
No preference
Families
No.
%
No.
%
No.
%
No.
%
No.
%
No.
%
Asteraceae
194
11.2
2
0.8
11
8
44
9.8
126
15.5
11
14.5
Rosaceae
144
8.3
5
2
20
14.6
62
13.8
50
6.2
7
9.2
Poaceae
115
6.7
23
9.2
0
0
29
6.4
59
7.3
4
5.3
Fabaceae
88
5.1
1
0.4
13
9.5
13
2.9
59
7.3
2
2.6
Cyperaceae
97
5.6
12
4.8
3
2.2
25
5.6
52
6.4
5
6.6
Ranunculaceae
65
3.8
14
5.6
7
5.1
23
5.1
20
2.5
1
1.3
Apiaceae
61
3.5
3
1.2
8
5.8
17
3.8
30
3.7
3
3.9
Lamiaceae
56
3.2
7
2.8
3
2.2
13
2.9
31
3.8
2
2.6
Orchidaceae
53
3.1
23
9.2
5
3.6
11
2.4
13
1.6
1
1.3
Caryophyllaceae
49
2.8
7
2.8
5
3.6
9
2
25
3.1
3
3.9
Ericaceae
39
2.3
9
3.6
0
0
12
2.7
16
2
2
2.6
Brassicaceae
35
2.0
7
2.8
4
2.9
7
1.6
17
2.1
0
0
Salicaceae
33
1.9
0
0
0
0
11
2.4
21
2.6
1
1.3
Orobanchaceae
32
1.9
5
2
5
3.6
4
0.9
13
1.6
5
6.6
Boraginaceae
29
1.7
7
2.8
5
3.6
7
1.6
8
1
2
2.6
Caprifoliaceae
29
1.7
4
1.6
7
5.1
8
1.8
10
1.2
0
0
Asparagaceae
27
1.6
7
2.8
0
0
8
1.8
8
1
4
5.3
Juncaceae
27
1.6
7
2.8
1
0.7
5
1.1
14
1.7
0
0
Plantaginaceae
27
1.6
2
0.8
2
1.5
4
0.9
19
2.3
0
0
Rubiaceae
27
1.6
7
2.8
1
0.7
5
1.1
13
1.6
1
1.3
Campanulaceae
24
1.4
3
1.2
2
1.5
3
0.7
14
1.7
2
2.6
Violaceae
22
1.3
7
2.8
0
0
8
1.8
6
0.7
1
1.3
Primulaceae
20
1.2
6
2.4
1
0.7
7
1.6
5
0.6
1
1.3
Other 94 families
433
25.1
82
32.8
34
24.8
115
25.6
184
22.6
18
23.7
Note: Shown are the total number of species and relative frequency for the 23 families with at least 20 species, grouped by forest affinity categories
(1.1, species of closed-­c anopy forests; 1.2, species of forest edges and forest clearings; 2.1, species of forests and open vegetation; 2.2/O, species of
mainly open vegetation, but sporadically in forests). For each species, the most common category across the 24 study regions was considered. If two
or more categories were equally frequent, the species was ranked as “no preference”.
e.g., to assess the variation in plant species diversity and composition
relative frequency) in European forests to various environmental
along edge-­to-­interior gradients in forests across Europe (Govaert
variables such as macroclimatic temperature and precipitation
et al., 2020), to study the migration of forest herbs along hedgerow
(e.g., from CHELSA; Karger et al., 2017), atmospheric nitrogen
corridors (Vanneste et al., 2020), to assess functional diversity of
(N) deposition and acidification (e.g., from EMEP; http://www.
understorey plant communities in fragmented landscapes (Vanneste
emep.int) and land-­use changes (e.g., from historical landscape
et al., 2019), to experimentally assess the effects of both macro-­and
analyses);
microclimatic changes on understorey plants with contrasting de-
2. To unravel how forest structure and microclimate gradients in-
gree of forest affinity (De Pauw et al., 2021), to model their popula-
fluence understorey plant community dynamics; for example,
tion growth rates under a changing climate system (Sanczuk et al.,
by assessing the relationship between temporal shifts in the dis-
2021), and to characterize the seed bank in forests along various
tribution of forest affinity categories in European forests (e.g.,
environmental gradients (Gasperini et al., 2021). Other potential ap-
using resurvey databases such as forestREplot; https://fores​trepl​
plications of this list are:
ot.ugent.be) to changes in forest canopy cover and subcanopy
temperatures over time;
1. To study the effects of global environmental change (climate
3. To study the ecology of forest plants; for example, by compar-
change, land-­
use change, atmospheric pollution, biological in-
ing Ellenberg indicator and thermal tolerance values among the
vasions, etc.) on forest biodiversity; for example, by linking
forest affinity categories. For instance, it is expected that spe-
the distribution of forest affinity categories (e.g., in terms of
cies of closed forests (1.1) would have lower Ellenberg indicators
12 of 16
| Journal of Vegetation Science
HEINKEN et al.
F I G U R E 6 Distribution of probability
across the 1,726 forest species in
EuForPlant that a species (a) stays
within the same category (graphs on the
diagonal) or (b) shifts to another category
in the 24 study regions. Categories
of forest affinity are: 1.1, species of
closed-­canopy forests; 1.2, species of
forest edges and forest clearings; 2.1,
species of forests and open vegetation;
2.2, species of mainly open vegetation,
but sporadically also occurring in
forests. All species that do not meet the
requirements for the four forest affinity
categories are classified as O species (Box
2). For each species, the most common
category across the 24 study regions was
considered. If two or more categories
were equally frequent, the species was
ranked as “no preference”
for light, nitrogen and reaction than species of open vegetation
species only in some regions (e.g., the Czech Republic, Germany
(Dzwonko, 2002; Schmidt et al., 2014). In addition, the list can be
and the Netherlands), whereas expert knowledge was a central
used to explore the evolutionary ecology of forest plant species;
methodology of classification in other regions. This is because in
4. Trait-­based analyses; for example, by establishing the link be-
the European Vegetation Archive (Chytrý et al., 2016) or other da-
tween forest affinity categories and functional life-­history traits
tabases: (1) some of the included countries, especially in northern
that are related to their competitiveness or dispersal ability. For
Europe, are not adequately covered; (2) the structural information
instance, Vanneste et al. (2019) demonstrated that the 1.1 species
needed to assign species to categories is often not available in the
were significantly shorter, produced heavier seeds and displayed
databases; and (3) many rare species are not sufficiently represented
lower specific leaf area than species in the other categories;
to classify them using fidelity measures. Following expert-­
based
5. Use in conservation planning and decision-­making; for example,
evaluation and correction of errors due to different interpretations
to evaluate biodiversity status of forests and to identify most
of the definitions by experts, the classification of forest species has
valuable, least-­transformed forests remaining in Europe. For in-
mainly been based on expert knowledge rather than an objective
stance, Culmsee et al. (2014) previously used the forest plant
mathematical algorithm. Expert assessments have been demon-
species list developed by Schmidt et al. (2011) to determine
strated to be invaluable and are still gaining momentum in ecolog-
the distribution of high-­nature-­value forests in Lower Saxony,
ical applications and conservation decision-­making (Kuhnert et al.,
Germany. In turn, this approach contributed to the identifica-
2010). Skilled experts have acquired extensive knowledge and ex-
tion of new conservation areas and to monitor the success of
perience, allowing them to perform a multicriteria analysis of the
existing forest conservation measures within the framework of
most relevant information for a given context in order to find the
Natura 2000.
most appropriate solution. However, groups of experts are not free
from biases and blind spots, and the usefulness of such expert judge-
Despite the extensiveness of the data set and its potential suit-
ments fluctuates (Burgman, 2005). For this reason, we looked for
ability for ecological research, several shortcomings must be con-
ways to independently support the expert assessments of the for-
sidered when using the EuForPlant. Most importantly, analyses of
est affinity categories using clear definitions and, as far as possible,
vegetation databases or comprehensive vegetation surveys based
vegetation databases. Moreover, the classification was coordinated
on databases were the primary source of the classification of forest
by the FLEUR network, a group of scientists who have met annually
HEINKEN et al.
for over 15 years and have local knowledge of the broad range of
Journal of Vegetation Science
| 13 of 16
species classification. Günter Gottschlich provided the taxonomy of
forest communities and habitat types covered in this list. Another
the Hieracium and Pilosella taxa and a basis for which species could
potential limitation related to this list is that intra-­regional shifts in
be included in EuForPlant. Technical support for creating the GIS
forest affinity are not considered. Even within regions, large climatic
maps was provided by Katja Lorenz and Jiří Šmída.
variability may be encountered, for instance, in mountainous areas
where gradients from lowland conditions to montane or (sub-­)alpine
DATA AVA I L A B I L I T Y S TAT E M E N T
climate occur. Finally, to date, our list covers only part of Europe,
The complete database and accompanying metadata are avail-
albeit a significant one, but not Southern and Eastern Europe. Filling
able from the Figshare repository, with the identifier https://doi.
in data from these areas will be an important task for the future.
org/10.6084/m9.figsh​are.80952​17.v1 (Heinken et al., 2019).
5
|
CO N C LU S I O N
ORCID
https://orcid.org/0000-0002-1681-5971
Thilo Heinken
https://orcid.org/0000-0001-8482-0679
Martin Diekmann
EuForPlant is a unique database of forest vascular plant species
Jaan Liira
covering the European temperate and boreal biome with a region-­
Anna Orczewska
https://orcid.org/0000-0001-8863-0098
https://orcid.org/0000-0002-7924-9794
specific classification scheme of their forest affinity. The list has
Marcus Schmidt
https://orcid.org/0000-0001-6712-6861
many possible applications in vegetation science, and is equally in-
Jörg Brunet
https://orcid.org/0000-0003-2667-4575
strumental in forest monitoring and conservation decision-­making.
Milan Chytrý
https://orcid.org/0000-0002-8122-3075
EuForPlant can be readily linked to other large vegetation-­
plot
Olivier Chabrerie
databases such as EVA (Chytrý et al., 2016) and sPlot (Bruelheide
Guillaume Decocq
https://orcid.org/0000-0001-9262-5873
et al., 2019; Sabatini et al., 2021), to plant trait data sets such as
Pieter De Frenne
https://orcid.org/0000-0002-8613-0943
https://orcid.org/0000-0003-0802-3509
https://orcid.org/0000-0002-8949-1859
TRY (Kattge et al., 2020) and ClimPlant (Vangansbeke et al., 2021),
Pavel Dřevojan
as well as to climate databases such as CHELSA (Karger et al., 2017)
Zbigniew Dzwonko
and SoilTemp (Lembrechts et al., 2020). The links between these da-
Jörg Ewald
tabases can help answer questions related to forest plant diversity
Bente Jessen Graae
https://orcid.org/0000-0002-5568-4759
patterns, structure and dynamics of understorey plant communities,
John-­Arvid Grytnes
https://orcid.org/0000-0002-6365-9676
long-­term vegetation changes in forests in response to global envi-
Martin Hermy
https://orcid.org/0000-0003-4944-0341
https://orcid.org/0000-0002-2758-9324
https://orcid.org/0000-0002-5403-0139
ronmental change, plant species niche breadth and range sizes, plant
Jonathan Lenoir
dispersal and colonization, and many others. Users of EuForPlant
Sigrid Lindmo
https://orcid.org/0000-0003-0638-9582
https://orcid.org/0000-0002-0077-2911
need to be aware of its limitations and potential biases that might
Damien Marage
affect answers to specific research questions. Finally, we see this list
Vitas Marozas
https://orcid.org/0000-0002-1311-7000
https://orcid.org/0000-0001-5687-5218
https://orcid.org/0000-0001-5118-3941
as dynamic and underpin the importance of expanding it towards
Thomas Niemeyer
other regions and biomes (e.g., the Mediterranean and the Balkans)
Jaanus Paal
to create the most comprehensive compilation of forest plant spe-
Petr Pyšek
https://orcid.org/0000-0001-8500-442X
cies at a pan-­European scale. The latest version of EuForPlant will
Jiří Sádlo
https://orcid.org/0000-0001-9723-3334
always be available on igshare (https://doi.org/10.6084/m9.figsh​
Joop H.J. Schaminée
are.80952​17.v1).
Torbjörn Tyler
Kris Verheyen
https://orcid.org/0000-0002-2067-9108
AU T H O R C O N T R I B U T I O N S
Monika Wulf
https://orcid.org/0000-0001-6499-0750
Thilo Heinken conceived and designed the study in close collabora-
Thomas Vanneste
https://orcid.org/0000-0003-0499-5757
https://orcid.org/0000-0002-0416-3742
https://orcid.org/0000-0002-7886-7603
https://orcid.org/0000-0001-5296-917X
tion with the FLEUR network and regional collaborators. All authors
were involved in the region delineation and species classification.
Thomas Vanneste and Thilo Heinken conducted the statistical analyses. Thilo Heinken and Thomas Vanneste wrote the first draft of the
paper, while all authors contributed to revisions.
AC K N OW L E D G E M E N T S
Special thanks go to Bernard Amiaud†, Gaëtan Duponchelle,
Frédéric Dupont, Bruno de Foucault, Dieter Frank, Jean-­Christophe
Hauguel, Peter Horchler, Karsten Horn, Arnault Lalanne, Jean-­Paul
Legrand, Antoine Meirland, Jean-­Charles Millouet, Frank Richter,
Christian Ries, Michael Ristow, Christiane Ritz, Maria-­Sofie Rohner,
Marc Wagner and Aymeric Watterlot for their assistance in the
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S U P P O R T I N G I N FO R M AT I O N
Additional supporting information may be found in the online
HEINKEN et al.
main information sources used for the delimitation of natural regions
and assignment of species
Appendix S2. Explanatory materials for forest definitions and forest
affinity categories as an assistance for the regional contributors
Appendix S3. Widespread plant species of different forest species
categories
Appendix S4. Species numbers of forest species groups in
the European Forest Plant Species List across the 24 regions
in temperate and boreal Europe, sorted by the total number of
species
How to cite this article: Heinken, T., Diekmann, M., Liira, J.,
Orczewska, A., Schmidt, M., Brunet, J., et al (2022) The
version of the article at the publisher’s website.
European forest plant species list (EuForPlant): Concept and
Appendix S1. The 24 regions for which the EuForPlant was
Available from: https://doi.org/10.1111/jvs.13132
established, the responsible coordinators and contributors, and the
applications. Journal of Vegetation Science, 33, e13132.
Supporting information to the paper
Heinken, T. et al. The European Forest Plant Species List (EuForPlant): Concept and applications. Journal of Vegetation Science.
Appendix S1. The 24 regions for which the EuForPlant was established, the responsible coordinators and contributors, and the main information sources used to delimitate natural regions and assignment of species. Colours:
assignment to biogeographic regions of EU Natura 2000 network. Light green: nemoral zone, atlantic; beige: nemoral zone, continental; dark green: nemoral zone, high mountains; light blue: hemiboreal zone; dark blue: boreal
zone. For abbreviated names of coordinators and primary contributors, we refer to the author list of this article. Full names of secondary contributors with particular expert knowledge on certain species groups and/or regions:
AL Arnault Lalanne, AME Antoine Meirland, AW Aymeric Watterlot, BA Bernard Amiaud†, BDF Bruno de Foucault, CRS Christian Ries, CRZ Christiane Ritz, DF Dieter Frank, FD Frédéric Dupont, FR Frank Richter, GDU Gaëtan
Duponchelle, JCH Jean-Christophe Hauguel, JCM Jean-Charles Millouet, JPL Jean-Paul Legrand, KH Karsten Horn, MR Michael Ristow, MSR Maria-Sofie Rohner, MW Marc Wagner. PH Peter Horchler.
Nr.
Region
Coordinator
Primary
contributor(s)
Secondary
contributor(s)
Delimitation of natural
landscapes
(additional sources)
Flora of natural regions,
identification of alien species
Assignment of species
Assignment of species according to
according to vegetation surveys floras
and databases
1
France, atlantic region
OC
GD, MH, DM
AL, AME, AW, BA,
BDF, GDU, FD,
JCH, JCM, JPL
http://www.tela-botanica.org/,
TAXREF v.11
http://www.tela-botanica.org/
Rameau et al. 1989, Provost 1998,
Lambinon et al. 2004, Tison & de
Foucault 2014
2
France, continental region
OC
GD, MH, DM
BA, BDF, GDU,
JCM
http://www.tela-botanica.org/,
TAXREF v.11
http://www.tela-botanica.org/
Rameau et al. 1989, Tison & de
Foucault 2014
3
France, mountains
OC
GD, MH, DM
BA, BDF, GDU
Aeschimann et al. 2004,
http://www.tela-botanica.org/,
TAXREF v.11
http://www.tela-botanica.org/
Rameau et al. 1994, Saule 2002,
Tison & de Foucault 2014
4
Belgium, lowlands
MH
https://www.vlaanderen.be/inbo/d
atasets/florabank/,
http://biodiversite.wallonie.be/fr/atl
as-enligne.html?IDD=6056&IDC=807
Van Landuyt et al. 2006,
https://www.vlaanderen.be/inbo/data
sets/florabank/,
http://biodiversite.wallonie.be/fr/atlas
-en-ligne.html?IDD=6056&IDC=807
5
Belgium, uplands and
mountains
MH
http://biodiversite.wallonie.be/fr/atl
as-enligne.html?IDD=6056&IDC=807
Lambinon et al. 2004,
http://biodiversite.wallonie.be/fr/atlas
-en-ligne.html?IDD=6056&IDC=807
6
Netherlands
JS
7
Luxembourg
TN
8
Germany, NW lowlands
TH
MS, WUK
PH, KH
9
Germany, NE lowlands
TH
MS, WUK
10
Germany, uplands and
mountains
TH
11
Germany, Alps
12
Poland, lowlands
Rameau et al. 1994
MH
https://www.verspreidingsatlas.nl/
vaatplanten
Schaminée’s series
Colling 2005, https://neobiota.lu/
Niemeyer et al. 2010
Ssymank (1994)
NetPhytD & BfN 2013,
https://www.floraweb.de
Dierschke`s series, PreisingVahles’s series
Müller et al. 2016a,b
BS, CRZ, DF, FR,
KH, MR, MSR, PH
Ssymank (1994)
NetPhytD & BfN 2013,
https://www.floraweb.de
Dierschke’s series, Passarge
1964, Passarge & Hofmann
1964, Hilbig-Schubert’s
series, Berg & Dengler 2001,
2004
Müller et al. 2016a,b
MS, WUK
PH, KH, FR
Ssymank (1994)
NetPhytD & BfN 2013,
https://www.floraweb.de
Dierschke’s series, HilbigSchubert’s series,
Oberdorfer’s series, PresingVahle’s series
Müller et al. 2016a,b
TH
JE
KH
NetPhytD & BfN 2013,
https://www.floraweb.de
Dierschke’s series, Ewald
1995, 1997, 2012
Müller et al. 2016a,b
AO
ZD
CRZ
Matuszkiewicz 2017
Szafer et al. 1988,
Rutkowski 2019, Popek 2002,
Zieliński 2004
CRS, MW
Szafer & Zarzycki 1977 Mirek et al. 2002, Tokarska-Guzik
et al. 2012
https://www.floravannederland.nl/
13
Poland, mountains
AO
ZD
Szafer & Zarzycki 1977 Mirek et al. 2002, Tokarska-Guzik
et al. 2012
Matuszkiewicz 2017
14
Czech Republic, lowlands
MC
JS, PD, PP
Kaplan (2012):
Thermophyticum
Danihelka et al. 2012, Pyšek et al. Chytrý’s series, Sádlo
2012
et al. 2007
15
Czech Republic, uplands and
mountains
MC
JS, PD, PP
Kaplan (2012):
Mesophyticum &
Oreophyticum
Danihelka et al. 2012, Pyšek et al. Chytrý’s series, Sádlo
2012
et al. 2007
16
Denmark, atlantic region
BG
JF
Mossberg & Stenberg 2003,
Frederiksen et al. 2012
Dierßen 1996
Mossberg & Stenberg 2003,
Frederiksen et al. 2012
17
Denmark, continental region
BG
JF
Mossberg & Stenberg 2003,
Frederiksen et al. 2012
Dierßen 1996
Mossberg & Stenberg 2003,
Frederiksen et al. 2012
18
Sweden, nemoral zone
JB
TT
Moen (1999)
Mossberg & Stenberg 2003, Tyler
et al. 2015
https://www.dyntaxa.se/
800 forest plots in Skåne (JB), Mossberg & Stenberg 2003, Fröberg
2006, Georgson et al. 1997, Tyler et
Tyler et al. 2020
al. 2007, 2020
19
Norway, hemiboreal zone:
atlantic coastline
BG
JAG, SL
Moen (1999)
Dierßen 1996
Lid & Lid, D 2005, Mossberg &
Stenberg (2007),
https://artskart.artsdatabanken.no,
https://www.biodiversity.no/alienspecies-2018
Lid & Lid, D 2005, Mossberg &
Stenberg 2007
20
Sweden, hemiboreal zone
JB
MD, TT
Moen (1999)
Mossberg & Stenberg 2003, Tyler
et al. 2015 on the basis of
Karlsson 1998-2014,
https://www.dyntaxa.se/
Mossberg & Stenberg 2003, Fröberg
2006, Georgson et al. 1997, Sterner
1938, , Tyler et al. 2020
21
Estonia
JL
ER, JP
22
Latvia
23
24
Dierßen 1996 Diekmann
1994, Tyler et al. 2020
Szafer et al. 1988,
Rutkowski 2019, Popek 2002,
Zieliński 2004
Kuusk et al. 1996, 2003, Kukk
1999, Laasimer et al. 1993
Kuusk et al. 1996, 2003, Kukk 1999,
Laasimer et al. 1993
ML
Kuusk et al. 1996, 2003,
Gavrilova & Šulcs 1999, Laasimer
et al. 1993
Kuusk et al. 1996, 2003, Laasimer et
al. 1993
Lithuania
VM
Kuusk et al. 1996, 2003, Laasimer Baleavičienė, 1991
et al. 2003
Gudžinskas, 1999, Kuusk et al.
1996, 2003, Laasimer et al. 1993
Sweden, boreal zone (except
'alpine')
JB
Mossberg & Stenberg 2003, Tyler
et al. 2015,
https://www.dyntaxa.se/
Tyler et al. 2015
TT
Moen (1999)
Dierßen 1996
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Hilbig, W., Knapp, H.D. & Reichhoff, L. 1982. Übersicht über die Pflanzengesellschaften des südlichen Teiles der DDR. XIV. Die thermophilen, mesophilen und azidophilen Saumgesellschaften. Hercynia N.F. 19:
212–248.
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Oberdorfer, E. (ed.) 1978. Süddeutsche Pflanzengesellschaften. Teil II: Sand- und Trockenrasen, Heide- und Borstgras-Gesellschaften, alpine Magerrasen, Saum-Gesellschaften, Schlag- und Hochstauden-Fluren.
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Supporting information to the paper
Heinken, T. et al. The European Forest Plant Species List (EuForPlant): Concept and applications. Journal of Vegetation Science.
Appendix S2. Explanatory materials for forest definitions and forest affinity categories as an assistance for the regional contributors
Definition and explanation of categories of forest affinity: see Box 2 and Fig. 2
Additional materials for explanation and assignment: tree rows and scrubs
Tree row with scrub structures:
Such a stand (and pure hedgerows with similar ecological conditions) is not covered by the forest definition, but inhabitated by a number of 1.1
species (here: Anemone nemorosa, A. ranunculoides and Hedera helix).
Such conditions should not be overestimated and lead to a 2.1 classification of the mentioned species.
On the other hand, seedlings and saplings of trees and shrubs in grasslands should not be considered for species assignments!
Note for the following examples:
Plant species mentioned do not necessarily behave the same way in your region!
Additional materials for explanation and assignment: Examples of 1.2-habitats
Distribution of habitat types on a dry, southwest exposed slope:
1.1: closed forest
1.2: open forest due to extreme site conditions
O: extreme rock outcrops, too dry and open to fulfill the forest definition
Thermophytic mixed oak and beech forests:
•
•
•
taxa that can be found mainly in the closed forest (1.1, here Luzula luzuloides),
taxa which are restricted to open forests due to extreme site conditions (1.2, here thermophytic species like Campanula persicifolia),
and taxa which occur in forests as well as in the open landscape (2.1, here Polygonatum odoratum, Anthericum liliago*, Silene
nutans*, Hieracium spp.) .
* maybe also 1.2 in a region if not present in an open landscape.
Skid trail in a beech forest: Canopy opening and soil compaction alter the forest understorey.
Species which ± only occur in such situations are classified as 1.2 (here: Rubus idaeus), but many 1.1 species (here: Athyrium filix-femina,
Stachys sylvatica) also show a positive response to management.
Additional materials for explanation and assignment: Afforestations, successional and disturbed forests (2.2 habitats)
Wooded meadow / wooded pasture: forest structure, but mown or grazed
•
•
•
group 1.1 species (here: Anemone nemorosa, Maianthemum bifolium),
group 2.1 species (here: Glechoma hederacea, Lathyrus linifolius),
group 2.2 species (here: Dactylis glomerata, Trifolium pratense, Ranunculus acris, Rumex acetosa, which would not be present
without grazing or mowing.
In case of problems with wooded pasture landscapes: Look for the relative abundance of species in more closed forests and the actual open
landscape.
Robinia forest on former dry grassland:
•
group 2.1 species (here: Urtica dioica), widespread also in late-successional forests,
•
group 2.2 species (here: Ballota nigra, Chelidonium majus, Silene latifolia),
in forests ± restricted to such anthropogenic types.
Additional materials for explanation and assignment: Afforestations, successional and disturbed forests (2.2 habitats)
Open successional oligotrophic/mesotrophic birch forest (quagmire):
•
•
group 2.1 species (here: Thelypteris palustris, Potentilla palustris, Carex echinata (also in closed swamp or bog forests),
group 2.2 species (Menyanthes trifoliata, Drosera rotundifolia, Carex lasiocarpa) that do not occur in closed forests.
Softwood riparian alluvial forest (Salicion albae):
•
•
group 2.1 species (here: Cornus sanguinea, Rubus caesius, Phalaris arundinaceae (also in closed forests),
group 2.2 species (here: Calystegia sepium, Stellaria aquatica, Bidens spp.) that do not occur in closed forests.
Additional materials for explanation and assignment: Afforestations, successional and disturbed forests (2.2 habitats)
Pine forest on a mixed deciduous forest site (→ successional forest):
•
•
•
group 1.1 species such as Pyroloidae,
group 2.1 species such as Vaccinium myrtillus, Vaccinium vitis-idaea,
group 2.2 species favoured by (former) grazing (e.g. Juniperus communis , Nardus stricta).
Successional pine forests on formerly open landscape, e.g. dunes:
•
•
group 2.2 species such as Corynephorus canescens, Carex arenaria and Empetrum nigrum,
group 1.1 species such as Moneses uniflora and other Pyrolaidae may also occur here.
Note for all afforestations, successional and disturbed forests (2.2):
no species classified as O should frequently occur in such forests!
Supporting information to the paper
Heinken, T. et al. The European Forest Plant Species List (EuForPlant): Concept and applications. Journal of Vegetation Science.
Appendix S3. Widespread plant species of different forest species categories. All species occurring in at least 20 regions are listed; in the case of species groups with a few species with ≥20 occurrences, species with
10-19 occurrences are also listed and shaded in gray. Within a forest species category, species are arranged according to layers; species of the tree layer, shrub layer and lianas (in this order) with frame. Alien species in
the whole study area are in bold and regionally alien species in italics. Species with habitat shift: black, ± equal distribution; green, mainly in the first category; blue, mainly in the last category; brown, mainly in the middle
category.
No habitat shift
1.1
Carpinus betulus
Fagus sylvatica
Pseudotsuga menziesii
Tilia cordata
Daphne mezereum
Actaea spicata
Adoxa moschatellina
Allium ursinum
Anemone ranunculoides
Blechnum spicant
Brachypodium sylvaticum
Bromopsis benekenii
Cardamine bulbifera
Carex digitata
Carex elongata
Carex remota
Carex sylvatica
Circaea alpina
Circaea lutetiana
Circaea x intermedia
Corydalis cava
Corydalis solida
Dactylis glomerata subsp. lobata
Drymochloa sylvatica
Elymus caninus
Galium odoratum
Goodyera repens
Gymnocarpium dryopteris
Hepatica nobilis
Hypopitys monotropa
Impatiens noli-tangere
Impatiens parviflora
Lathraea squamaria
Lathyrus vernus Bernh.
Luzula pilosa
1.2 (down to 10 regions)
2.1
2.2
O
Humulus lupulus
Atropa bella-donna
Campanula persicifolia
Cardamine flexuosa
Digitalis purpurea
Lathyrus niger
Senecio ovatus
Silene baccifera
Stachys alpina
Trifolium rubens
Vicia dumetorum
Vicia pisiformis
Acer campestre
Acer pseudoplatanus
Alnus glutinosa
Betula pendula
Betula pubescens
Fraxinus excelsior
Pinus nigra subsp. nigra
Aesculus hippocastanum
Atriplex prostrata
Bellis perennis
Blysmus compressus
Carex cuprina
Carex hostiana
Carex lepidocarpa
Dactylorhiza majalis
Eriophorum latifolium
Malva moschata
Myosotis ramosissima
Phleum pratense
Salvia verticillata
Scorzoneroides autumnalis
Pinus sylvestris
Populus alba
Populus nigra
Populus tremula
Prunus avium
Prunus padus
Prunus serotina
Malus pumila
Populus x canadensis
Salix alba
Salix euxina
Salix pentandra
Salix x rubens
Cytisus scoparius
Juniperus communis
subsp. communis
Salix purpurea
Salix triandra
Salix viminalis
Syringa vulgaris
Quercus robur
Calystegia sepium
Sambucus nigra
Sorbus aucuparia
Berberis vulgaris
Corylus avellana
Cotoneaster integerrimus
Crataegus laevigata
Crataegus monogyna
Euonymus europaea
Frangula alnus
Ligustrum vulgare
Prunus spinosa
Rhamnus cathartica
Rosa canina
Rubus fruticosus agg.
Salix aurita
Salix cinerea
Viburnum lantana
Viburnum opulus
Lonicera periclymenum
Aegopodium podagraria
Agrimonia eupatoria
Agrostis stolonifera
Alchemilla spp.
Arrhenatherum elatius subsp. elatius
Carex leporina
Chelidonium majus
Cirsium vulgare
Danthonia decumbens
Epilobium ciliatum subsp. adenocaulon
Epilobium parviflorum
Festuca rubra
Galium mollugo
Holcus lanatus
Impatiens glandulifera
Luzula campestris
Persicaria minor
Phragmites australis
Poa pratensis
Prunella vulgaris
Reynoutria japonica
Habitat shift
4-5 assignments
Comarum palustre
Gnaphalium sylvaticum
Silene nutans
Ribes rubrum
Geranium sylvaticum
Hypericum montanum
Poa chaixii
Allium scorodoprasum
Carex ericetorum Pollich
Huperzia selago
Arctostaphylos uva-ursi
Lycopodium annotinum
Maianthemum bifolium
Mercurialis perennis
Milium effusum
Moehringia trinervia
Moneses uniflora
Neottia nidus-avis
Onoclea struthiopteris
Orthilia secunda House
Oxalis acetosella
Paris quadrifolia
Phegopteris connectilis
Polygonatum multiflorum
Polystichum aculeatum
Pyrola minor
Rumex sanguineus
Sanicula europaea
Veronica montana
Viola mirabilis
Viola reichenbachiana
Ajuga reptans
Alliaria petiolata
Angelica sylvestris
Avenella flexuosa
Calamagrostis canescens
Calluna vulgaris
Caltha palustris
Cardamine amara
Carex acutiformis
Carex canescens
Carex elata
Carex muricata
subsp. muricata
Carex pallescens
Carex paniculata
Carex pilulifera
Cirsium oleraceum
Crepis paludosa
Dactylorhiza fuchsii
Deschampsia cespitosa
Epilobium montanum
Epipactis atrorubens
Ficaria verna
Galeopsis bifida
Galium aparine
Galium palustre
Geranium robertianum
Geum rivale
Geum urbanum
Glechoma hederacea
Hieracium lachenalii
Hieracium murorum
Holcus mollis
Iris pseudacorus
Lactuca muralis
Reynoutria sachalinensis
Silene flos-cuculi
Solidago gigantea
Continuation 2.1
Lamium maculatum
Lapsana communis
Lathyrus linifolius
Luzula multiflora
Lycopus europaeus
Lysimachia nummularia
Lysimachia vulgaris
Molinia caerulea
Orchis mascula
Peucedanum palustre
Platanthera bifolia
Poa trivialis
Polypodium vulgare
Primula veris
Pteridium aquilinum
Ranunculus polyanthemos
Ranunculus repens
Scirpus sylvaticus
Scutellaria galericulata
Silene dioica
Solanum dulcamara
Solidago virgaurea
Stellaria alsine
Thelypteris palustris
Urtica dioica
Vaccinium vitis-idaea
Veronica chamaedrys
Veronica officinalis
Vicia sepium
Viola hirta
Habitat shift, 2 assignments
1.1&1.2
1.1&2.1
1.2&2.1
2.1&2.2
2.1&O
2.2&O
Malus sylvestris
Taxus baccata
Ribes spicatum
Abies alba
Acer platanoides
Alnus incana
Sambucus racemosa
Rubus caesius
Rubus idaeus
Populus x canescens
Robinia pseudoacacia
Anagallis minima
Arabis alpina
Asplenium adiantum-nigrum
Salix repens
Lycium barbarum
Aegonychon purpurocaeruleum
Bromopsis ramosa
Campanula latifolia
Carex pendula
Cephalanthera damasonium
Cypripedium calceolus
Picea abies
Pinus strobus
Quercus petraea
Tilia platyphyllos
Ulmus glabra
Cornus sanguinea
Aquilegia vulgaris
Cardamine impatiens
Chaerophyllum temulum
Epilobium angustifolium
Fragaria moschata
Fragaria vesca
subsp. adiantum-nigrum
Asplenium ruta-muraria
Asplenium trichomanes
Calla palustris
Carex colchica
Conopodium majus
Achillea ptarmica
Agrimonia procera
Agrostis gigantea
Agrostis vinealis
Alisma plantago-aquatica
Allium vineale
Salix caprea
Symphoricarpos albus
Agrostis capillaris
Anthoxanthum odoratum
Anthriscus sylvestris
Berula erecta Coville
Brachypodium pinnatum
Achillea millefolium
Galium sylvaticum
Melittis melissophyllum
Orobanche hederae
Schedonorus giganteus
Stachys sylvatica
Stellaria nemorum
subsp. montana
Ribes uva-crispa
Geranium sanguineum
Hedera helix
Hypericum hirsutum
Anemone nemorosa
Myosotis sylvatica
Athyrium filix-femina
Polygonatum odoratum
Calamagrostis arundinacea
Torilis japonica
Cephalanthera longifolia
Vaccinium uliginosum
Chrysosplenium alternifolium
Convallaria majalis
Dryopteris carthusiana
Dryopteris dilatata
Dryopteris filix-mas Schott
Equisetum sylvaticum
Galanthus nivalis
Leucojum vernum
Lycopodium clavatum
Melica nutans
Neottia ovata
Polygonatum verticillatum
Pyrola rotundifolia
Scrophularia nodosa
Stellaria holostea
Stellaria nemorum subsp. nemorum
Vaccinium myrtillus
Calamagrostis epigejos
Callitriche palustris agg.
Cardamine pratensis
Carex acuta
Carex flacca
Carex nigra
Carex pseudocyperus
Carex riparia
Carex rostrata
Cirsium palustre
Dactylis glomerata
subsp. glomerata
Epilobium palustre
Equisetum fluviatile
Equisetum palustre
Eriophorum angustifolium
Eupatorium cannabinum
Fallopia dumetorum
Galeopsis tetrahit
Glyceria fluitans
Glyceria maxima
Hieracium sabaudum
Hieracium umbellatum
Hottonia palustris
Juncus effusus
Lythrum salicaria
Mentha aquatica
Myosoton aquaticum
Origanum vulgare
Phalaroides arundinacea
Potentilla erecta
Serratula tinctoria
Stellaria media
Taraxacum sect. Taraxacum
Trifolium medium
Valeriana officinalis
Viola palustris
Cystopteris fragilis
Dactylorhiza maculata
Hieracium laevigatum
excl. species americ. borealis
Myosotis sparsiflora
Orobanche rapum-genistae
Polypodium x mantoniae
Scutellaria minor
Veronica hederifolia
Continuation 2.2&O
Lythrum portula
Malva alcea
Medicago falcata
Medicago lupulina
Mentha arvensis
Mentha longifolia
Myosotis arvensis
Myosotis laxa
Myrrhis odorata
Nardus stricta
Ochlopoa annua
Oenanthe aquatica
Ononis spinosa subsp. spinosa
Ononis spinosa
subsp. procurrens
Ophioglossum vulgatum
Ophrys insectifera
Orchis militaris
Oxalis stricta
Parnassia palustris
Pedicularis sylvatica
Picris hieracioides
Pilosella lactucella
Pilosella officinarum
Pilosella piloselloides
Pinguicula vulgaris
Plantago lanceolata
Plantago major
Plantago media
Poa angustifolia
Poa compressa
Poa palustris
Polygala amarella
Polygala vulgaris
Potentilla reptans
Potentilla tabernaemontani
Alopecurus pratensis
Angelica archangelica
Anisantha sterilis Nevski
Antennaria dioica
Anthyllis vulneraria
Arabidopsis arenosa subsp. aren.
Arabis hirsuta
Arctium lappa
Arctium minus
Arctium tomentosum
Arnica montana
Artemisia campestris
Artemisia vulgaris
Asparagus officinalis
Avenula pubescens
Ballota nigra
Bidens tripartitus
Bistorta officinalis
Botrychium lunaria
Briza media
Bromopsis erecta
Campanula glomerata
Carduus crispus
Carex caryophyllea
Carex demissa
Carex diandra
Carex dioica
Carex disticha
Carex flava
Carex hirta
Carex lasiocarpa
Carex panicea
Carex pulicaris
Carlina vulgaris
Centaurea jacea
Centaurium erythraea
Cerastium fontanum subsp. vulgare
Cicuta virosa
Cirsium acaulon
Cirsium arvense
Colchicum autumnale
Corynephorus canescens
Cruciata laevipes
Cuscuta europaea
Cynoglossum officinale
Dactylorhiza incarnata
Daucus carota
Ranunculus acris
Rhinanthus minor
Rorippa amphibia
Rumex acetosa
Rumex acetosella
Rumex hydrolapathum
Rumex obtusifolius
Sanguisorba minor
Sanguisorba officinalis
Saponaria officinalis
Saxifraga granulata
Scabiosa columbaria
Scorzonera humilis
Securigera varia
Sedum acre
Sedum album
Sedum sexangulare
Selinum carvifolia
Senecio leucanthemifolius
subsp. vernalis
Senecio viscosus
Senecio vulgaris
Silene latifolia
Solidago canadensis
Sparganium erectum
Stellaria graminea
Stellaria palustris
Succisa pratensis
Tanacetum parthenium
Thymus pulegioides
Thymus serpyllum
Trichophorum cespitosum
Trifolium pratense
Trifolium repens
Tussilago farfara
Typha latifolia
Utricularia vulgaris
Verbascum nigrum
Verbascum thapsus
Veronica beccabunga
Veronica serpyllifolia
Vicia cracca
Vicia tenuifolia
Vicia tetrasperma
Dianthus deltoides
Drosera rotundifolia
Elytrigia repens
Epilobium hirsutum
Epilobium obscurum
Epilobium tetragonum
Epipactis palustris
Equisetum variegatum
Erigeron acris
Erigeron annuus
Erigeron canadensis
Fallopia convolvulus
Galeopsis ladanum
Galium pumilum
Galium verum
Glyceria notata
Gnaphalium uliginosum
Gymnadenia conopsea
Helianthemum nummularium
Hypericum perforatum
Hypochaeris radicata
Inula helenium
Jacobaea vulgaris
Jasione montana
Juncus articulatus
Juncus bufonius
Juncus conglomeratus
Juncus filiformis
Juncus inflexus
Juncus squarrosus
Juncus subnodulosus
Juncus tenuis
Knautia arvensis
Koeleria pyramidata
Lamium album
Lathyrus pratensis
Lathyrus tuberosus
Lemna trisulca
Leontodon hispidus
Leucanthemum vulgare
Linaria repens
Linaria vulgaris
Linum catharticum
Lotus corniculatus
Lotus pedunculatus
Habitat shift, 3 assignments
1.1,1.2,2.1
Larix decidua
Crataegus rhipidophylla
Lonicera xylosteum
Ribes alpinum
Ribes nigrum
Campanula trachelium
Carex montana
Cephalanthera rubra
Epipactis helleborine
Equisetum hyemale
Gagea lutea Ker Gawl.
Lilium martagon
Melampyrum pratense
Phyteuma spicatum
Poa nemoralis
Ranunculus auricomus coll.
Rubus saxatilis
Vinca minor
Viola riviniana
1.1,2.1,2.2
1.1,2.1,2.2
1.2,2.1,2.2
1.2,2.1,O
2.1,2.2,O
Platanthera chlorantha
Luzula luzuloides
Scilla bifolia
Gymnocarpium robertianum
Equisetum telmateia
Sorbus intermedia
Cotoneaster bullatus
Hippophaë rhamnoides
Rosa micrantha
Spiraea alba
Pyrus communis subsp. communis
Rosa dumalis
Rosa rubiginosa
Rosa villosa
Clematis vitalba
Arctium nemorosum
Carex spicata
Clinopodium vulgare
Hesperis matronalis
Lathyrus sylvestris
Lithospermum officinale
Melampyrum nemorosum
Pleurospermum austriacum
Senecio sylvaticus
Stellaria neglecta
Vicia sylvatica
Viola odorata
Arabis glabra
Aristolochia clematitis
Bidens frondosus
Botrychium matricariifolium
Botrychium multifidum
Hieracium caesium
Lilium bulbiferum
Parietaria officinalis
Phytolacca americana
Sambucus ebulus
Trifolium aureum
Continuation 2.1,2.2,O
Myosotis scorpioides
Ornithogalum umbellatum
Persicaria hydropiper
Petasites hybridus
Pimpinella major
Pimpinella saxifraga
Ranunculus flammula
Ranunculus lingua
Sedum rupestre
Silene viscaria
Silene vulgaris
Stachys officinalis
Stachys palustris
Symphytum officinale
Thalictrum flavum
Thalictrum minus
Vaccinium oxycoccos
Valeriana dioica
Viola canina
Agrostis canina
Allium oleraceum
Andromeda polifolia
Asplenium septentrionale
Astragalus glycyphyllos
Campanula rapunculoides
Campanula rotundifolia
Carex appropinquata
Carex echinata
Carex vesicaria
Centaurea scabiosa
Empetrum nigrum
Epilobium roseum
Equisetum arvense
Eriophorum vaginatum
Euphorbia cyparissias
Festuca ovina
Filipendula ulmaria
Galeopsis speciosa
Galium boreale
Galium uliginosum
Glyceria declinata
Heracleum sphondylium
Hieracium levicaule
Hydrocotyle vulgaris
Hylotelephium telephium
Hypericum maculatum
Hypochaeris maculata
Lemna minor
Lysimachia thyrsiflora
Menyanthes trifoliata
Supporting information to the paper
Heinken, T. et al. The European Forest Plant Species List (EuForPlant): Concept and applications. Journal of Vegetation Science.
Appendix S4. Species numbers of forest species groups in the European forest species list across the 24 regions in temperate and boreal
Europe, sorted by the total number of species. Forest species are divided according to the forest affinity categories (1.1 – species of closed
forests, 1.2 – species of forest edges and forest clearings, 2.1 – species of forests and open vegetation, 2.2/O – species of mainly open
vegetation, but sporadically in forests). See Fig. 4 for the reason of merging the categories 2.2 and O Moreover, species classified as O
necessarily have a 2.2 habitat preference in at least one region, otherwise, they would not be part of the forest species list.
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