Ecology of Closely Related Plant Species. Proceedings of the 40th Symposium of the International...

Stellaria nemorum L. and S. montana Pierrat (Caryophyllaceae) in the Forest Communities ofSloveniaAuthor(s): Igor Dakskobler, Andrej Seliškar and Branko VrešSource: Folia Geobotanica, Vol. 34, No. 1, Ecology of Closely Related Plant Species. Proceedingsof the 40th Symposium of the International Association of Vegetation Science (1999), pp. 115-125Published by: SpringerStable URL: http://www.jstor.org/stable/4201350 .

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Folia Geobotanica 34: 115-125, 1999

STELLARIA NEMORUM L. AND S. MONTANA PIERRAT (CARYOPHYLLACEAE) IN THE FOREST COMMUNITIES OF SLOVENIA

Igor Dakskobler, Andrej Seliskar1) & Branko Vres2)

Institute of Biology, Centre for Scientific Research of the Slovenian Academy of Sciences and Arts, Novi trg 5, SI-1000 Ljubljana, Slovenia; tel. +386 61 1256 068, fax +386 61 1257 797, E-mail 1) [email protected]; 2) [email protected]

Keywords: Distribution, Phytosociology, Stellaria nemorum group, Taxonomy, Forest vegetation

Abstract: Morphological and coenological differences between closely related species Stellaria nemorum L. and S. montana PIERRAT are described. Stellaria montana is frequent in Slovenia and occurs mostly in the montane belt (altitude 600-1200 m). It has been recorded in thirty two forest communities (associations or lower syntaxa). Although widely distributed in Europe S. nemorum is more rare in Slovenia. It occurs mostly in upper-montane (altimontane) and subalpine belt (altitude 1200-1600 m) in ten forest or shrubby communities. Both, S. nemorum and S. montana are valuable diagnostic species for certain associations. They characterize sites with fresh soil, rich in nitrogen. As an example we present two syntaxa of beech forests of Slovenia, in which the species of the S. nemorum group have high constancy and cover value.

INTRODUCTION

In the present paper, two closely related species, Stellaria nemorum L. s. str. and S. montana PIERRAT, are discussed in connection with their relations to the forest communities of Slovenia. The aim of this work has been to search for new morphological characteristics of both species, their chorology and the relation of these plants in two beech forest communities, using the BRAUN-BLANQUET (1964) and MATUSZKIEWICZ & MATUSZKIEWICZ (1981) approach.

Stellaria nemorum was first described by LINNAEUS (1753). In 1880, the related species Stellaria montana PIERRAT was described (PIERRAT 1880), and later (MURBECK 1891) the subspecies Stellaria nemorum subsp. glochidisperma MURB. (= S. glochidisperma (MURB.) FREYN). In the past, they were subject to changes in their names as well as taxonomic rank (FREYN 1892, HEGI 1911, PETERSON 1936). Recent reports (GREEN 1954, ANDREAS 1956,

KOOPMANS & SLIM 1968, VRES 1991, BABIJ et al. 1997) show that both S. glochidisperma and S. montana represent the same taxon, which should be strictly distinguished from S. nemorum L. s. str. In the rank of species, the oldest valid name is Stellaria montana PIERRAT (= S. nemorum subsp. montana (PIERRAT) BERHER).

In Slovenia, recent chorological data show that Stellaria montana is more frequent than S. nemorum (Fig. 1). In contrast, a different situation has been observed in relation to their whole distribution range in Europe. Throughout Europe, S. montana occurs rarely and is

concentrated in Denmark and on the Balkan peninsula, being more or less dispersed elsewhere (JALAS & SUOMINEN 1983). On the other hand, S. nemorum is a widely spread species.

According to the herbarium (LJU, LJM), floristic and phytosociological data (e.g. MARIN*EK & DAKSKOBLER 1988, MARINfEK et al. 1989, VRES 199 1, ZUPANcIK' 1980), Stellaria



116 I. Dakskobler et al.

46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65

91 15 4 _ -7 = |||J7

92 L 93 94

95

96

97 1; 0 ! 9- ; > kt S S~~L; VENIA

0 Stellafia mrntana

04

05 141_ __7_ __ __

Fig. 1. The distribution of Stellaria nemorum L. and S. montana PIERRAT in Slovenia, according to the mapping of flora of Central Europe.

nemorum is considerably rare in Slovenia. It has been recorded in forest communities of montane, upper-montane, and subalpine belt (600 to 1600 m), mostly at altitudes between 1200 and 1600 (-1800) m. The altitudinal range of the forest communities in which S. montana occurs extends from the submontane to the upper-montane belt (200 to 1400 m), the median being in the montane belt (600 to 1200 m) (Fig. 2).

MORPHOLOGICAL DIFFERENCES BETWEEN STELLARIA NEMORUM S. STR. AND S. MONTANA

In Slovenia, the morphological features of the Stellaria nemorum group were studied during the previous ten years. The specimens of S. nemorum and S. montana from herbarium LJU and LJM were examined and measured (VRES 1991). The species usually differ clearly in their habitus (Fig. 3) and some morphological characters (Tab. 1). The most important differentiating characters are the ratio between the length and the width of the leaf at the node below the first branching of inflorescence, the length and pubescence of the sepals, the length of the papillae on ripe seeds and the presence or absence of hooks on the papillae of ripe seeds. Both species can be satisfactorily distinguished on the basis of the combination of these features. In Stellaria nemorum, the leaf at the node under the first branching of inflorescence is sessile, ovate, truncate to angustate at base, (1.7-)2-2.6(-3.2) times longer than wide (2.4 ? 0.3 times in average). In S. montana, leaves are stalked, cordate with a cordate basis, and only (1.3-)1.6-2(-2.4) times longer than wide (1.8 ? 0.2 times in average). In S. montana, the bracts decrease abruptly in size after the first dichotomy (the second pair is usually less than 1/3 as long as the first), but in S. nemorum they decrease gradually (the second pair is usually almost 1/2 as long as the first). The sepals of flowers are slightly longer in S. nemorum (5.6-7 mm), with a nude apex (pubescent mostly only at the base), compared to S. montana (4.6-5.5 mm), which has a pubescent apex. The ripe seeds are covered with cylindrical or hemispherical papillae. In addition, the two species differ from each other in the length, the width and the shape of papillae (Fig. 4). In S. nemorum, the seeds are furnished



Stellaria nemorum and S. montana in the forest communities of Slovenia 117

2001-2100 1901-2000 1801-1900 * S. nmom 1701-1800 O S. montana 1601-1700

1501-1600

1401-1500

1301-1400

1201-1300

1101-1200

1001-1100

901-1000 801-900

701-800

601-700 501-600

401-500

301-400 201-300 101-200

0-100

-25.0 -20.0 -15.0 -10.0 -5.0 0.0 5.0 10.0 15.0 20.0

Frequency (%/6)

Fig. 2. The distribution frequency of Stellaria montana and S. nemorum s. str. in Slovenia in correlation to altitude (data sources: literature, herbarium data). n=35 (S. montana), n=67 (S. nemorum).

with hemispherical unarmed papillae, being (0.3-)0.6-1.5(-3) times as long as wide (1.0 ? 0.4 times in average), whereas in S. montana the seeds have long cylindrical papillae with barbate caps, being (1.2-)2-4(-12) times longer than wide (3.8 ? 1.4 times in average).

THE COMPARISON OF BOTH SPECIES AFFINITY TO VEGETATION TYPES

Characteristic habitats of S. montana are in communities with Fraxinus excelsior, Acer pseudoplatanus, and Ulmus glabra. It is an important diagnostic species for the association Hacquetio-Fraxinetum MARINtEK in WALLNOFERet al. 1993 and for the suballiance Polysticho setiferi-Acerenion pseudoplatani BORHIDI et KEVEY 1996. This species also grows in humid forms of the montane and upper-montane beech, fir-beech, and maple-beech forests. It can be found in fir forests on colluvial soil, in humid moderately acidophilic fir-beech forests, or in the moderate freezing dolines and medium acidophilic spruce forests of the Dinaric phytogeographic region. It is very scarce in hornbeam forests and submontane beech forests.

Stellaria nemorum is recorded in the forest communities of the montane, upper-montane, and subalpine belt but mostly thrives in the subalpine maple-beech forests or in the more humid upper-montane and subalpine beech communities. In the Alpine and pre-Alpine phytogeographical region of Slovenia, it grows in the subalpine spruce forests or in shrubby communities of Alnus viridis on moderately fresh and acid soil.

THE PRESENCE OF STELLARIA NEMORUM AND STELLARIA MONTANA IN THE FOREST COMMUNITIES OF SLOVENIA

Generally, the Stellaria nemorum group most frequently thrives in the communities of the order Fagetalia sylvaticae PAWLOWSKI et al. 1928. Stellaria nemorum s. str. is classified as a characteristic species of the alliance of the hygrophilic communities of central Europe (Alnion incanae PAwLowsKI et al. 1928 (=Alno-Ulmion BR.-BL. et R. Tx. 1943), e.g.



118 1. Dakskobler et al.

Table 1. Mean, standard deviation, range (in brackets), in mm, number of observations of morphological characters of Stellaria nemorum L. s.str. and S. montana PIERRAT in Slovenia; all differences among samples are highly significant (P < 0.001).

Variable Stellaria nemorum S. montana

Length of the first stem leaf under branching of inflorescence 66.7?17.5 (27-115); n=93 47.5?15.6 (11-90); n=95

Length of the second stem leaf under branching of inflorescence 66.8?19.4 (32-117); n=89 47.5?13.4 (17-82); n=95

Length of peduncle of the first stem leaf under branching of inflorescence 0.6?1.8 (0-10); n=93 18.7+9.6 (2-50); n=94

Length of peduncle of the second stem leaf under branching of inflorescence 6.9?8.1 (0-31); n=89 29.9?9.8 (10-60); n=95

Length of bracts in the first branching of inflorescence 53.8?16.2 (15-92); n=90 43.3?16.1 (6-85); n=95

Length of bracts in the second branching of inflorescence 30.5?12.2 (4-64); n=185 13.0?8.0 (1.7-36); n=156

Length of bracts in the third branching of inflorescence 13.4?7.5 (2.5-42); n=303 3.4?2.0 (1-10); n=86

Petal limb length 11.0?0.7 (9-12); n=59 11.8?1.5 (7.5-15); n=101

Calyx length 6.1?0.6 (4.5-7.8); n=535 4.9+0.5 (3.5-7); n=562

Fruit length 8.5?1.0 (6-11); n=260 7.4?1.2 (4.5-10); n=284

Peduncle length 24.2?6.3 (10-50); n=403 19.5?7.7 (5-55); n=469

Papillae length 0.06?0.02 (0.03-0.13); n=651 0.10?0.02 (0.04-0.15); n=752

Papillae width 0.06?0.01 (0.03-0.10); n=595 0.03?0.01 (0.01-0.07); n=685

OBERDORFER (1983: 370-371), ELLENBERG (1991: 145), WALLNOFER et al. (1993: 91)). Stellaria montana has, due to its considerable distribution in south-eastern Europe, certain diagnostic value in mesophilic beech communities (alliance Aremonio-Fagion BORHIDI in TOROK et al. 1989), and especially in the communities of sycamore maple (suballiance Polysticho setiferi-Acerenion BORHIDI et KEVEY 1996) - BORHIDI & KEVEY (1996: 114). Stellaria nemorum s. str. mostly thrives in similar communities both in Central and western Europe (e.g. ACCEITO 1991, Tab. 2).

The presence of both species in the forest communities of Slovenia is represented in Tab. 2. Sources for the table were published phytosociological data. The forest associations were grouped on the basis of floristic and ecological similarity (columns 1 and 2). Authors of the sources are stated in the column 3. The next two columns specify the frequency (Fr) of each species (0-100%), except when there exists only one releve or synoptic table published, and the presence degree class (Cl) (I-V). The importance of each species (Im) for alliances and suballiances is indicated by values 1-5 (1 means rare and diagnostically unimportant, 5 means frequent and characteristic for group of communities).

It was not possible to verify completely the reliability of the determination of both taxa of the Stellaria nemorum group in the phytosociological material applied. The herbarium material, gathered in different regions of Slovenia and in the stands of numerous syntaxa, indicates that the determination was generally correct. We established false determination with certainty




iomm momm

Fig. 3. Habitus of Stellaria montana (a) and Stellaria nemorum (b) (del. B. VRES).

only in two cases, considering the corrigenda in the Tab. 2. It is possible that both taxa of the S. nemorum group occur together in the stands of some communities, thriving in the upper-montane belt, while the authors identified only one. In the synoptic table (Tab. 3) there is an example of two beech communities in Slovenia, in which the species of the Stellaria nemorum group grow optimally.

The first one (Tab. 3, column 1) is classified in the association Lamio orvalae-Fagetum (HORVAT 1938) BORHIDI 1963. The stands of this community are located in the area of Kalski gozd (9948/2, UTM VMO0), a rather extensive forest complex in the northern part of the Banjsice plateau in western Slovenia. They thrive on diversified rocky High Karst surface, mostly on sinkhole edges at the altitude of 800 to 1050 m. The parent material is Jurassic and partly Cretaceous limestone. The prevailing soil type is brown calcareous soil (Chromic Cambisols). Stellaria montana has high constancy and cover value in herb layer of these stands (see also DAKSKOBLER 1996a: 31-34).

As an example of beech forest, where both species of the S. nemorum group treated thrive, we present upper-montane beech community, which is classified in the association Ranunculo platanifolii-Fagetum MARIN-EK et al. 1993 (Tab. 3, column 2). The stands of this community have been found in the south-eastern part of the Julian Alps, mainly on the north-western slopes of Porezen (9849/2, UTM VM21) in the Baca Valley. They thrive in the upper-montane belt, up to the border of the subalpine belt (altitude 1200-1450 m). Parent material is Cretaceous



Table 2. The presence of Stellaria nemorum and S. montana in the forest communities of Slovenia. Fr. - frequency; Cl. - class, Im. - Importance of species.

Alliance, suballiance, association; geographical variant, subassociation References S. montana S. nemorum Fr. Cl. Im. Fr. Cl. Im.

Tilio-Acerion KLIKA 1955 4 Hacquetio-Fraxinetum MARINCEK in WALLNOFER et al. 1993; var. geogr. Dentaria pentaphyllos MARINCEK (1990) 61 IV Hacquetio-Fraxinetum MARINCEK in WALLNOFER et al. 1993; var. geogr. Dentaria pentaphyllos, var. Carpinus betulus MARINCEK (1995b) 13 1 Acer pseudoplatanus-Ulmus glabra ass. (nom. prov.) MARINCEK (1995a) 100 V Corydalo cavae-Aceretum pseudoplatani MOOR 1938; var. geogr. Dentaria enneaphyllos ZUPANCIC (1996) 100 V

Erythronio-Carpinion (HORVAT 1958) MARINCEK in WALLNOFER et al. 1993 1 Pruno padi-Carpinetum betuli MARINCEK 1994 MARINCEK & ZUPANCIC (1984) 1 1 I

Lamio orvalae-Fagenion BORHIDI ex MARINeEK et al. 1993 3 Lamio orvalae-Fagetum (HORVAT 1938) BORHIDI 1963; var. geogr. Dentaria pentaphyllos DAKSKOBLER (1996a) 4 I Lamio orvalae-Fagetum (HORVAT 1938) BORHIDI 1963; var. geogr. Dentaria pentaphyllos, stellarietosum montanae DAKSKOBLER - Tab. 3, col. I 100 V Lamio orvalae-Fagetum (HORVAT 1938) BORHIDI 1963; var. geogr. Dentaria polyphyllos MARINCEK et al. (1983) 10 I Omphalodo-Fagetum (TREGUBOV 1957) MARINCEK et al. 1993; aceretosum PUNCER et al. (1974) 60 III Omphalodo-Fagetum (TREGUBOV 1957) MARINCEK et al. 1993; aceretosum facies Petasites albus PUNCER (1980) 80 IV Omphalodo-Fagetum (TREGUBOV 1957) MARINCEK et al. 1993; typicum PUNCER (1980) 8 I Omphalodo-Fagetum (TREGUBOV 1957) MARINCEK et al. 1993; galietosum odorati, var. Hacquetia epipactis PUNCER (1980) 6 I Omphalodo-Fagetum (TREGUBOV 1957) MARINCEK et al. 1993; thelypteretosum limbospermae AccErro (1978) 18 I Cardamini savensi-Fagetum KoAm 1962 KotIR (1979) 19 1 Cardamini savensi-Fagetum KoaIR 1962; var. geogr. Abies alba KOtIR (1979) 41 III Isopyro-Fagetum KOaIR 1962; var. geogr. Arum maculatum KoaIR (1979) 1 1 I Isopyro-Fagetum Ko0IR 1962; var. geogr. Adenostyles alliariae KoSIR (1979) 31 II

Saxifrago rotundifoliae-Fagenion MARINCEK et al. 1993 3 3 Homogyno sylvestris-Fagetum MARINtEK et al. 1993; var. Stellaria montana MARINCEK & DAKSKOBLER (1988) 100 V Ranunculo platanifolii-Fagetum MARIN?EK et al. 1993; var. geogr. Hepatica nobilis MARINCEK et al. (1989) I Ranunculo platanifolii-Fagetum MARINEEK et al. 1993; var. geogr. Hepatica nobilis, cortusetosum DAKSKOBLER (1996b) 25 II 12 I Ranunculo platanifolii-Fagetum MARIN?EK et al. 1993 DAKSKOBLER - Tab. 3, col. 2 54 III 100 V Stellario montanae-Fagetum (ZUPANCIC) MARINCEK et al. 1993 ZUPANdIC (1967, 1969) V I Knautio drymeiae-Fagetum ZUPANCIC (1969) 1994 ZUPANCIC (1967, 1969) V Aconito paniculati-Fagetum (ZUPANdIP 1969) MARINCEK et al. 1993 ZUPANCIC (1969) V Polysticho lonchitis-Fagetum (HORVAT 1938) MARINCEK in POLDINI et NARDINI 1993; var. geogr. Salix waldsteniana MARINCEK (1980) 40 II Polysticho lonchitis-Fagetum (HORVAT 1938) MARINCEK in POLDINI et NARDINI 1993; var. geogr. Allium victorialis MARINCEK (1996) 21 II

Luzulo-Fagenion LOHM. et R. Tx. 1954 1 Luzulo-Fagetum MEUSEL 1937; var. geogr. Cardamine trifolia, abietetosum MARINCEK & DAKSKOBLER (1988) 3 I Luzulo-Fagetum MEUSEL 1937; var. geogr. Cardamine trifolia, abietetosum, var. Galium rotundifolium MARINCEK (1995a) 10 I

Vaccinio-Piceion BR.-BL. 1939 2 2 Dryopterido (pseudomas)-Abietetum KotIR 1994 KoaIR (1994) Polysticho setiferi-Abietetum KoaIR 1994 KoaIR (1994) 25 A Stellario montanae-Piceetum ZUPANCIC (1976) 1994 ZUPANCIC (1980, 1994) 57 O Hacquetio-Piceetum ZUPAN-IC (1976) 1994 ZUPANcIC (1980, 1994) 5 v Luzulo sylvaticae-Piceetum M. WRABER 1963 WRABER (1963) 22 II CD

CD Alnion viridis AICHINGER 1933 4 Alnetum viridis BR.-BL. 1918 PISKERNIK (1982) 73 IV




V..~~~~~~~~

a 01 mmI b

Fig. 4. The form and papillae length: Stellaria montana (a), S. nemorum (b) (del. B. VRES).

platy limestone with admixture of red marl and chert. The soils are fresh, humose, loamy (brown calcareous soil and eutric brown soil). These beech stands are characterized by high cover of the species of the Stellaria nemorum group. Stellaria montana predominates at the altitude of 1200 to 1350 m, whereas above 1350 m, in the transition area to the subalpine beech stands of the association Polysticho lonchitis-Fagetum (HORVAT 1983) MARINO-EK in POLDINI et NARDINI 1993, Stellaria nemorum is dominant. In one releve an intermediate form between both related species was reported. However, due to a lack of readily available plant material it can only be analyzed in the future.

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Encl. Appendix pp. 124-125




APPENDIX

Table 3. Synoptic table of two beech communities from westem Slovenia in which there is a growing optimum of the species of the Stellaria nemorum group. 1 - Lamio orvalae-Fagetwn (HORVAT 1938) BORHIDI 1963; 2- Ranunculo platanifolii-Fagetum MARINCEK et al. 1993. Nomenclature of vascular plants in this paper follows TRPrN & VRED (1995) and that of mosses DOLL (1991).

Association number 1 2 Association number 1 2 Number of releves 18 13 Number of relev6s 18 13

Stellaria montana 100 54 Fraxinus excelsior E2 17 Stellaria nemorum . 100 Fraxinus excelsior Ei 72 Stellaria nemorum x S. montana (?) . 8 Mercurialis perennis 72 15

Phyllitis scolopendrium 72 Aremonio-Fagion Cardamine impatiens 72 31 Lamium orvala 100 38 Dentaria pentaphyllos 67 Dentaria enneaphyllos 100 69 Pulnonaria officinalis 61 31 Helleborus odorus 89 Polystichum xluerssenii Cardamine trifolia 61 85 (= P braunii x P aculeatum) 56 23 Cyclamen purpurascens 39 . Veronica montana 56 Aremonia agrimonoides 28 . Corydalis solida 50 8 Isopyrum thalictroides 28 Sanicula europaea 50 8 Rhamnus fallax E2 17 . Carex sylvatica 50 8 Anemone trifolia 6 . Symphytum tuberosum 44 92 Primula vulgaris 6 15 Lathyrus vemus 44 8 Polystichum setiferum . 8 Festuca altissima 39 8

Neottia nidus-avis 28 31 Saxifrago rotundifoliae-Fagenion Prunus avium E3 6 Adenostyles glabra 11 85 Prunus avium El 22 Saxifraga rotundifolia 11 85 Melica nutans 17 Polygonatum verticillatum . 100 Brachypodium sylvaticum 17 Adenostyles alliariae . 77 Asarum europaeum subsp. caucasicum 17 Luzula sylvatica subsp. sylvatica . 38 Salvia glutinosa 17 Cicerbita alpina 15 Circaea intermedia 17 Ranunculus platanifolius . 15 Stachys sylvatica 11

Euporbia dulcis 11 Fagetalia sylvaticae Dryopteris affinis subsp. borreri 11 8 Fagus sylvatica E3 100 100 Galium laevigatum 11 38 Fagus sylvatica E2 72 100 Campanula trachelium 6 31 Fagus sylvatica El 78 8 Prenanthes purpurea 6 31 Dentaria bulbifera 100 62 Ranunculus lanuginosus 6 62 Corydalis cava 100 69 Poa nemoralis 6 31 Dryopteris filix-mas 100 92 Lilium martagon . 92 Arum maculatum 100 . Leucojum vernum . 31 Galium odoratum 100 15 Polystichum aculeatum 100 77 Querco-Fagetea Adoxa moschatellina 100 77 Anemone nemorosa 100 100 Geranium robertianum 100 38 Camptothecium lutescens 89 31 Paris quadrifolia 100 100 Isothecium alopecuroides 89 54 Circaea lutetiana 94 8 Ctenidium molluscum 83 54 Myosotis sylvatica agg. 94 46 Gagea lutea 67 31 Scrophularia nodosa 94 69 Anemone ranunculoides 56 69 Galeobdolon flavidum 94 85 Lonicera xylosteum E2 33 Milium effusum 94 62 Corylus avellana 33 Epilobium montanum 94 62 Vinca minor 28 Actaea spicata 94 85 Clematis vitalba E2 28 Epipactis helleborine 94 . Hypericum montanum 17 Mycelis muralis 94 31 Crocus napolitanus (= C. vittatus) 17 Sambucus nigra E2 89 . Sesleria autumnalis 17 Viola reichenbachiana 89 . Moehringia trinervia 11 8 Daphne mezerum E2 89 8 Arabis turrita 11 Chrysosplenium alternifolium 78 31 Platanthera bifolia 11 Polygonatum multiflorum 78 . Hepatica nobilis 11 Acer pseudoplatanus E3 6 62 Carex digitata 11 Acer pseudoplatanus E2 39 46 Calamintha sylvatica 11 Acer pseudoplatanus El 78 62 Moehringia muscosa 11 Fraxinus excelsior E3 6 . Galanthus nivalis 6 8




Association number 1 2 Association number 1 2

Cirsium erisithales 31 Polypodium vulgare 50 8 Laburnum alpinum E2 15 Cymbalaria muralis 28

Sedum hispanicum 17 Vaccinio-Piceetea Asplenium viride 46 Oxalis acetosella 89 77 Luzula luzuloides 83 85 Other species Gymnocarpium dryopteris 56 38 Galeopsis pubescens 83 Dryopteris expansa 56 15 Rubus hirtus E2 61 Hypnum cupressiforme 39 31 Cardamine flexuosa 56 Dryopteris carthusiana 28 8 Galeopsis speciosa 50 54 Polytrichum formosum 28 8 Rubus ideus E2 44 54 Maianthemum bifolium 22 Solanum dulcamara 39 Gentiana asclepiadea 17 62 Sambucus racemosa E2 33 8 Luzula luzulina 11 8 Atropa bella-donna 33 Saxifraga cuneifolia 11 Sorbus aucuparia E2 17 23 Picea abies E3 . - 38 Sorbus aucuparia Ei 28 Picea abies E2 11 8 Fragaria vesca 28 Abies alba E3 8 Verbascum lanatum 17 Abies alba E2 11 Deschampsia cespitosa .17 46 Atrichum undulatum 11 8 Galeopsis tetrahit 11 Thelypteris phegopteris 6 8 Bromus ramosus 11 Dryopteris dilatata 85 Dactylis glomerata 6 8 Aposerisfoetida 69 Hypericwn hirsutum 38 Calamagrostis arundinacea 54 Polystichum xillyricum 23 Veronica urticifolia 54 Dactylorhiza maculata s.l. 15 Valeriana tripteris 23 Lamium maculatum 15 Thelypteris limbosperma 15 Crocus albiflorus 15 Vaccinium myrtillus 15 Poa alpina 15.

Betonica alopecuros 15 Adenostyletalia Senecio ovatus 100 100 Mosses and lichens Urtica dioica 100 46 Grimmia pulvinata 94 54 Athyrium filix-femina 83 100 Brachythecium velutinum 89 31 Impatiens noli-tangere 50 31 Plagiochila asplenioides 67 46 Scrophularia vemalis 39 8 Collema sp. 50 8 Angelica sylvestris 28 46 Neckera crispa 44 Ribes alpinum E2 17 Plagiothecium denticulatum 39 23 Veratrum album subsp. lobelianum 11 92 Plagiomnium affine 39 Thalictrum aquilegiifolium 62 Thamnobryum alopecurum 39 Phyteuma ovatum 62 Brachytheciun rutabulum 39 31 Chaerophyllum hirsutum subsp. hirsutum 54 Mnium marginatum 39 31 Rumex arifolius 46 Mniun sp. 33 8 Aconitum vulparia 38 Madotheca platyphylla 33 Doronicum austriacum 38 Anomodon viticulosus 28 Carduus personata 38 Plagiomnium undulatum 28 8 Alnus viridis 31 Bryum capillare 22 23 Myrrhis odorata 23 Conocephalum conicum 22 38 Geum rivale 23 Plagiomnium cuspidatum 22 23 Crepis paludosa 15 Peltigera canina 17 8 Melandrium rubrun 15 Cladonia sp. 11 15 Salix appendiculata E2 15 Metzgeria sp. 11 Chaerophyllum hirsutum subsp. villarsii 15 Anomodon attenuatus 11 8

Fissidens cristatus 6 46 Asplenietea trichomanis Plagiothecium sylvaticum 6 31 Cystopteris fragilis 78 77 Paraleucobryum sauteri 46 Asplenium trichomanes 72 23 Tortella tortuosa 38



Ecology of Closely Related Plant Species. Proceedings of the 40th Symposium of the International...

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