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303
Regional species pools of vascular plants in habitats of the Czech Republic
Druhy cévnatých rostlin v biotopech České republiky
Jiří S á d l o 1, Milan C h y t r ý 2 & Petr P y š e k 1,3
1
Institute of Botany, Academy of Sciences of the Czech Republic, CZ-252 43 Průhonice,
Czech Republic, e-mail: saadlo@volny.cz, pysek@ibot.cas.cz; 2Department of Botany
and Zoology, Masaryk University, Kotlářská 2, CZ-611 37 Brno, Czech Republic, e-mail:
chytry@sci.muni.cz; 3Department of Ecology, Charles University, Viničná 7, CZ-128 01
Praha, Czech Republic
Sádlo J., Chytrý M. & Pyšek P. (2007): Regional species pools of vascular plants in habitats of the
Czech Republic. – Preslia 79: 303–321.
Based on a combination of data from the Czech National Phytosociological Database and expert
knowledge, a database of vascular plant species pools for 88 habitats, representative of the diversity
of Czech vegetation, was compiled. This database contains 1820 native species, 249 archaeophytes
and 278 neophytes, each assigned to one or more habitats. Besides the data on species occurrence in
different habitats, the database contains information on a species’ ecological optimum in the habitat
or its dominance. The largest pools of native species were found in rather rare habitats of dry and
warm herbaceous or woody habitats at low altitudes, some of which contain > 530 species (maximum of 695 species for thermophilous forest fringes). These were followed by common habitats on
mesic soils. The smallest pools of native species were in saline, aquatic and bog habitats (< 90 species). Species pool sizes of archaeophytes and neophytes for different habitats were positively, yet
weakly, correlated with the species pool sizes of native species. Habitats with native species pools
< 350 species contained any number of archaeophytes. Habitats with < 100 native species contained
< 5, and often no neophytes, but habitats with 100–350 native species contained different numbers
of neophytes. Habitats with > 350 native species always contained > 5 archaeophytes and > 5 neophytes, and often many more. Two hundred and thirty two native species, 18 archaeophytes and 30
neophytes were identified as potential dominants in at least one habitat. However, potentially dominant species made up less than 3% of the species pool for 78 out of 88 habitats. Larger percentages
(up to 14.6%) of potential dominants were included in habitats with small species pools and speciespoor stands (e.g., aquatic, saline and mire habitats). The number of habitats in which a species occurred was used as a measure of its ecological range. Most ecological generalists were found among
the native species, less among the archaeophytes and least among the neophytes. Out of the 36 species that occur as dominants in three or more habitats, 34 were native (many are grasses), one was an
archaeophyte (Cirsium arvense) and one was a neophyte (Impatiens parviflora).
K e y w o r d s : alien, archaeophyte, habitat classification, local and regional processes, native, neophyte, species richness, vegetation type
Introduction
Species pool is the set of species that are potentially capable of coexisting in a particular
community (Zobel 1997). The concept of species pool explains local species richness in
terms of historical and evolutionary processes operating at large spatial and temporal
scales (Taylor et al. 1990, Cornell & Lawton 1992, Zobel 1992, Ricklefs & Schluter
1993). According to the species pool theory, species richness of a local plant community
not only depends on local processes existing within this community, such as competition
(Palmer 1994), but also to a large extent, on the availability of propagules of those species
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that are able to disperse to and grow in the target community. In addition to studies on the
variation in local species richness, the species pool theory is important in studies of habitat
invasibility by alien plants (Chytrý et al. 2005, Stohlgren et al. 2006). Consistent with the
species pool theory is the recognition in invasion ecology that the levels of invasion of particular habitats not only depend on habitat properties (habitat susceptibility to invasion),
but also propagule pressure, i.e. the size of alien species pool (Williamson 1996, Lonsdale
1999, Daehler 2006, Richardson & Pyšek 2006, Richardson 2006). The species pool concept is also of great importance in applied fields such as restoration ecology (Zobel et al.
1998), where the success of restoration projects, especially those relying on spontaneous
succession (Prach et al. 2001, Prach 2003, Ruprecht 2005), strongly depends on the migration of species from the local species pools into the restored sites.
When taking species pool effects into account, the key issue is the identification of the
set of species that are in the species pool of the target community (Pärtel et al. 1996). Of
the different attempts to tackle this problem (see an overview in Zobel et al. 1998), perhaps
the most reliable is a combination of the relevé data existing in large phytosociological databases (Hennekens & Schaminée 2001) and expert knowledge of the habitat affinities of
the species in the regional flora. In this way a relatively standardized list of species belonging to the species pool can be identified for each habitat occurring in the target region. If
the target region is larger than a landscape in which species can migrate to a particular site
within a few years, such lists correspond to the definition of regional species pool sensu
Pärtel et al. (1996) and Zobel (1997) rather than to a local or actual species pool. The lists
of regional species pools for particular habitats can be useful both in theoretical studies on
the control of local species richness or habitat invasibility and in practical applications
such as nature conservation or ecological restoration. However, even a simple comparison
of the sizes and compositions of regional species pools among habitats can provide interesting ecological insights into the history of the formation of regional floras.
To facilitate future research into diversity and invasibility of Central European plant
communities and future restoration projects, a database of the regional species pools of
vascular plant for 88 habitats, which are representative of the vegetation diversity of the
Czech Republic, was compiled. In this paper, statistical summary of the regional species
pools of Czech habitats, extracted from this database, and some basic correlations between
numbers of native and alien species are presented.
Materials and methods
The habitat classification used in this study (Appendix 1) was prepared in co-operation
with I. Kühn, S. Klotz (Halle) and G. Karrer (Vienna) for the purpose of describing habitat
affinities in the future database of species traits of German, Czech and Austrian flora. Habitat delimitation is largely based on phytosociological syntaxa, mainly at the level of alliance. Appendix 1 lists only the habitats occurring in the Czech Republic.
Species in the Czech flora were assigned to habitats in several steps. First, 24,283
relevés from the Czech National Phytosociological Database (Chytrý & Rafajová 2003)
were assigned to habitats, using the assignment of these relevés to phytosociological
syntaxa provided by the relevé’s authors. Then, the percentage occurrence frequency and
fidelity (according to Tichý & Chytrý 2006) of each species to each habitat, based on the
Sádlo et al.: Habitat plant species pools
305
relevé data, were calculated using the JUICE 6.4 program (Tichý 2002). This provided the
first approximation, because many rare species may be under-represented in relevés, as are
species occurring in ecotonal or successional situations or small patches, which do not satisfy the phytosociological requirement for homogeneous plots (Westhoff & van der
Maarel 1973).
This database was further improved and corrected by the first author, using handbooks
on Czech flora (Hejný & Slavík 1988–1992, Slavík 1995–2000, Kubát et al. 2002, Slavík
& Štěpánková 2004), papers on individual species, local and national vegetation studies
(e.g., Moravec et al. 1995, Moravec 1998–2003, Kočí 2001, Havlová et al. 2004, BottaDukát et al. 2005, Havlová 2006, Kropáč 2006, Lososová et al. 2006, Chytrý 2007), and
expert knowledge of species habitat affinity in the field. Often there were no reliable data
on the occurrence of some species in some habitats; in such cases, species presence or absence in the habitat was assessed, with some degree of uncertainty, from the reliable information on its presence or absence in ecologically similar and close by habitats.
Some species recorded in single relevés from some habitats were deleted if such records represented sporadic exceptions to the habitat range of that plant (e.g., Coronilla
vaginalis in a forest clearing or Dryopteris carthusiana growing in saline grassland).
Seedlings and saplings were not included in the species pools of habitats in which they had
little chance of survival (e.g., Abies alba in a meadow). Trees and shrubs were generally
not included in species pools of meadows, pastures, annual anthropogenic vegetation,
cliffs and very wet or inundated marshes.
The following groups of taxa were excluded from the database: (1) Species which disappeared from the flora of the Czech Republic before 2000 (Holub 2000). (2) Hybrids or
hybridogenous taxa, with 19 exceptions, those that differ in habitat affinities from their parents or have at least one parent species not occurring in the Czech Republic (e.g., Circaea
×intermedia, Pinus ×pseudopumilio, Prunus ×fruticans, Sorbus ×sudetica and Viola
×wittrockiana were all included). (3) Agamospecies of Rubus and Taraxacum were excluded and replaced by series and sections, respectively. However, agamospecies of
Alchemilla, Hieracium and Sorbus were included. (4) Of 817 casual neophytes present in
the Czech flora (Pyšek et al. 2002a), 119 relatively frequent species were included and 698
were excluded from the database. Excluded casual neophytes were those that occur as garden escapees but do not survive for long (e.g., Allium cepa and Petunia hybrida), do survive
for long as garden escapees but are rare (e.g., Arabis procurrens), used to escape from gardens but do not escape today (e.g., Chenopodium foliosum and Lathyrus ochrus), are beginning to escape but are still rather rare (e.g., Amaranthus caudatus), or lack clear habitat affinity regardless of how common they are and for how long they have been present as casuals
(e.g., Artemisia dracunculus, Spiraea douglasii, Ribes aureum). In total, 2347 species were
included in the database, of which 1820 are native, 249 archaeophytes and 278 neophytes.
Each species-habitat assignment was classified to one of the following categories: (1)
occurrence – species can grow in the habitat, but it is not ecological optimum for this species, which often is rare in this habitat; (2) optimum – the habitat or part of it is ecological optimum for this species; (3) dominant – species can be assigned to the previous category and
at the same time frequently attains a cover above 25% in areas ≥ 10 m2 or ≥ 100 m2 in herbaceous or woody vegetation, respectively; (4) constant dominant – same as for the previous category but also determines the general appearance of the habitat (e.g., Calluna
vulgaris in heathlands), occurring in ≥ 40% of the localities of the habitat.
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Preslia 79: 303–321, 2007
The database reflects species’ habitat affinities as recorded or observed in 1990–2007,
i.e. in the current period of landscape development, following the socio-economic change
of 1989. The only exception is the inland vegetation of succulent halophytes, which had
disappeared by the 1970s (Šumberová 2007), but was included in the habitat list and its
species pool reconstructed based on a detailed phytosociological survey from the 1960s
(Vicherek 1973).
Taxonomy and nomenclature of plants follow Kubát et al. (2002), except for taxa of
Centaurea sect. Jacea, which follow Štěpánek & Koutecký (2004). Classification of species as native/alien follows the list in Pyšek et al. (2002a) and recent terminological proposals (Richardson et al. 2000, Pyšek et al. 2004). Exceptions include Arrhenatherum
elatius, which is now considered to be an archaeophyte rather than a neophyte (Chytrý et
al. 2005), and Cytisus scoparius, Imperatoria ostruthium, Mimulus guttatus and Myrrhis
odorata, which are considered to be naturalized rather than invasive. Names of
phytosociological syntaxa follow Moravec (1998–2003) for forests, Chytrý (2007) for
grassland and heathland vegetation and Moravec et al. (1995) for other vegetation types.
The basic statistical figures characterizing the database presented in this paper were obtained using the programs MS Excel, MS Access (www.microsoft.com) and Statistica
(www.statsoft.com). Numbers of archaeophytes and neophytes had a strongly rightskewed distribution, which was transformed to normal by log10 transformation before correlation and regression analyses. Number of native species was normally distributed and
therefore used without transformation. Normality was tested using the KolmogorovSmirnov test at P < 0.05.
Results and discussion
Pools of native species
The largest regional pools of native species (Table 1) were identified in dry and warm herbaceous or woody habitats at low altitudes (thermophilous forest fringe vegetation – 695
species, tall mesic and xeric scrub – 672, broad-leaved dry grasslands – 549, peri-Alpidic
basiphilous thermophilous oak forests – 535). It is remarkable that these habitats harbour
so many species although occupying restricted areas in the Czech Republic. Concentration
of many species in the dry low-altitude habitats may reflect their long historical continuity.
Similar habitats may have occurred there in the Pleistocene and most of the species in their
regional species pools may have survived in situ up to the present, because at low altitudes
in the Czech Republic there were presumably many open areas and open-canopy forests
throughout the Holocene (Ložek 1973, Sádlo et al. 2005). Thus, regional species pool
sizes for these habitats may be a consequence of historical inertia rather than contemporary effects of the size of the area of these particular habitats (Aarssen & Schamp 2002,
Pärtel 2002, Ewald 2003). Additional reasons for these habitats having large regional species pools include: (i) each of these habitats occur at a wide range of diverse sites (e.g.,
soils of varying quality), therefore they include a number of species with different niches;
(ii) these habitats occur at the border of forest and naturally treeless areas, which results in
ecotone effects on species diversity.
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Sádlo et al.: Habitat plant species pools
Table 1. – Numbers of species and dominant species occurring in particular habitats. Note that casual aliens were only assessed if
they exhibited a clear affinity for a particular habitat (see Materials and methods for details). Delimitations of individual habitats are
described in Appendix 1.
Neophytes naturalized
Neophytes invasive
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0 0
0 3
1 2
0 2
0 3
0 6
1 10
0
11
10
11
4
33
11
1
7
39
69
1 0 1 2 41
0 1 5 6 224
2 2 9 13 289
1 3 5 9 183
2 2 4 8 180
0 11 19 30 330
1 6 13 20 293
0 1 2 3 109
0 3 6 9 316
1 3 9 13 289
7 10 21 38 342
5
3
6
3
3
3
2
1
7
8
6
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
1
1
0
1
0
0
0
0
0
0
0
1
6
3
7
3
3
3
2
1
7
8
9
0 11 5 16
0 15 4 19
3 34 11 48
0 4 9 13 164
1 7 15 23 358
1 10 23 34 274
3
3
3
0
0
0
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
3
0
0
3
3
3
7
1 0 0 2 2 117
3 0 2 4 6 273
0 0 0 1 1 174
4 0 2 1 3 325
3 0 3 2 5 331
1 0 1 1 2 230
0 0 0 0 0 90
1 0 0 0 0 111
49 9 15 15 39 593
9 3 5 7 15 335
58 12 20 9 41 573
39 4 8 9 21 424
12 1 6 8 15 382
19 0 3 7 10 468
64 4 12 15 31 439
2 0 1 2 3 301
0
7
2
4
4
5
8
4
5
3
5
3
4
6
3
4
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
7
2
4
4
5
8
4
6
3
5
3
4
6
3
4
4
0
0
0
0
0
0
0
0
4
0 0 0
0 9 2
0 5 5
0 6 5
0 3 1
0 25 8
0 8 3
0 1 0
0 3 4
5 28 6
6 52 11
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1 0
2 1
0 0
3 1
2 1
1 0
0 0
1 0
41 8
6 3
52 6
34 5
10 2
15 4
53 11
2 0
0 10
2 12
0
4
4
8 430
All species
Neophytes casual
0
0
0
0
0
0
0
Neophytes all
Archaeophytes all
0
0
1
0
0
0
0
All species
0
0
0
0
0
0
0
Neophytes all
0
3
0
2
3
6
9
Neophytes invasive
25 1 4 4 9 286
19 0 3 4 7 396
41 10 10 13 33 176
40 1 2 5 8 326
0 0 0 0 0 107
3 0 0 5 5 334
0 3 0 1 4 86
Neophytes naturalized
4
5
7
8
0
3
0
Neophytes casual
20
13
34
32
0
0
0
Archaeophytes all
Archaeophytes invasive
Archaeophytes invasive
7B
Archaeophytes naturalized
5A
5B
5C
5D
5E
5F
5G
5H
6A
6B
6C
6D
6E
6F
6G
7A
Archaeophytes casual
4J
4K
4L
1
1
0
0
0
0
0
No. of dominant species
Native species
3B
3C
4A
4B
4C
4D
4E
4F
4G
4H
4I
252
Calcareous cliffs
370
Siliceous cliffs and block fields
102
Walls
278
Mobile calcareous screes
107
Alpine grasslands on siliceous bedrock
Subalpine tall-forb and tall-grass vegetation 326
82
Macrophyte vegetation of eutrophic and
mesotrophic still waters
39
Macrophyte vegetation of water streams
Vegetation of oligotrophic lakes and pools 207
266
Reed-beds of eutrophic still waters
163
Halophilous reed and sedge beds
168
Eutrophic vegetation of muddy substrata
267
Riverine reed vegetation
262
Reed vegetation of brooks
Mesotrophic vegetation of muddy substrata 105
300
Tall-sedge beds
Vegetation of low annual hygrophilous herbs 237
235
Vegetation of nitrophilous annual
hygrophilous herbs
135
River gravel banks
316
Petasites fringes of montane brooks
Nitrophilous herbaceous fringes of lowland 192
rivers
114
Hard-water springs with tufa formation
264
Lowland and montane soft-water springs
173
Alpine and subalpine soft-water springs
318
Calcareous fens
Acidic moss-rich fens and peatland meadows 323
227
Transitional mires
90
Raised bogs
110
Wet peat soils and bog hollow
505
Mesic Arrhenatherum meadows
311
Montane mesic meadows
474
Pastures and park grasslands
364
Alluvial meadows of lowland rivers
355
Wet Cirsium meadows
439
Intermittently wet Molinia meadows
344
Vegetation of wet disturbed soils
296
Subalpine and montane acidophilous
grasslands
410
Submontane Nardus grasslands
Archaeophytes naturalized
1A
1B
1C
1D
2A
2B
3A
Archaeophytes casual
Native species
No. of species
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Native species
Archaeophytes casual
Archaeophytes naturalized
Archaeophytes invasive
Archaeophytes all
Neophytes casual
Neophytes naturalized
Neophytes invasive
Neophytes all
All species
0
0
0
0
0
0
0
0
0
0
3
2
5 7
3 8
3 6
1 13
1 5
0 5
0 1
1 4
4
4
4
5
8
6
3
4
555
600
446
757
303
327
115
326
6
2
5
6
1
3
1
3
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
6
2
5
7
1
3
1
3
5 64
0
4
3
7 341
4
0
0
0
0
0
0
0
0
4
1
6
0
0
0
0
37
3
0
1
0
1
0
0
0
0
4
0 0
9 46
2 10
2 8
0 0
2 2
1 5
7 27
0
1
0
0
0
0
0
3
0 0 0
6 11 18
1 0 1
2 3 5
0 1 1
0 2 2
2 2 4
4 18 25
2
6
3
4
4
5
5
6
Archaeophytes all
Neophytes casual
Neophytes naturalized
471
549
389
695
232
280
99
253
1
0
0
0
1
0
0
1
4
4
5
7
7
4
1
7
270
1 58
0
31
Archaeophytes casual
6
3
5
68
36
44
43
57
36
12
64
16
15
13
19
14
11
4
9
All species
0
0
4
2
Neophytes all
0
0
3 41
3 25
Neophytes invasive
0
0
Archaeophytes invasive
3
2
0 38
0 22
63
32
39
36
49
32
11
56
No. of dominant species
4 14 439
4 9 467
384
433
Native species
8A Hercynian dry grasslands on rock outcrops
8B Submediterranean dry grasslands on rock
outcrops
8C Narow-leaved sub-continental steppes
8D Broad-leaved dry grasslands
8E Acidophilous dry grasslands
8F Thermophilous forest fringe vegetation
9B Open vegetation of acidic sands
9C Festuca grasslands on acidic sands
9D Pannonian sand steppes
9E Acidophilous vegetation of spring
therophytes and succulents
9F Basiphilous vegetation of spring therophytes
and succulents
10G Continental vegetation of annual halophilous
grasses
10H Inland vegetation of succulent halophytes
10I Inland saline meadows
10J Inland saline steppes
11A Dry lowland to alpine heathlands
11D Subalpine acidophilous Pinus mugo scrub
11H Subalpine deciduous scrub
11I Willow carrs
11J Willow galleries of loamy and sandy river
banks
11L Tall mesic and xeric scrub
11N Low xeric scrub
11R Scrub and pioneer woodland of forests
clearings
12A Alder carrs
12B Alluvial forests
12C Oak-hornbeam forests
12D Ravine forests
12E Herb-rich beech forests
12F Limestone beech forests
12G Acidophilous beech forests
12H Peri-Alpidic basiphilous thermophilous
oak forests
12I Sub-continental thermophilous oak forests
12J Acidophilous thermophilous oak forests
12K Acidophilous oak forests
12L Boreo-continental pine forests
12O Peri-Alpidic pine forests
12P Peatland pine forests
12Q Peatland birch forests
12R Acidophilous spruce forests
Archaeophytes naturalized
No. of species
15
269
57
443
160
243
238
288
0 0
0 37
0 8
0 6
0 0
0 0
0 4
0 20
15
333
68
456
161
247
247
340
2
6
2
3
4
5
5
6
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
672
463
431
4 57 13 74 6 16 20 42 788
2 31 6 39 2 2 6 10 512
4 76 16 96 25 36 47 108 635
12
6
13
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
2
1 13
0 6
2 15
315
449
481
453
362
341
268
535
0
2
2
1
0
0
0
2
3
10
5
8
6
1
2
4
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
0
0
0
0
0 3
1 11
1 6
1 9
0 6
0 1
0 2
0 4
460
458
404
322
298
67
118
211
1 10
2 20
1 9
0 5
0 7
0 0
0 0
0 0
4
2
9
7
5
4
2
4
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
3
14
11
15
3
3
1
22
1
5
4
4
2
1
0
8
4
21
17
20
5
4
1
32
5 16
7 29
1 11
1 6
1 8
0 0
0 0
0 0
0 1 4 5 324
5 16 24 45 515
4 13 6 23 521
1 16 5 22 495
0 4 2 6 373
0 1 1 2 347
0 2 4 6 275
1 6 3 10 577
1
0
1
0
0
0
0
0
2
8
8
3
2
0
0
0
3 6 482
4 12 499
6 15 430
3 6 334
0 2 308
0 0 67
1 1 119
0 0 211
4
2
9
7
5
4
2
4
309
Sádlo et al.: Habitat plant species pools
Native species
Archaeophytes casual
Archaeophytes naturalized
Archaeophytes invasive
Archaeophytes all
Neophytes casual
Neophytes naturalized
Neophytes invasive
Neophytes all
All species
0
0
1
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
2
5
0
1
2
0
0
2
7
0
2
6
0
2
2
0
0
2
7
3
10
7
3
5
10
4
2
13
19
Archaeophytes all
Archaeophytes casual
0 47
9 56
2 12 25 39 553
16
0
0
1
1
0
0
0
0 17
0 0
21 40
29 81
8 14
9 18
27 148
12 74
14 46
37 144
45 119
All species
0
2
0
0
0
5
1
0
5
0
Neophytes all
0
0
0
0
0
2
0
0
3
0
Neophytes invasive
0
2
0
0
0
3
1
0
2
0
Neophytes naturalized
0
0
0
0
0
0
0
0
0
0
Neophytes casual
Archaeophytes invasive
3
6
1
3
3
3
3
2
6
12
0
14
32
5
9
45
25
17
58
48
458
No. of dominant species
125
421
431
369
584
561
398
302
715
620
125 0 0 0 0 0
293 1 73 14 88 5
289 1 47 13 61 20
344 0 7 4 11 1
525 0 34 7 41 0
202 16 178 17 211 76
146 16 145 17 178 37
154 6 84 12 102 15
386 6 160 19 185 49
404 4 75 18 97 26
Native species
12S Basiphilous spruce and fir forests
12T Robinia pseudacacia plantations
12U Plantations of broad-leaved non-native trees
12V Spruce plantations
12W Pine and larch plantations
13A Annual vegetation of ruderal habitats
13B Annual vegetation of arable land
13C Annual vegetation of trampled habitats
13D Perennial thermophilous ruderal vegetation
13E Perennial nitrophilous herbaceous
vegetation of mesic sites
13F Herbaceous vegetation of forests
clearings and Rubus scrub
Archaeophytes naturalized
No. of species
These rare dry and warm habitats are followed, in terms of the regional species pool
size, by widespread habitats on mesic soils (Pinus and Larix plantations – 525 species,
mesic Arrhenatherum meadows – 505, oak-hornbeam forests – 481, pastures and park
grasslands – 474). It is important to note that of the 525 native species in Pinus-Larix plantations, only 86 have their optima in this habitat. The rich regional species pool is therefore
composed of species that find their suboptima there, which is a general feature of the occurrence of native species in human-made habitats.
In contrast, the smallest regional species pools of native species are found in saline,
aquatic and bog habitats (inland vegetation of succulent halophytes – 15 species; continental vegetation of annual halophilous grasses – 31; macrophyte vegetation of water
streams – 39; inland saline steppes – 57; peatland pine forests – 67; macrophyte vegetation
of eutrophic and mesotrophic still waters – 82; raised bogs – 90). All of these habitats are
relatively rare and subject to strong environmental stress. To some extent, however, the
low numbers of species assigned to regional pools for some of these habitats can be due to
limited data. Rarity or even extinction in the case of the halophilous habitats makes it difficult to identify all the species that should be in regional species pools.
The sizes of regional species pools are well reflected in the local species richness of
particular habitats. However, the local species richness and regional species pool size are
not formally compared in this paper, because the data on local species richness, in the form
of phytosociological relevés, are less reliable than the present species pool data. Relevés
originate from different sources (Chytrý & Rafajová 2003), from plots of varying size
(Chytrý & Otýpková 2003) and were made by preferential sampling, which results in
a bias towards species-richer sites (Diekmann et al. 2007, Hédl 2007, but see Botta-Dukát
310
Preslia 79: 303–321, 2007
Fig. 1. – Relationships between the number of archaeophytes and neophytes, respectively, and the number of native species in the regional species pools of particular habitats. Regressions and correlations were calculated using log-transformed numbers of archaeophytes and neophytes; labels on vertical axes are back-transformed to
original values.
et al. 2007) or artificial interactions between the magnitude of the bias and plot size
(Chytrý 2001). Still, there are rough estimates of local species richness of individual vegetation types made during the preparation of the vegetation survey of the Czech Republic
(Chytrý 2007), based on the relevés in the national phytosociological database (Chytrý &
Sádlo et al.: Habitat plant species pools
311
Rafajová 2003). These analyses indicate good agreement between the local species richness of particular vegetation types and the regional species pool sizes presented here. For
example, broad-leaved dry grasslands, basiphilous thermophilous oak forests and forest
fringes were among the locally richest vegetation types, while bogs, saline and aquatic
vegetation were among the poorest. These patterns might be indicative of the effect of the
regional species pool size on the local species richness, although there are several pitfalls
in the interpretation of correlations between these two variables (Zobel 1997, Herben
2000, Grace 2001).
Pools of alien species
There are significant positive correlations between the regional species pool sizes of
archaeophytes and native species, and neophytes and native species (Fig. 1). These results
are different from those obtained from phytosociological relevés of the Czech Republic
(Chytrý et al. 2005), in which the relationships between the number of archaeophytes or
neophytes, respectively, and the number of native species occurring in the relevés were insignificant, if calculated across all habitats. This difference between the regional species
pools and the relevé data probably relates to the more general issue of scale-dependence of
the native-alien relationships (Fridley et al. 2004, Herben et al. 2004, Stohlgren et al.
2006). In addition, correlation between neophytes and native species in the
phytosociological data may be weak or absent due to the fact that neophytes are generally
poorly represented in small plots, although making up a considerable proportion of the
country’s flora (Pyšek et al. 2002a, Chytrý et al. 2005). However, the relationships revealed in the present study are rather weak even for the regional species pools (Fig. 1).
The scatter plots of Fig. 1 indicate that habitats with native species pools < 350 species
may contain any number of archaeophytes. Habitats with < 100 native species contain < 5,
and often no neophytes, but habitats with 100–350 species may contain different numbers of
neophytes. It is remarkable that habitats with > 350 native species always contain > 5
archaeophytes and > 5 neophytes, and often many more. Thus, habitats supporting many native species also tend to support aliens. This is consistent with the currently accumulating evidence of a positive relationship between native and alien species richness existing at larger
spatial scales (Planty-Tabacchi et al. 1996, Lonsdale 1999, Stohlgren et al. 1999, 2006,
Pyšek et al. 2002b, Deutschewitz et al. 2003, Kühn et al. 2003, Dark 2004, Espinosa-García
et al. 2004, Herben et al. 2004, Gilbert & Lechowicz 2005, Pino et al. 2005, Palmer 2006).
In spite of the generally positive relationships between the regional species pool sizes
of alien and native species, the largest pools of alien species are found in habitats other
than the largest pools of native species (Table 1). Largest numbers of both archaeophytes
and neophytes occur in herbaceous ruderal vegetation, on arable land, in scrub and pioneer
woodland in forest clearings and plantations of non-native trees. In contrast, no aliens are
included in the regional species pools of natural spruce forests, bogs, alpine grasslands
and vegetation consisting of succulent halophytes. There is a strong positive relationship
between the regional species pool sizes of archaeophytes and neophytes (r = 0.804,
P < 0.001). The size of alien pools for particular habitats closely corresponds to the proportions of alien species recorded in phytosociological relevés from the Czech Republic
(Chytrý et al. 2005). In that study, relevé data also indicated significant positive relationship between archaeophytes and neophytes.
312
Preslia 79: 303–321, 2007
Fig. 2. – Percentage frequency distribution of species representation in regional species pools of different habitats. The longer the right tail of the distribution (such as in native species), the more habitat generalists there are in
that species group.
Pools of dominant species
Species classified as dominants (including constant dominants) make up less than 3% of
the regional species pool for 78 out of 88 habitats. Larger percentages of dominants were
found only in habitats with small regional species pools and species-poor stands
(macrophyte vegetation of water streams – 14.6%, inland vegetation of succulent
halophytes – 13.3%, macrophyte vegetation of eutrophic and mesotrophic still waters –
11.6%, continental vegetation of annual halophilous grasses – 10.8%, raised bogs – 8.9%,
peatland pine forests – 6.0%, inland saline steppes – 4.4%). Most species that are potential
dominants are included in the species pools of nutrient-rich and disturbed habitats (Table
1). Most native dominants occur in herbaceous vegetation of forest clearings and Rubus
scrub (16 species), scrub and pioneer woodland of forest clearings (13), perennial
nitrophilous herbaceous vegetation of mesic sites (12) and tall mesic and xeric scrub (12).
Most dominant archaeophytes were found in perennial thermophilous ruderal vegetation
313
Sádlo et al.: Habitat plant species pools
Table 2. – Native and alien species occurring in most habitats, and the number of habitats (n = 88). Asterisks indicate alien species that were recorded in the highest numbers of habitats in the previous study (Chytrý et al. 2005),
which was based on species occurrences in vegetation relevés.
Native species
Festuca rubra
Taraxacum sect. Ruderalia
Agrostis stolonifera
Deschampsia cespitosa
Calamagrostis epigejos
Cerastium holosteoides
subsp. triviale
Dactylis glomerata
Veronica chamaedrys
Agrostis capillaris
Equisetum arvense
Urtica dioica
Galium aparine
Poa angustifolia
Avenella flexuosa
Achillea millefolium
subsp. millefolium
Aegopodium podagraria
Cirsium palustre
Galium album
Galeopsis bifida
Heracleum sphondylium
Ranunculus repens
Rubus idaeus
Rumex acetosa
Elytrigia repens
Hypericum perforatum
Poa trivialis
Galeopsis tetrahit s. str.
Angelica sylvestris
Anthoxanthum odoratum s. str.
Carex hirta
Epilobium montanum
Archaeophytes
Neophytes
78
77
73
67
66
63
*Arrhenatherum elatius
*Cirsium arvense
*Lapsana communis
*Medicago lupulina
*Tanacetum vulgare
*Fallopia convolvulus
62
44
40
40
39
38
*Epilobium ciliatum
*Impatiens parviflora
*Conyza canadensis
*Trifolium hybridum
*Robinia pseudacacia
*Bidens frondosa
45
45
34
28
24
22
62
62
61
60
60
59
59
57
56
Plantago major
*Convolvulus arvensis
*Mentha arvensis
Myosotis arvensis
*Lamium album
*Linaria vulgaris
*Lactuca serriola
Tripleurospermum inodorum
Veronica arvensis
38
37
37
35
33
33
31
30
29
*Agrostis gigantea
*Aster lanceolatus
*Medicago sativa
*Cytisus scoparius
*Juncus tenuis
Lolium multiflorum
Impatiens glandulifera
Pinus nigra
Rumex thyrsiflorus
21
19
18
17
17
17
16
16
16
56
56
55
55
55
55
55
55
54
54
54
53
52
52
52
52
*Echium vulgare
Vicia hirsuta
Chelidonium majus
Melilotus albus
Ballota nigra
Aethusa cynapium
Arctium lappa
A. tomentosum
Bromus hordeaceus
Capsella bursa-pastoris
Carduus acanthoides
Melilotus officinalis
Vicia angustifolia
V. sativa
*Cirsium vulgare
Geranium columbinum
28
28
26
26
25
24
24
24
24
24
24
24
24
24
23
23
Sisymbrium loeselii
Populus × canadensis
*Veronica persica
Ribes rubrum
Heracleum mantegazzianum
*Lupinus polyphyllus
Matricaria discoidea
Oxalis fontana
Quercus rubra
*Solidago canadensis
S. gigantea
Veronica filiformis
*Aster novi-belgii s. str.
*Erigeron annuus
Galinsoga parviflora
G. quadriradiata
Lonicera caprifolium
Sedum spurium
Sisymbrium strictissimum
Solanum decipiens
16
15
15
15
14
14
14
14
14
14
14
14
13
13
13
13
13
13
13
13
(5) and annual vegetation in ruderal habitats (5), and most dominant neophytes in perennial nitrophilous herbaceous vegetation of mesic sites (7) and plantations of broad-leaved
non-native trees (6).
Two hundred and thirty two native species, 18 archaeophytes and 30 neophytes were
identified as potential dominants in at least one habitat. These figures correspond, respectively, to 12.7%, 7.2% and 10.8% of the total number of species within these categories,
which suggests that there are hardly any differences in the ability of native and alien species to become dominants.
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Preslia 79: 303–321, 2007
Table 3. – Native and alien species that are dominant (including constant dominants) in ≥ 3 habitats, with numbers
of habitats in which species are dominant and % of habitats in which they are dominant relative to all the habitats
in which they occur.
Species
No. of habitats
% as dominant
Status
9
9
8
7
6
6
5
5
5
5
5
4
4
4
4
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
3
16
19
13
14
10
19
12
11
10
13
16
6
11
7
9
5
7
6
8
11
14
13
8
7
11
7
4
8
19
9
9
12
18
7
9
9
native
native
native
native
native
native
native
neophyte
native
native
native
native
native
native
native
native
native
native
native
native
native
native
native
archaeophyte
native
native
native
native
native
native
native
native
native
native
native
native
Avenella flexuosa
Vaccinium myrtillus
Urtica dioica
Festuca ovina
Agrostis capillaris
Calamagrostis villosa
Brachypodium pinnatum
Impatiens parviflora
Picea abies
Pinus sylvestris
Quercus petraea agg.
Calamagrostis epigejos
Phalaris arundinacea
Poa angustifolia
P. nemoralis
Aegopodium podagraria
Betula pendula
Calamagrostis arundinacea
Calluna vulgaris
Calystegia sepium
Carex humilis
C. rostrata
Chaerophyllum hirsutum
Cirsium arvense
Fagus sylvatica
Festuca pratensis
F. rubra
Impatiens noli-tangere
Lemna minor
Molinia caerulea s. str.
Nardus stricta
Petasites albus
P. hybridus
Phragmites australis
Potentilla anserina
Quercus robur
Species with the broadest habitat range
The number of habitats in which a species occurs can be used as a measure of the ecological range of that species. This measure is not without problems, because species occurring
in, e.g., five forest habitats probably do not have a broader ecological range than a species
occurring in three habitats, including one forest habitat, one grassland habitat and one
man-made habitat. Still, a comparison of the number of habitats occupied by different species (Fig. 2, Table 2) provides valuable information on the distribution of ecological generalists and specialists among the plants in the Czech flora. Most generalists are native species, less are archaeophytes and least are neophytes. The low number of generalists among
neophytes may reflect, to some extent, the short residence time of these species in this
Sádlo et al.: Habitat plant species pools
315
country, which has prevented them from reaching all the habitats in which they are potentially able to grow. Ecological generalists in the native flora include many grasses, while
the group of alien generalists mainly consists of dicotyledonous herbs.Very similar patterns were found in a previous study based on phytosociological relevés and a different
habitat classification (Chytrý et al. 2005); the similarity of the results of these two studies
indicate the robustness of our estimates of species ecological range.
Native species and grasses are also the most common dominants in different habitats
(Table 3). Among 36 species that are dominant in three or more habitats, there is only one
archaeophyte (Cirsium arvense) and one neophyte (Impatiens parviflora). These 36 species include two genera represented by three species (Calamagrostis and Festuca) and five
genera represented by two species (Impatiens, Petasites, Poa, Carex and Quercus). This
pattern suggests that the ability of a species to become dominant in plant communities depends on phylogenetical relatedness. Most of these species are dominant in less than 15%
of the habitats in which they occur.
Conclusions
This study provides a basic description of the regional species pools for 88 habitats occurring in the Czech Republic. This data set will be used for various studies on the diversity of
Central European flora and vegetation, its historical formation and patterns of plant invasions. In particular, there is great potential for comparative studies when similar databases
become available for other regions of the world (e.g., species assignments to habitats and
phytosociological syntaxa in the German BIOLFLOR database; Klotz et al. 2002). However, a knowledge of regional species pools of different habitats is also important for planning and monitoring of ecological restoration. For example, the same methods as used in the
preparation of the present database were recently used to design seed mixtures for grassland
restoration projects in different regions of the Czech Republic (Blažková et al. 2006).
Acknowledgements
We thank Ilona Knollová for her help with the preparation of the data from the Czech National Phytosociological
Database, Ladislav Mucina and Karel Prach for comments on previous version of the manuscript, Ingolf Kühn,
Stephan Klotz and Gerhard Karrer for their co-operation in the preparation of the habitat classification and Tony
Dixon for improving our English. This research was supported by institutional long-term research plans AV0Z
60050516 (Academy of Sciences of the Czech Republic), MSM 0021622416 (Masaryk University, Brno), MSM
0021620828 (Charles University, Prague), and the European Union within the FP 6 Integrated Project ALARM
(GOCE-CT-2003-506675; Settele et al. 2005) and STREP project DAISIE (SSPI-CT-2003-511202).
Souhrn
Článek shrnuje základní údaje získané z nově vytvořené databáze, která obsahuje tzv. „species pools“ cévnatých
rostlin pro biotopy České republiky, tedy seznamy druhů, které jsou potenciálně schopny růst v daném biotopu.
Pro tento účel jsme použili klasifikaci biotopů do 88 kategorií (appendix 1), které většinou odrážejí fytocenologické jednotky na úrovni svazů nebo skupin svazů. Databáze vychází z údajů obsažených v České národní fytocenologické databázi, které byly kriticky revidovány a rozsáhle doplněny s využitím terénních znalostí o výskytu
druhů v různých biotopech. Databáze zahrnuje 1820 původních druhů, 249 archeofytů a 278 neofytů, z nichž každý je přiřazen k jednomu nebo více biotopům. Kromě údajů o prostém výskytu druhů v jednotlivých biotopech
databáze obsahuje také informace o tom, zda má druh v těchto biotopech svoje ekologické optimum nebo zda se
v nich vyskytuje jako dominanta porostů, případně jako častá dominanta porostů (tab. 1).
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Preslia 79: 303–321, 2007
Největší „species pools“ původních druhů byly zjištěny v suchomilné a teplomilné bylinné i dřevinné vegetaci nižších nadmořských výšek (teplomilné bylinné lesní lemy – 695 druhů, vysoké mezické a suché křoviny – 672
druhů, širokolisté suché trávníky – 549 druhů, perialpidské bazifilní teplomilné doubravy – 535 druhů). Tyto relativně vzácné biotopy jsou podle velikosti „species pools“ následovány hojnými biotopy mezických půd (borové
a modřínové lesní kultury – 525 druhů, mezofilní ovsíkové louky – 505 druhů, dubohabřiny – 481 druhů, pastviny
a parkové trávníky – 474 druhů). Nejmenší „species pools“ (méně než 90 druhů) mají biotopy vodní, slaniskové
a rašeliništní. Velikosti „species pools“ archeofytů a neofytů pro jednotlivé biotopy jsou pozitivně, i když slabě,
korelovány se „species pools“ původních druhů (obr. 1). Biotopy obsahující méně než 350 původních druhů mohou obsahovat různý počet archeofytů. Biotopy s méně než 100 původních druhů obsahují méně než 5, a často
žádné neofyty, zatímco biotopy se 100–350 druhy mohou obsahovat velmi různé počty neofytů. Biotopy s více
než 350 původními druhy vždy obsahují více než 5 archeofytů a více než 5 neofytů (často mnohem více).
Druhy, které se vyskytují jako dominanty porostů, jsou zastoupeny 232 původními druhy, 18 archeofyty a 30
neofyty. Tyto druhy tvoří méně než 3 % ze „species pool“ pro 78 z 88 biotopů. Větší procentická zastoupení (až
14,6 %) dominantních druhů byla zjištěna jen u biotopů s malým „species pool“ a druhově chudými porosty, např.
u vodních, slaniskových a rašeliništních biotopů.
Počet biotopů, v nichž se druh vyskytuje, lze použít jako hrubou míru ekologické amplitudy druhu. Při použití
tohoto kritéria bylo nejvíce druhů se širokou ekologickou amplitudou zjištěno ve skupině původních druhů, méně
mezi archeofyty a nejméně mezi neofyty (obr. 2, tab. 2). Z 36 druhů, které se mohou vyskytovat jako dominanta ve
3 a více biotopech (tab. 3), je 34 druhů původních, jeden je archeofyt (Cirsium arvense) a jeden neofyt (Impatiens
parviflora).
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Received 9 July 2007
Revision received 15 August 2007
Accepted 1 September 2007
Sádlo et al.: Habitat plant species pools
319
Appendix 1. – Habitat classification used in this study and phytosociological syntaxa occurring in particular habitats. Missing items in the sequence of habitat codes indicate habitats that do not occur in the Czech Republic.
1.
1A.
1B.
1C.
1D.
Vegetation of cliffs, screes and walls
Calcareous cliffs (Potentillion caulescentis, Cystopteridion), including rare cases of walls colonized by
natural vegetation of these alliances (e.g. ruins of medieval castles distant from the settlements)
Siliceous cliffs and boulder fields (Agrostion alpinae, Androsacion vandellii, Asplenion serpentini), including rare cases of nutrient-poor acidic mobile screes
Walls (Centrantho-Parietarion), excluding grassy or annual vegetation on wall crowns
Mobile calcareous (to a small extent also siliceous) screes (Stipion calamagrostis)
2.
2A.
2E.
Alpine and subalpine grasslands
Alpine grasslands of siliceous bedrock (Juncion trifidi, Nardo strictae-Caricion bigelowii)
Subalpine tall-grass and tall-forb vegetation (Calamagrostion villosae, Calamagrostion arundinaceae,
Adenostylion alliariae, Dryopterido filicis-maris-Athyrion distentifolii)
3.
3A.
Aquatic vegetation
Macrophyte vegetation of eutrophic and mesotrophic still waters (Lemnion minoris, Utricularion vulgaris,
Hydrocharition, Nymphaeion albae, Magnopotamion, Parvopotamion, Batrachion aquatilis ); rare and occasional occurrences of terrestrial species which do not survive for longer time when flooded are excluded
Macrophyte vegetation of water streams (Batrachion fluitantis)
Vegetation of oligotrophic lakes and pools (Isoëtion lacustris, Littorellion uniflorae, SphagnoUtricularion), including transitions to 4H and 4I
3B.
3C.
4.
4A.
4B.
4C.
4D.
4E.
4F.
4G.
4H.
4I.
4J.
4K.
4L.
5.
5A.
5B.
5C.
5D.
5E.
5F.
5G.
5H.
Wetland and riverine herbaceous vegetation
Reed-beds of eutrophic still waters (Phragmition communis), including Phragmites australis stands in terrestrial habitats, but excluding wet meadows overgrown by Phragmites
Halophilous reed and sedge beds (Scirpion maritimi)
Eutrophic vegetation of muddy substrata (Oenanthion aquaticae)
Riverine reed vegetation (Phalaridion arundinaceae), including ruderalized stands of disturbed stream banks
Reed vegetation of brooks (Sparganio-Glycerion fluitantis)
Mesotrophic vegetation of muddy substrata (Carici-Rumicion hydrolapathi)
Tall-sedge beds (Magnocaricion elatae, Caricion gracilis, Caricion rostratae), including transitions to
6D, 6E and 6F and wetlands with Phalaris arundinacea outside stream banks
Vegetation of low annual hygrophilous herbs (Eleocharition ovatae, Radiolion linoidis, Nanocyperion
flavescentis), including transitions to 4I
Vegetation of nitrophilous annual hygrophilous herbs (Bidention tripartitae, Chenopodion glauci)
River gravel banks (Myricarietum germanicae, Calamagrostietum pseudophragmitis)
Petasites fringes of montane brooks (Petasition officinalis); Petasites stands at low altitudes are assigned
to 13E
Nitrophilous herbaceous fringes of lowland rivers (Senecionion fluviatilis), including ruderalized stands of
disturbed habitats
Vegetation of springs and mires
Hard-water springs with tufa formation (occurrences of Caricion davallianae vegetation on tufa sediments, Lycopodo-Cratoneurion commutati)
Lowland and montane soft-water springs (Cardamino-Montion, Cardaminion amarae)
Alpine and subalpine soft-water springs (Swertio-Anisothecion squarrosi)
Calcareous fens (Caricion davallianae vegetation outside tufa sediments, Cladietum marisci), including
transitions to 6F
Acidic moss-rich fens and peatland meadows (Caricion fuscae, Caricion lasiocarpae, Caricion demissae,
Drepanocladion exannulati, Sphagno warnstorfiani-Tomenthypnion)
Transitional mires (Eriophorion gracilis, Sphagno recurvi-Caricion canescentis)
Raised bogs (Sphagnion medii, Oxycocco-Empetrion hermaphroditi)
Wet peatsoils and bog hollows (Rhynchosporion albae, Leuko-Scheuchzerion palustris), including transitions to 3C and to different types of meadows
320
6.
6A.
6B.
6C.
6D.
6E.
6F.
6G.
Preslia 79: 303–321, 2007
Meadows and mesic pastures
Mesic Arrhenatherum meadows (Arrhenatherion elatioris), including intensively managed mesic
meadows
Montane mesic meadows (Polygono bistortae-Trisetion flavescentis)
Pastures and park grasslands (Cynosurion cristati)
Alluvial meadows of lowland rivers (Deschampsion cespitosae), including intensively managed wet
meadows
Wet Cirsium meadows (Calthion palustris)
Intermittently wet Molinia meadows (Molinion caeruleae)
Vegetation of wet disturbed soils (Ranunculo repentis-Rumicenion crispi)
7.
7A.
7B.
Acidophilous grasslands
Subalpine and montane acidophilous grasslands (Nardion strictae, Nardo strictae-Agrostion tenuis )
Submontane Nardus grasslands (Violion caninae, Nardo strictae-Juncion squarrosi ), including
acidophilous grasslands on forest clearings (Rumici-Avenellion flexuosae)
8.
8A.
8B.
Dry grasslands
Hercynian dry grasslands on rock outcrops (Alysso-Festucion pallentis)
Submediterranean dry grasslands on rock outcrops (Bromo pannonici-Festucion pallentis, Diantho
lumnitzeri-Seslerion)
Narow-leaved sub-continental steppes (Festucion valesiacae)
Broad-leaved dry grasslands (Bromion erecti, Cirsio-Brachypodion pinnati)
Acidophilous dry grasslands (Koelerio-Phleion phleoidis, Hyperico perforati-Scleranthion perennis)
Thermophilous forest fringe vegetation (Geranion sanguinei, Trifolion medii), including some
acidophilous types of herbaceous forest fringes
8C.
8D.
8E.
8F.
9.
9B.
9C.
9D.
9E.
9F.
10.
10G.
10H.
10I.
10J.
Sand grasslands and rock-outcrop vegetation
Open vegetation of acidic sands (Corynephorion canescentis, Thero-Airion)
Festuca grasslands on acidic sands (Armerion elongatae)
Pannonian sand steppes (Festucion vaginatae)
Acidophilous vegetation of spring therophytes and succulents (Arabidopsion thalianae), including transitions to dry grasslands and 13A
Basiphilous vegetation of spring therophytes and succulents (Alysso alyssoidis-Sedion), including transitions to dry grasslands and 13A
Saline vegetation
Continental vegetation of annual halophilous grasses (Cypero-Spergularion salinae)
Inland vegetation of succulent halophytes (Salicornion prostratae)
Inland saline meadows (Juncion gerardii)
Inland saline steppes (Puccinellion limosae)
11. Heathlands and scrub
11A. Dry lowland to alpine heathlands (Euphorbio cyparissiae-Callunion vulgaris, Genisto pilosae-Vaccinion,
Loiseleurio procumbentis-Vaccinion)
11D. Subalpine acidophilous Pinus mugo scrub (Pinion mugo), including rare Pinus mugo scrub with tall forbs
11H. Subalpine deciduous scrub (Salicion silesiacae, Salicetum lapponum)
11I. Willow carrs (Salicion cinereae), including wet scrub of Frangula alnus and Spiraea salicifolia
11J. Willow galleries of loamy and sandy river banks (Salicion albae, Salicion triandrae)
11L. Tall mesic and xeric scrub (Berberidion), including scrub of neophytic Sarothamnus scoparius
11N. Low xeric scrub (Prunion spinosae)
11R. Scrub and pioneer woodland of forests clearings (Sambuco-Salicion capreae), including scrub in ruderal
habitats
12.
12A.
12B.
12C.
Forests
Alder carrs (Alnion glutinosae)
Alluvial forests (Alnion incanae)
Oak-hornbeam forests (Carpinion)
Sádlo et al.: Habitat plant species pools
12D.
12E.
12F.
12G.
12H.
12I.
12J.
12K.
12L.
12O.
12P.
12Q.
12R.
12S.
12T.
12U.
12V.
12W.
321
Ravine forests (Tilio-Acerion), including forests on man-made screes
Herb-rich beech forests (Eu-Fagenion, Acerenion ), including herb-rich fir forests (Galio-Abietenion)
Limestone beech forests (Cephalanthero-Fagion )
Acidophilous beech forests (Luzulo-Fagion), including acidophilous fir forests (e.g. Luzulo pilosaeAbietetum) and Larix decidua forests in the area of its native distribution
Peri-Alpidic basiphilous thermophilous oak forests (Quercion pubescenti-petraeae)
Sub-continental thermophilous oak forests (Aceri tatarici-Quercion, Potentillo albae-Quercetum )
Acidophilous thermophilous oak forests (Sorbo torminalis-Quercetum, Genisto pilosae-Quercetum
petraeae)
Acidophilous oak forests (Genisto germanicae-Quercion)
Boreo-continental pine forests (Dicrano-Pinion), including rare acidophilous pine forests on sand
Peri-Alpidic pine forests (Erico-Pinion)
Peatland pine forests (Vaccinio uliginosi-Pinetum sylvestris)
Peatland birch forests (Sphagno-Betulion pubescentis)
Acidophilous spruce forests (Piceion excelsae)
Basiphilous spruce and fir forests (Athyrio alpestris-Piceion)
Robinia pseudacacia plantations (Robinietea), including plantations of Ailanthus altissima
Plantations of broad-leaved non-native trees (Acer negundo, Juglans nigra, Populus ×canadensis and
Quercus rubra), including stands in parks and gardens
Spruce plantations (Picea abies; rarely also plantations of Picea pungens and Pseudotsuga menziesii)
Pine and larch plantations (Pinus sylvestris, P. nigra, P. strobus and Larix decidua)
13. Anthropogenic vegetation
13A. Annual vegetation of ruderal habitats (Sisymbrion officinalis, Bromo-Hordeion murini, Malvion neglectae,
Salsolion ruthenicae, Eragrostion)
13B. Annual vegetation of arable land (Caucalidion lappulae, Sherardion, Veronico politae-Euphorbion,
Fumario-Euphorbion, Spergulo-Oxalidion, Aphanion, Scleranthion annui, Panico-Setarion, VeronicoTaraxacion)
13C. Annual vegetation of trampled habitats (Polygonion avicularis p. p.)
13D. Perennial thermophilous ruderal vegetation (Onopordion acanthii, Dauco-Melilotion, Arction lappae,
Convolvulo-Agropyrion), including transitions to 6A
13E. Perennial nitrophilous herbaceous vegetation of mesic sites (Galio-Alliarion, Aegopodion podagrariae,
Rumicion alpini), including herbaceous forest fringes on nutrient-rich soils and transitions to 13D
13F. Herbaceous vegetation of forests clearings and Rubus scrub (Carici piluliferae-Epilobion angustifolii,
Atropion bellae-donnae), including transitions to 7B, 11A, 11L, 11R and 13E