Journal of Vegetation Science 19: 331-342, 2008
doi: 10.3170/2008-8-18372, published online 24 January 2008
© IAVS; Opulus Press Uppsala.
331
Classification and phytogeographical differentiation of broad-leaved ravine forests in southeastern Europe
Kosir, Petra1*; Carni, Andraz12 & Di Pietro, Romeo3
^Institute of Biology, Scientific Research Centre of the Slovenian Academy of Sciences and Arts, Novi trg 2, P. B. 306,
SI-1001 Ljubljana, Slovenia; 2E-mail carni@zrc-sazu.si; 3 Department ITACA, University of Rome 'La Sapienza', Via Flaminia 70, IT-00196 Rome, Italy; E-mail romeo.dipietro@uniromal.it; "Corresponding author; E-mailpetrako@zrc-sazu.si
Abstract
Question: How do broad-leaved ravine forests in SE Europe differentiate phytogeographically? Do they differ from analogous European forests? What is their distribution pattern? Location: southeastern Europe, Apennine-Balkan province. Methods: The initial data set of 2189 releves was stratified geographically andphytosociologically; 614releves remaining after stratification were classified with aTWINSPAN and cluster analysis, wich resulted in four clusters and eight subclusters. Average Pignatti indicator values for releves of each subcluster were subj ected to PCA to show ecological relationships among the clusters. The spectra of geoelements and sociological species groups of individual subclusters were calculated to show phytogeographical and sociological relationships between them. The diagnostic species combination was calculated by a fidelity measure (0-coefficient) and presented in a synoptic table. Results: Broad-leaved ravine forests in southeastern Europe form a separate group within the European broad-leaved ravine forests. They are well differentiated by the species with a southeast European distribution, as well as by many other species that reflect their different ecological affinities. Conclusions: The phytosociological and phytogeographical relationships between the Apennines and the Balkan peninsula that have already been recognized for other vegetation types have been confirmed for broad-leaved ravine forests. According to the numerical analysis, two suballiances of broad-leaved ravine forests in southeastern Europe are proposed, both belonging to the alliance Tilio-Acerion: an amphi-Adriatic xerothermo-philous suballiance Ostryo-Tilienion platyphylli suball. nova andamesophilous suballiance Lamio orvalae-Acerenion suball. nova, the latter appearing only on the Balkan Peninsula.
Keywords: Acer; Apennines; Balkan Peninsula; Biogeogra-phy; Fidelity; Fraxinus; Numerical analysis; Phytosociology; Syntaxonomy; Tilia; Ulmus.
Abbreviation: BLRF = Broad-leaved ravine forest.
Nomenclature: Tutin et al. (1964-1980); except Stellaria montana Pierrat and Dryopteris affinis (Lowe) Fraser-Jenkins. Fagus moesiaca is included in Fagus sylvatica; syntaxonomy follows Mucina et al. (1993), except for the syntaxa under consideration. New names are based on the nomenclature rules in Weber etal. (2000).
Introduction
Broad-leaved ravine forests (BLRFs) grow on spatially restricted sites with specific soil conditions. They occur on slopes, on the foot of slopes, in sinkholes, gorges and hollows with colluvial, skeletal and primarily unstable soil, which allow the broad-leaved trees Acer platanoides,A.pseudoplatanus, Fraxinus excelsior, Tilia cordata, T. platyphyllos and Ulmus glabra to replace otherwise competitively stronger tree species, above all Fagus sylvatica (Clot 1990; Muller 1992; Mucina et al. 1993; Ellenberg 1996).
BLRFs have already been thoroughly studied in many parts of Europe. In central Europe they are classified into the Tilio-Acerion alliance, within which two ecological groups of associations are distinguished, recognized as suballiances by some authors (Muller 1992; Willner 1996): a group of mesophilous Acer associations and a group of xerothermophilous associations with Tilia and Corylus (Mucina et al. 1993; Borhidi 2003). Forests of both groups of associations are found on sites with similar soils, but they differ in temperature and moisture requirements. There have been some locally focused publications on BLRFs in southeastern Europe, but there is still is no synthetic review of these forests in the region available.
We focus on southeastern Europe (Apennine-Balkan province, Fig. 1), the distribution area of zonal Carpinus forests of the alliance Erythronio-Carpinion and Fagus forests of the alliance Aremonio-Fagion, both of the order Fagetalia sylvaticae, and of fhermophilous forests of the alliance Carpinion orientalis of the order Quercetalia pubescentis. All of these communities are very rich in species and occur on both sides of the Adriatic Sea in the Apennines and the Balkans. They are characterized by numerous relict and endemic species that survived Quaternary glaciations in southern European refugia (Bennet et al. 1991; Trinajstic 1992; Tzedakis 1993; Magri 1998; Petit et al. 2002).
It has already been established that mesophilous
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deciduous forests of southeastern Europe differ from forests in central Europe, and vicariant alliances have been described. The southeastern European alliances Aremonio-Fagion and Erythronio-Carpinion are vicariant to the central European alliances Fagion sylvaticae and Carpinion betuli within the order Fagetalia sylvaticae. In that respect, the question of phytogeographic differentiation within European BLRFs is raised. Should southeastern European BLRFs be classified like zonal vegetation of Fagus- and Carpinus forests in a vicariant alliance regarding central European forests? This paper addresses this question by reviewing and discussing BLRFs in southeastern Europe, above all their floristic composition, chorology, syntaxonomy and phytogeo-graphical differentiation in comparison with the central European BLRFs.
Methods
Forest vegetation relevés made by applying the Braun-Blanquet approach (Braun-Blanquet 1964), classified by their authors as broad-leaved ravine forests and/or dominated by broad-leaved species, and originating from SE and C Europe (Fig. 1), were collected from the literature (n = 2636). Relevés were entered into the TURBOVEG (Hen-nekens & Schaminée 2001) database. The relevés with an incomplete list of herb species indicated by the authors were not included into the analyses. With regard to the definition of BLRFs by dot (1989), we excluded the relevés whose dominant tree species (cover value 4 and 5) are species of climazonal and other forest types (Abies alba, Alnus glutinosa, A. incana, Carpinus betulus, Castanea saliva, Fagus sylvatica, Fraxinus ornus, Ostrya carpinifolia, Picea abies, Quercus cerris, Q. ilex, Q.petraea, Q.pubescens and Q. robur), as well as those where none of the tree species characteristic of BLRFs (Acer platanoides, A. pseudopla-tanus, Fraxinus excelsior, Tilia cordata, T. platyphyllos and Ulmus glabra) had a cover value of at least 2 (Chytrý et al. 2002a). There were 2189 BLRF relevés remaining after this selection.
This data set of 2189 relevés was then stratified. Stratified resampling was made by combining the geographical stratification with stratification by phytosocio-logical association (Knollová et al. 2005). This means up to ten relevés of one association in one area were selected in such a way that different authors, different publications and different locations within the area were represented. We took the biogeographic map of Europe (Rivas-Martinez et al. 2004) as the basis of geographical stratification, where sectors were used as geographical strata. The associations were defined according to synthetic works (Clot 1990; Mucina et al. 1993; Willner 1996) or expert assignments in case they were not treated
in any of those synthetic works. Syntaxa described in older publications under the names Aceri-Fraxinetum, Aceri-Tilietum and Tilio-Fraxinetum were considered separately according to the author and publication. After stratification, 614 releves remained (App. 1), originating from 87 combined geographic-habitat strata.
As many authors did not record mosses, we excluded them from our analysis before numerical processing. For the purpose of numerical analysis and in the synoptic table we unified the system of layer division, which differs from author to author. All sublayers of the tree layer were incorporated into one, whereas the herb and scrub layers, where scrub species, tree saplings, seedlings and lianas occur, were united into one scrub layer.
Fig. 1. The study area on the Biogeographical map of Europe (Rivas-Martmez et al. 2004). Legend: The Apennine-Balkan province (9; shaded) is divided into the Apennine (9a), Pada-nian (9b), Illyrian (9c), Hndan (9d) andBulgarian (9e) sectors. Neighbouring provinces and sectors include the central European province (5) with the middle European sector (5b), Alpine province (8) with the western Alpine (8b), central Alpine (8c) and Eastern Alpine (8d) sectors, Pannonio-Carpathian province (10) with the Pannonian (10a) and Carpathian (10b) sectors, Italo-Thyrrhenian (20), Adriatic (21) and Graeco-Aegean (22) province. Broad-leaved ravine forest suballiances distinguished in this paper: A.Ostryo-Tilienion, ♦ Lamio orvalae-Acerenion, • Tilienion, # Lunario-Acerenion.
- Classification and phytogeographical differentiation of broad-leaved forests -
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When processing and analysing the BLRF relevés of SE and C Europe we carried out a TWINSPAN classification (Hill 1979), run under the JUICE 6.4 program (Tichý 2002). TWINSPANpseudospecies cutlevels for species abundances were set to 0 -2 -5 -10 -20% scale units as proposed by McCune & Grace (2002). Initially, six division levels were chosen and the minimum group size for division was set to five relevés. As we were investigating classification of BLRFs on the level of superior syntaxa, the level of division was raised until groups of relevés still interpretable regarding ecology and phytogeography were obtained. Three levels of division were accepted, resulting in eight groups of relevés.
As TWINSPAN cannot perform a fully hierarchically organized classification tree (it does not consider cluster heterogeneity), the relationships between the resulting eight TWINSPAN groups were further investigated by cluster analysis, using the SYN-TAX program (Podani 2001). Percentage frequencies of species occurrences (constancies) in the eight TWINSPAN groups were used as input data for cluster analysis. We used several methods: /3-flexible with parameter j3 = -0.25, group average (UPGMA), complete link (farthest neighbour) method with similarity ratio as the resemblance measure, all leading to similar results. Only the result of the complete link method is presented in this paper. One TWINSPAN group with a small number of relevés, set apart on the first level of cluster analysis, was not interpreted (see explanation later in the text).
According to the result of cluster analysis of seven TWINSPAN groups (subclusters), four clusters were obtained, since several authors suggested that there are four groups based on the main ecological and phytogeographical gradient of these forests in the area. Further, these four main clusters were also supported by the spectra of geo-elements and sociological species groups (see below).
Diagnostic species of each of the seven subclusters and four clusters were determined in the JUICE 6.4 program (Tichý 2002) by calculating the fidelity of each species to each cluster and subcluster (Bruel-heide 1995, 2000; Chytrý et al. 2002b; Havlová et al. 2004), using the <p-coefficient as fidelity measure and presented in Table 1. In these calculations, each cluster was compared with the other relevés in the data set, which were taken as a single, undivided group. Each of the seven subclusters and four clusters was virtually adjusted to 1/7 or 1/4 of the size of the entire data set, respectively, while holding the percentage occurrences of a species within and outside a target cluster the same as in the original data set (Tichý & Chytrý 2006). Species diagnostic for the BLRFs of SE Europe (calculated after merging subclusters 2.1,2.2 and 4.1, whose relevés geographically belong to this area) are also indicated in Table 1. For this analysis, fidelity
was calculated based on the merged clusters and after adjusting the number of releves to 1/2 of the size of the entire data set. We also calculated Fisher's exact test and gave zero fidelity value to the species with significance P < 0.001. The range of the 0-coefficient is -1 to 1, but the values were multiplied by 100 in Table 1. The threshold 0-value for the species to be considered as diagnostic was set at 30.0.
For further interpretation of the seven subclusters and four clusters, unweighted average indicator values for releve groups (Pignatti 2005), calculated in the JUICE program, were passively projected onto a Principal Components Analysis biplot (PCA from CANOCO 4.5; ter Braak & Smilauer 2002) to show ecological relationships among these clusters and to explain environmental gradients underlying the main ordination axes. Square-root transformed percentage frequencies were used as the input data.
We also calculated the spectra of geo-elements and sociological species group composition of individual subclusters. Spectra of geo-elements were calculated according to Poldini (1991) and Pignatti (2005). Five sociological species groups - diagnostic species of the class Querco-Fagetea, orders Fagetalia sylvaticae and Quercetaliapubescentis, and alliances Carpinion orientalis and Aremonio-Fagion - were taken from various literature sources (Poldini 1988; Marincek et al. 1993; Mucina et al. 1993; Oberdorfer 1994; Dzwonko & Loster 2000; Marincek & Carni 2000; Blasi et al. 2001, 2004, 2005; Biondi et al. 2002). In general, the categories of geo-elements proposed by Pignatti (2005) were taken into consideration but some adjustments were made, such as Eurymediterranean (incorporating Stenomediterranean), Mediterranean-montane, Eurasian (incorporating montane European, montane central European, montane south European), separately elaborating SE European (incorporating montane SE European) and Pontic, Atlantic (incorporating montane SW-European), Endemic, Cosmopolitan, Palaeotropic and Adventive.
In the calculations, we considered only the species occurring in at least three releves within individual subclusters (Dzwonko et al. 1999). The spectra of geo-elements and sociological species groups of individual subclusters are presented as proportions (percentage) of the entire species composition of individual subclusters. The proportions of geoele-ments in individual subclusters are plotted (similarly as average indicator values) as supplementary data on the PCA ordination diagram of seven subclusters to show phytogeographical relationships. The sociological species group spectra are indicated at the head of the synoptic table to show sociological relationships between subclusters.
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Table 1. Synoptic table of the classification results (Fig. 2). Species values are percentage frequencies. Diagnostic species for the clusters and subclusters (defined as those with 0 > 30.0) are shown, ranked by decreasing value of the 0-coefficient, indicated by asterisks (for subclusters) and shading (for clusters and subclusters). t = tree layer, s = shrub layer, h = herb layer. Some of the species diagnostic for BLFs of southeastern Europe are, at the same time, diagnostic for only one cluster (cluster 2 or 4) and therefore appear twice in the table: they are indicated with r.
Subcluster number No. of releves Cluster number
Sociological type (% of all species in the subcluster)
Carpinion orientalis Aremonio-Fagion Quercetalia pubescentis Fagetalia sylvaticae Querco-Fagetea
1.1	2.1	2.2	3.1	3.2	3.3	4.1
59	141	126	116	49	44	62
1	1 2	1	3			1 4
0.5	1 7.1	4.5 1	0.5	0.6	0.7	0.6
1.6	8.5	9.1	4.8	1.3	2	1 15.6
15.9	13.3	12.9	2.7	5	0.7	0.6
34.9	30.9	32.7	51.3	48.4	52.7	55.7
19.6	17.7	17.8	17.6	12.6	9.6	10.8
Character species of Tilio-Acerion
Fraxinus excelsior	t	51		71	60
Fraxinus excelsior		42		35	52
Ulmus glabra	t	39		48	40
Ulmus glabra		36		37	32
Tilia cordata	t	42		11	44
Tilia cordata		29		4	38 *
Acer platanoides	t	58		40	29
Acer platanoides		49		33	25
Tilia platyphyllos	t	76		55	56
Tilia platyphyllos		39		31	27
Acer pseudoplatanus	t	46		70	67
Acer pseudoplatanus	s	37		38	54
Species diagnostic for one cluster					
Cluster 1					
Poa nemoralis	h	81	*	14	32
Sedum telephium	h	44	*	6	15
Quercus petraea	t	42	*	15	11
Quercus petraea	s	17		5	8
Campanula persicifolia	h	34	*	8	6
Carpinus betulus	t	63	*	40	23
Carpinus betulus	s	39		20	14
Cardaminopsis arenosa	h	29	*	2	6
Galium aparine	h	42	*	24	3
Euonymus verrucosus	s	31	*	11	11
Rosa canina agg.	s	27	*	5	9
Impatiens parvifiora	h			4	2
Chelidonium mqjus	h			15	2
Cluster 2					
Primula vulgaris	h			29	25
Ostrya carpinifolia	t			16	34 *
Ostrya carpinifolia	s			2	5
Hedera helix	s	22		64 *	48
Fraxinus ornus	t	8		27	28
Fraxinus ornus	h	7		17	17
Acer campestre	t	17		51 *	22
Acer campestre	s	29		35	22
Crataegus monogyna	t			1	
Crataegus monogyna	s	8		33	21
Rosa arvensis	s	2		18	22
Melica uniflora	h	12		47 *	18
Cyclamen purpurascens	h	2		10	56 *
Tamus communis	s	2		27	21
Helleborus odorus	h	7		30	16
Festuca heterophylla	h	2		17	20
Daphne laureola	h	3		24	13
Hepatica nobilis	h	25		30	51
Euonymus latifolius	s			13	26
Ligustrum vulgare	h	3		18	12
Melittis melissophyllum	h	5		13	19
Clematis vitalba	s	20		34	32
Cluster 3					
Impatiens noli-tangere	h	20		4	5
Viola bifiora	h				
Polygonatum verticillatum	h				6
Chaerophyllum hirsutum	h	2		2	2
78	67	25	42
64	63	20	26
63	47	25	40
34	45	16	42
5	10	11	2
3	10	2	3
11	2	2	10
16	8	2	8
25	6	14	8
9	6	14	5
89	98	98	95
67	55	59	82
23	47	41	2
5		2	
3			24
4			15
	4		6
7	2		3
3			
8	4	2	
12			
2			3
1			2
25	2	2	10
3			11
3	2		11
3			
6		2	2
4			13
4			3
1			5
1	2		
1			2
10	6	2	6
		2	6
9	10		
55 *	47	36	42
2	16	45 *	
10	37	55	16
10	22	59	13
- Classification and phytogeographical differentiation of broad-leaved forests -
335
Subcluster number No. of releves Cluster number
Sociological type (% of all species in the subcluster)
Carpinion orientalis Aremonio-Fagion Quercetalia pubescentis Fagetalia sylvaticae Querco-Fagetea
1.1
59
2.1 141
2.2 126
3.1 116
3.2 49
3.3 44
4.1
62
0.5 1.6
7.1 8.5
4.5
9.1
0.5 4.8
0.6 1.3
0.7 2
0.6 15.6
15.9	13.3	12.9	2.7	5	0.7	0.6
34.9	30.9	32.7	51.3	48.4	52.7	55.7
19.6	17.7	17.8	17.6	12.6	9.6	10.8
Cluster 4
Stellaria montana	h		4
Symphytum tuberosum	h	5	21
Cardamine bulbifera	h	10	39
Doronicum austriacum	h		
Chrysosplenium alternifolium	h	8	9
Lamium orvala	h		13
Cardamine trifolia	h		1
Adoxa moschatellina	h	7	16
Anemone nemorosa	h	3	21
Cardamine enneaphyllos	h	7	28
Cardamine waldsteinii	h		1
Leucojum Vernum	h		4
Scopolia carniolica	h		1
Dryopteris carthusiana agg.	h	3	2
Oxalis acetosella	h	19	16
Omphalodes verna	h		1
Polystichum braunii	h		1
Anthriscus nitida	h	5	1
Dryopteris affinis	h		1
Gentiana asclepiadea	h		1
Corydalis solida	h	5	13
Lunaria rediviva	h	15	8
Veratrum album	h		
Sambucus nigra	t	2	2
Sambucus nigra	s	24	53
Polystichum x illyricum	h		
Species diagnostic for one subcluster			
Acer obtusatum	t		19
Acer obtusatum	s		1
Lathyrus venetus	h	7	23
Galanthus nivalis	h	7	35
Ruscus hypoglossum	s	2	18
Tilia tomentosa	t	3	18
Tilia tomentosa	s	2	16
Crataegus laevigata	s	5	28
Anemone ranunculoides	h	12	34
Viola alba	h		17
Quercus cerris	t	3	16
Quercus cerris	s	2	4
Ornithogalum umbellatum	h		12
Ranunculus ficaria	h	10	28
Carex digitata	h	12	15
Luzula nivea	h		
Lonicera xylosteum	s	15	10
Asplenium ruta-muraria	h	2	1
Solidago virgaurea	h	3	4
Galium laevigatum	h		
Sorbus aria agg.	t	7	7
Sorbus aria agg.	s	2	3
Anemone trifolia	h		3
Knautia dipsacifolia	h		
Phyteuma spicatum	h		4
Silene vulgaris	h	8	1
Crepis pyrenaica	h		
Adenostyles glabra	h		
Vicia sylvatica	h	2	
Calamagrostis varia	h		1
Bromus ramosus agg.	h	15	10
Valeriana officinalis	h	7	6
Hordelymus europaeus	h	7	8
Cirsium oleraceum	h		4
Pimpinella major	h	3	1
Vicia sepium	h	2	5
Rubus saxatilis	s		
8 22 13
3 2
25
9 13
6 27
2 2
5 33
2
4
6 10
1
5
3 1
29
4
63	*
24	*
72	*
18	*
50	*
17	*
28	
23	*
19	*
5	
16	
6	
1	
19	
18	
6	
1	
3	
4	
3	
1
5
7
24
2
8
9 9
5
7
2 9
53
3 7
6 5
48
3 11
3 39
10
1
2 14
1
8 37
2 10
14 4
5
30 9 9
27 64
5 2 7
25 23
9 2
15
12		16 2	6
31		23	21
2			
49		34 2	2 3
4		9	
6		5	5
45	*	23	
63	*	27	8
31	*	5	
22	*	5	
37	*	18	2
20	*	2	
41	*	16	2
51	*	5	2
41	*	36	
35	*	7	
35	*	23	6
22	*	2	
27	*	7	
20	*	11	
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Subcluster number		1.1	2.1	2.2	3.1	3.2		3.3		4.1		
No. of releves		59	141	126	116	49		44		62		
Cluster number		1    i 1	2	1	3				1	4		1
Sociological type (% of all species												
in the subcluster)												
Carpinion orientalis		0.5	1 7.1	4.5 1	0.5	0.6		0.7		0.6		
Aremonio-Fagion		1.6	8.5	9.1	4.8	1.3		2	1	15.6		1
Quercetalia pubescentis		1 15.9	13.3	12.9	2.7	5		0.7		0.6		
Fagetalia sylvaticae		34.9	30.9	32.7	51.3	48.4		52.7		55.7		1
Querco-Fagetea		19.6	17.7	17.8	17.6	12.6		9.6		10.8		
Carduus defloratus	h			2		16	*	7				
Brachypodium sylvaticum	h	14	31	28	26	65	*	9		24		
Buphthalmum salicifolium	h			2		14	*					
Laserpitium latifolium	h		6	4	1	22	*	2				
Saxifraga rotundifolia	h	2	13	8	13	6		70	*	18		
Polystichum lonchitis	h			1	3	8		41	*	6		
Adenostyles alliariae	h		1		3			36	*	8		
Stellaria nemorum	h	2	1	6	12	12		50	*	19		
Lonicera alpigena	h		1	20	7	16		52	*	8		
Primula elatior	h	2	1	2	10	8		34	*			
Cicerbita alpina	h				2			23	*	5		
Polystichum aculeatum	h	3	21	29	38	24		80	*	40		
Epilobium montanum	h	12	5	17	23	20		59	*	16		
Paris quadrifolia	h		9	35	28	24		75	*	63		
Picea abies	t	5	4	13	16	31		50	*	32		
Picea abies	s	3	3	10	10	31		52	*	34		
Streptopus amplexifolius	h			1	1	6		20	*			
Rubus idaeus	s	20	1	14	14	22		52	*	16		
Rumex alpestris	h		1		1	6		20	*	2		
Lonicera nigra	s			2	1	8		20	*	2		
Salix appendiculata	s			1		2		16	*	2		
Valeriana montana	h					2		14	*			
Crepis paludosa	h		1		2	4		18	*	2		
Cystopteris montana	h							11	*			
Cardamine pentaphyllos	h			22	9	8		41	*	19		
Aconitum variegatum ssp. paniculatum	h			8		18		27	*	6		
Veronica urticifolia	h		1	28	5	33		43	*	5		
Ranunculus platanifolius	h			1		18		20	*	3		
Species diagnostic for more than one subcluster												
Aconitum lycoctonum	h	5	18	10	6	69	*	68	*	10		
Petasites albus	h	2	3	6	16	39		68	*	60		*
Athyrium filix-femina	h	2	9	20	33	31		80	*	90		*
Species diagnostic for broad-leaved ravine		forests of SE Europe										
r Symphytum tuberosum	h	5	21	22	5						66	*
r Lamium orvala	h		13	25	2						56	*
r Cardamine enneaphyllos	h	7	28	27	5	2		9			63	*
r Cyclamen purpurascens	h	2	10	56 *	4						13	
Polystichum setiferum	h	2	28	13	1						23	■
Other species with high frequency												
Fagus sylvatica	t	32	55	45	47	51		27		63		
Fagus sylvatica	s	17	26	37	41	35		14		52		
Lamiastrum galeobdolon	h	47	52	71	68	92		98		89		
Geranium robertianum	h	85	56	61	75	73		73		45		
Dryopteris filix-mas	h	51	38	67	69	53		89		87		
Mercurialis perennis	h	39	52	62	71	80		41		48		
Corylus avellana	s	59	60	84	51	35		16		44		
Urtica dioica	h	63	33	16	61	33		48		71		
Galium odoratum	h	39	44	29	66	61		41		42		
Mycelis muralis	h	51	40	52	31	65		48		27		
Senecio nemorensis	h	29	9	47	44	43		75		53		
Actaea spicata	h	8	21	50	33	41		66		60		
Salvia glutinosa	h	10	26	55	25	59		9		45		
Campanula trachelium	h	37	33	43	16	59		18		8		
Aegopodium podagraria	h	10	38	24	28	51		25		32		
Asarum europaeum	h	25	44	42	30	8		5		29		
Asplenium scolopendrium	h	3	27	39	36	6		27		42		
Polygonatum multiflorum	h	19	48	46	27	4		5		29		
Viola reichenbachiana	h	10	47	33	12	45		11		19		
Pulmonaria officinalis	h	27	26	39	22	6		14		37		
Aruncus dioicus	h	7	9	32	32	18		43		29		
Arum maculatum	h	10	41	14	29	2		2		53		
Fragaria vesca	h	22	20	38	9	37		18		5		
Cystopteris fragilis	h	31	9	17	22	8		41		21		
Asplenium trichomanes	h	24	21	48	21	10		11		13		
Geum urbanum	h	34	34	14	18	18		11		11		
- Classification and phytogeographical differentiation of broad-leaved forests -
337
Results
Clusters and their interpretation
Fig. 2 shows the result of the cluster analysis of the eight TWINSPAN groups of releves (subclusters). The subclusterX comprises 17 releves of xerothermophilous forests, 13 of which are from one author (Hierholzer 1957; Acereto-Coryletum avellanae, Aceri-Fraxinetum) from the northwestern Apennines with a high cover of Corylus avellana. Because of the small size, specific floristic composition and unclear interpretation of this subcluster, we decided to eliminate it from further analyses. Excluding subclusterX, the xerothermophilous forests and mesophilous forests divide at the next first level. At the second level, the two groups also divide geographically into a group of southeastern European and a group of central European forests, resulting in four main clusters. The list of the syntaxa in each subcluster and cluster is given in App. 1.
The classification of broad-leaved ravine forests is presented in a synoptic table (Table 1), in which statistically determined diagnostic species are indicated and ranked by decreasing fidelity.
Cluster 1 corresponds to subcluster 1.1 and is represented mainly by xerothermophilous forests from the middle European and Pannonian sector. The cluster is characterized by a group of thermophilous and nitrophil-ous species with Eurasian distribution, growing at lower altitudes in the Quercus petraea and Carpinus betulus belt. The central European Tilia associations Aceri-Carpinetum, Aceri-Tilietum cordatae, Aceri-Tilietum platyphylli, Mercuriali-Tilietum, Roso pendulinae-Tilietum and Seslerio albicantis-Tilietum cordatae are classified within this cluster.
Cluster 2 is characterized by thermophilous species of southeastern distribution, e.g. Primula vulgaris, Ostrya carpinifolia, Fraxinus ornus, Cyclamen purpurascens, Helleborus odorus and Euonymus latifolius. It comprises of two subclusters. Subcluster 2.1 is represented mainly by xerothermophilous forests from the Apennine, Illyrian and Pannonian sectors, and subcluster 2.2 by xerothermophilous forests from the Eastern Alpine, Illyrian, central Alpine and middle European sectors as it comprises the relict association Asperulo taurinae-Tilietum in the broader sense. Diagnostic species of subcluster 2.1 are thermophilous species with southeastern distribution {Acer obtusatum, Lathyrus venetus, Galanthus nivalis, Tilia tomentosa) andEurymediterranean species (Ruscus hypoglossum, Viola alba, Quercus cerris). Diagnostic species of subcluster 2.2 indicate that the distribution of this subcluster is mainly related with the Alps (e.g. Luzula nivea, Galium laevigatum, Anemone trifolia).
	0.8
	073
	0.7
	0 65
	0.6
	0 55
	0.5
	
■c	
	0 45
1	0.4
	035
<x	
c	0.3
	0 25
	0.2
	0.15
	0.1
	0 05
	0
"3,
0
Fig. 2. Cluster analysis of the eight TWINSPAN groups of releves (subclusters) of BLFs in southeastern and central Europe, resulting in four clusters.
Cluster 3 is characterized by a small group of mesophilous Eurasian species indicating colder and wetter sites. It comprises of three subclusters. Subcluster3. lis represented by mesophilous forests from the middle European, Eastern Alpine, western Alpine and central Alpine sectors; subclusters 3.2 and 3.3 are represented by mesophilous forests from the Eastern, western and central Alpine sectors. Subcluster 3.1 has no diagnostic species, as it represents, ecologically and phytogeographically, the core of central European BLRFs represented by the associations Aceri-Fraxinetum, Arunco-Aceretum, Corydalido-Aceretum and Phylitido-Aceretum. Subcluster 3.2 is characterized by species with Eurasian and Eurosiberian distribution indicating light and warm sites at higher altitudes; its releves mainly belong to the central European associations Sorbo-Aceretum and Asperulo taurinae-Aceretum. Subcluster 3.3 is characterized by a species group indicating cold sites at higher altitudes, represented by the central European association Ulmo-Aceretum.
Cluster 4 corresponds to subcluster 4.1 and is represented almost exclusively by mesophilous forests from the Illyrian sector (Aceri-Fraxinetum illyricum s.lat; Chrysanthemo macrophylli-Aceretum, Dryop-terido affinis-Aceretum, Hacquetio-Fraxinetum, Lamio orvalae-Aceretum,Omphalodo-Aceretum).Thecluster is characterised by the so-called Illyrian species, i.e. relic endemics of mesophilous forest sites with southeastern distribution, including Stellaria montana, Lamium or-vala, Cardamine trifolia, C. enneaphyllos, C. waldsteinii, Scopolia carniolica and Omphalodes verna.
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Phytosociological analyses
According to the phytosociological composition (Table 1), most species in all subclusters are classified as Querco-Fagetea and Fagetalia sylvaticae species, which is in accordance with the classification of these forests into the mentioned higher rank syntaxa. A relatively high proportion of Quercetaliapubescentis species is characteristic of xero-thermophilous BLRFs of clusters 1 and 2. The proportion of Fagetalia sylvaticae is therefore greater in mesophilous BLRFs (clusters 3 and 4). A high proportion of Carpinion orientalis species characterizes both subclusters of cluster 2 (southeastern European Tilia forests). The proportion of Aremonio-Fagion species is considerably higher in cluster 4 (Illyrian Acer forests) than in other clusters of mesophilous forests. Within the xerothermophilous forests the proportion of Aremonio-Fagion species is the highest in both subclusters of cluster 2.
Indicator values and geo-elements
PCAs are presented of the seven subclusters with mean Pignatti indicator values (Fig. 3) and spectrum of geo-elements (Fig. 4) plotted as supplementary variables on the ordination diagram. Eigenvalues of the first two axes are 0.411 and 0.213. Xerothermophilous and mesophilous BLRFs are separated along the axis 1, while along axis 2 these forests are separated according to phytogeography.
Temperature and moisture (Fig. 3) show a high correlation with PCA axis 1, light and nutrient show moderately high correlation with axis 2. Tilia forests occur on warmer and drier sites, Acer forests on colder and wetter sites. Forests of SEEurope (Apennine-Balkan province) occur on nutrient richer soil and more shaded sites than C-European BLRFs. Nutrient richer soils are probably associated with thriving in the region of a warmer macroclimate. Together with the relatively wet sites, this suggests that decomposition of organic matter proceeds faster. Acer and Tilia forests, and forests generally, are better preserved in the Illyrian sector and less changed due to economic influences, and perhaps more shade.
The proportions of Pontic, Eurymediterranean and Eu-rosiberian species are highly correlated with axis 1 and the proportions of SE-European and Mediterranean-montane species are correlated with Axis 2 (Fig. 4). This means that Pontic and Eurymediterranean species are characteristic of subclusters of Tilia forests on the left side of the diagram, Eurosiberian species of subclusters of Acer forests on the right side, SE-European and Mediterranean-montane species of Apennine-Balkan subclusters in the upper part, and Eurasian species of C-European subclusters in the lower part of the diagram.
			O 4.1 ^Nutrients / Moisture
Temperature	.-'2.1 \ \ °	\	/ 3 V-
		^P? 1 \ / /	\             3.3 /
	O 1.1	Light	\ 3.2 /' \ o /
-1.5			1.0
Fig. 3. Principal components analysis (PCA) of subclusters and Pignatti indicator values plotted as supplementary data on the diagram. The subclusters are numbered as in Table 1. Clusters are framed. Only indicator values with the highest correlations with the first two PCA axes are shown. The highest correlations with the first axis have the indicator values for temperature (-0.988) and moisture (0.824), with the second axis the values for light (-0.889) and nutrients (0.789).
Discussion
Many authors have found that broad-leaved ravine forest communities in SE Europe differ from C-European communities. However, there are different views on the synsystematic classification of BLRFs:
1. Classification within the alliance of Fagus forests Aremonio-Fagion (Horvat 1938,1962; Glavac 1958; Accetto 1991; Vukelic &Raus 1998);
2. Classification within the separate suballiance Polys-ticho setiferi-Acerenion, still within Fagus forests of the alliance Aremonio-Fagion (Kosir 1972; Marincek 1990; Borhidi & Kevey 1996; Dakskobler 1999; Kosir & Marincek 1999; Kosir 2002);
3. Classification into the broadly conceived central European alliance Tilio-Acerion (Zupancic & Zagar 1999; Taffetani 2000; Biondi et al. 2002; Catorci et al. 2003; Angiolini et al. 2005);
4. Classification into an independent SE- European alliance Fraxino-Acerion Fukarek 1969 (Fukarek 1969; Jovanovic et al. 1986; Stefanovic 1986; Saric 1997; Kojic et al. 1998; Kosir 2005 a, b) or heterogeneous alliance Euonymo latifolii-Fagion (Ubaldi 2003), vicariating with Tilio-Acerion in the Apennines.
The idea of classifying the SE-European BLRFs into the alliance Aremonio-Fagion was proposed after a study of Acer dominated forests occurring intrazonally within an area of Illyrian Fagus forests (Horvat 1938;Marincek
- Classification and phytogeographical differentiation of broad-leaved forests -
339
SE-European Mediterranean-montane
O 4.1
Eury mediterranean
Eurasian
-1.0 1.0
Fig. 4. PCA of subclusters and proportion of geo-elements in subclusters plotted as supplementary data on the diagram. The subclusters are numbered as in Table 1. Clusters are framed. Only geo-elements with the highest correlations with the first two PCA axes are shown. The highest correlations with the first axis have the values for Eurymediterranean (-0.897), Pontic (-0.903) and Eurosiberian (0.928) geoelement, with the second axis the values for SEEuropean (0.853), Mediterranean-montane (0.804) and Eurasian (-0.719) geoelement.
1990). However, not all the BLRFs are spatially related to Fagus communities. In fact, Tilia dominated forests are more often linked to the zonal forests belonging to the alliances Erythronio-Carpinion or Carpinion orientalis. This suggests that classifying the southeastern European BLRFs into the alliance Aremonio-Fagion is not the most suitable solution.
Classification into an independent alliance, as proposed by Fukarek (1969), is not a suitable solution as we could not confirm it by the numerical analyses. Numerical analyses (Fig. 2) show that xerothermophilous or mesophilous BLRFs of different phytogeographic regions are more similar than xerothermophilous and mesophilous BLRFs of a single region. Despite this, there are a few species linking together mesophilous and xerothermophilous BLRFs of southeastern Europe (e.g. Polystichum setiferum, see Table 1).
Cluster analysis has suggested that BLRFs in southeast Europe differ from central European ones and that they form separate groups (Fig. 2). However, the central European alliance Tilio-Acerion is characterized by the dominance of some tree species such as Acer platanoides, A. pseudoplatanus, Fraxinus excelsior, Tilia cordata, T. platyphyllos, Ulmus glabra, which dominate ravine forests not only in central Europe but also in a large part of SE Europe. Therefore, we propose the assignment of BLRFs
of southeastern Europe into the alliance Tilio-Acerion, and a further subdivision into two new suballiances of southeastern European distribution, one of which includes xerothermophilous and the other mesophilous BLRFs. These suballiances, named Lamio orvalae-Acerenion and Ostryo-Tilienion (see description below), are parallel to the already established central European suballiances Lunario-Acerenion and Tilienion (Table 1).
There are more plant species in BLRFs of SE Europe than in those of C Europe due to the general richness of woody species of SE Europe due to the effect of glacial refugia and the possible relic character of some stands (Trepp 1947). SE-European suballiances are, therefore, defined by groups of geographical differential species which are also ecologically coherent. We think that the classification based on ecological, as well as geographical, differential groups is more generally applicable than the classification which considers only ecological differences. After all, the differences in geographical position that result in different macroclimatic conditions are always reflected together with ecological ones.
Syntaxonomical classification
The suggested syntaxonomical classification of broad-leaved ravine forests in central and southeastern Europe is as follows:
Class: Querco-Fagetea Br.-Bl. et Vlieger in Vlieger 1937
Ordo: Fagetalia sylvaticae Pawlowski et al. 1928
Alliance: Tilio-Acerion Klika 1955
Suballiance: Tilienion platyphylli (Moor 1975) Th.
Miiller 1992 (cluster 1 in Table 1)
Suballiance: Lunario-Acerenion pseudoplatani (Moor
1973) Th. Miiller 1992; (cluster 3 in Table 1)
Suballiance: Lamio orvalae-Acerenion pseudoplatani
suball. nova hoc loco (cluster 4 in Table 1)
Suballiance: Ostryo carpinifoliae-Tilienion platyphylli
suball. nova hoc loco (cluster 2 in Table 1)
The nomenclatural type (holotypus) of the Lamio orvalae-Acerenion pseudoplatani is the association Omphalodo vernae-Aceretum R Kosir et Marincek 1999 holotypus hoc loco. Illyrian mesophilous BLRFs are classified into this suballiance.
The holotypus of the Ostryo carpinifoliae-Tilienion platyphylli is the association Saxifrago petraeae-Tili-etum platyphylli Dakskobler 1999 holotypus hoc loco. Xerothermophilous BLRFs with the distribution centre in SE Europe (Apennine-Balkan province) are classified into this suballiance.
Mesophilous forests of the suballiance Lamio orvalae-Acerenion are geographically bound to the Illyrian subprovince (Fig. 1), as they are defined with meso-
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philous Illyrian species which are gradually disappearing towards the South and the east of the region as a result of the unfavourable climate (higher temperatures, unfavourable precipitation regime). Stands of Acer forests appear intrazonally within the range of zonal Illyrian beech forest communities of alliance Aremonio-Fagion.
The suballiance Ostryo-Tilienion includes xerofher-mophilous BLRFs from a great part of the Apennine-Balkan province (Fig. 1), especially from the regions with sub-mediterranean climate. They are defined by a diagnostic species combination which is partly shared with the alliance Carpinion orientalis. Only few associations of the Ostryo-Tilienion can be found also outside the limits of the Apennine-Balkan province as scattered islands of thermophilous relic BLRFs (e.g. association Asperulo taurinae-Tilietum in the Central Alpine and Eastern Alpine sectors and more sporadically in the Middle European sector, or associations Scutellaria altissimae-Aceretum and Tilio tomentosae-Fraxinetum orni at the edge of the Pannonian sector).
In the southern Apennines and Balkans, the presence and cover of tree species characteristic of the Tilio-Acerion drastically decrease, while other tree species such as Acer obtusatum, A. lobelii, Aesculus hippocastanum, Alnus cordata, Tilia platyphyllos ssp. pseudorubra and T. to-mentosa become dominant (Bonin 1978; Bergmeier 1990; Mazzoleni & Ricciardi 1995; Brullo et al. 2001; Corbetta etal. 2004; Amanatidou2005). The syntaxonomic affinity of these tree species is not yet clear. As the alliance Tilio-Acerion is defined by dominance of Acer platanoides, A. pseudoplatanus, Fraxinus excelsior, Tilia cordata, T. platyphyllos and Ulmus glabra, we support the opinion that communities from the extreme south of Europe should be classified into a separate alliance vicariating with Tilio-Acerion (Brullo et al. 2001) or a separate suballiance within the alliance Carpinion orientalis (Blasi et al. 2006).
Vegetation in SE Europe (the Apennine-Balkan province) is chorologically uniform. This is because the refugia where the Apennine-Balkan vegetation, as well as European Tilia species (Tilia cordata, T. platyphyllos, T. rubra, T. tomentosa), survived the Quaternary glaciation were in the Apennine and the Balkan peninsula and on the southern and southeastern edges of the Alps (Lang 1994, 2003; Sercelj 1996; Magri 1998).
The distribution centre of mesophilous BLRFs is in the Central European and Alpine provinces. We can find most southern mesophilous stands only in the Illyrian sector of the Apennine-Balkan province. In contrast the distribution centre of xerothermophilous BLRFs is in southeastern Europe, namely in the Apennine-Balkan province.
Conclusions
It was found that broad-leaved ravine forests in the Apennine-Balkan province differ from the Central European BLRFs and can be differentiated as two suballiances within the alliance Tilio-Acerion, characterized geographically, ecologically and chorologically, sharing the same history. There are still some open questions about classification of BLRFs from the southern Apennine and Balkan peninsula that need further investigations. In the future, classification of BLRFs in southeastern Europe, especially within suballiance Ostryo-Tilienion, should be done also at the level of associations. This article establishes the similarity between the vegetation on both sides of the Adriatic Sea and thus confirms certain contemporary phytogeographical findings.
Acknowledgements. For discussion, comments and literature we thank Lojze Marinček (SI), Joso Vukelič (HR), Senka Barudanovič (BA), Igor Dakskobler (SI), IstvanDancza (HU), Josip Franjič (HR), Paul Heiselmayer (AT), Walter Keller (CH), BalázsKevey (HU),UvioPoldini (IT),Andreas Stampfli (CH), Zeljko Škvore (HR), Fabio Taffetani (IT), Jean-Paul Theurillat (CH) and for language revision Mr. Alan McConnell-Duff. We would also like to thank Milan Chytrý, Wolfgang Willner and Radim Hédl for their suggestions and comments on previous versions of the manuscript.
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Received 17 April 2007; Accepted 26 July 2007; Co-ordinating Editor: M. Chytrý.
For App. 1, see JVS/AVS Electronic Archives;
www.opuluspress.se/
I
App. 1. Sources of data used for the analysis. Subcluster codes refer to Fig. 2 and Table 1. Subcluster 1.1
Jovanovic (1985), Tilio-Fraxinetum excelsioris, (relevé 3), Schwarz (1991), Aceri-Tilietum cordatae, Table 5 (relevés 2, 7, 8), Schwarz (1991), Aceri-Tilietumplatyphylli, Table 5 (relevés 11,12, 14, 17,18), Reitter-Hebenstreit (1984), Aceri-Tilietumplaty-phylli, Table 5 (relevé 61), Isépy (1968), MercuriaJi-Tilietum, Table 2 (relevé 7), Horánszky (1964), Mercuriali-Tilietum, Table 5 (relevés 3,4), Fekete (1963), Phyllitidi-Aceretum, Table 1 (relevé 5), Fekete & Járai-Komlódi (1962), Mercuriali-Tilietum, Table (relevés 5, 18, 23, 27), Kovács (1968), Mercuriali-Tilietum, Table 1 (relevés 1, 3, 6), Jakucs (1967), Phyllitidi-Aceretum, Table 3 (relevés 19,21, 24, 29), Csiky et al. (2001), Rosopendulinae-Tilietum, Table 4 (relevés 4-11), Knapp (1944), Acereto-Fraxinetum timokense viburnetosum (one relevé), Willner (1996), Aceri Carpinetum, Table 1 (relevés G106, G313, L511, JELI), Moravcová-Husová (1964), Fago-Aceretum lunarietosum (Lunario-Aceretum), Table 1 (relevé 18), Husová et al. (2003), Aceri-Carpinetum, Table 6.6.1. (relevés 8, 9, 14, 16), Husová et al. (2003), Lunario-Aceretum, Table 6.6.3 (relevé 7), Husová et al. (2003), Arunco-Aceretum, Table 6.6.5 (relevé 10), Husová et al. (2003), Seslerio albicantis-Tilietum cordatae, Table 6.6.8 (relevé 2), Chytrý & Sádlo (1997), Seslerio albicantis-Tilietum cordatae, Table 1 (relevés 9, 12, 14, 26), Husová (1982), Aceri-Carpinetum, Table 2 (relevés 22, 26, 32, 38), Pekanovié (1991), Tilio-Fraxinetum excelsioris, Table 17 (relevés 1, 2, 7), Pekanovié (1991), Aceri-Fraxinetum excelsioris, Table 16 (relevés 1, 3).
Subcluster 2.1
Zupančič & Zagar (1999), Arunco-Aceretum, Table (relevés 3, 4), Blečié & Lakušič (1967), Aceri-Fraxinetum montenegrium Table 5 (relevé 1), Jovanovic (1985), Tilio-Fraxinetum excelsioris, (relevé 2), Jovanovic (1985), Aceri-Fraxinetum, (two relevés), Lasen & Urbinati (1995), Hacquetio-Fraxinetum, Table 1, group H (relevés 30,38), Busnardo & Lasen (1994), Aceri-Fraxinetum et Aceri-Tilietum, Table (relevé 1), Košir & Surina (2005), Paeonio officinalis-Tilietum platyphylli, Table 1 (relevés 3, 4, 5, 6, 10, 13, 14, 15, 17, 18), Košir (2002), Hacquetio-Fraxinetum, Table 1 (relevé 1), Sturm (1978), Aceri-Fraxinetum corydaletosum cavae, Table 6 (relevés 96-100), Rauscher (1990), Aceri-Fraxinetum stachyetosum sylvaticae, Table 9 (relevé 10), Tezner (1958), Gipfeleschenwald, Table p. 65 (relevés G88, G6, G9), Taffetani (2000), Ornithogalo sphaerocarpi-Aceretum pseudoplatani, Table 24 (relevés 3-5), Antonietti (1968), Cirsio erisithali-Ulmetum, Table 1 (relevé 38), Biondi et al. (2002), Fraxino excelsioris-Aceretum obtusati, Table 5 (relevés 1-3, 5-6, 8, 10-12), Biondi et al. (2002), Aceretum obtusati-pseudoplatani, Table 6 (relevés 1,2), Kevey (1985), Scutellario altissimae-Aceretum, Table 1 (relevés 1,4), Isépy (1968), Phyllitidi-Aceretum transdanubicum, Table 1 (relevés 1,4, 13), Kevey (1997), Scutellario altissimae-Aceretum, Table 1 (relevés 13, 19, 21), Košir (2004), Hacquetio-Fraxinetum, Table 6 (relevé 4), Kevey & Borhidi (1998), Tilio tomentosae-Fraxinetum orni, Table 3 (relevés 2,4, 5, 8, 13, 14,16-19), Kevey (1993), Scutellario altissimae-Aceretum, Table 1 (relevés 2, 10, 20, 21, 26), Willner (1996), Aceri-Carpinetum, Table 1 (relevés G123, G101, E4, G302, G521), Kiridus (1987), unknown ass. subass. Acer pseudoplatanus var. Allium ursinum, Table (relevés 216,224), Moravcová-Husová (1964), Fago-Aceretum typicum, Table 1 (relevé 16), Clot (1989), Corydalido-Aceretum, Table 4 (relevés 116, 120), Keller (1974), Asperulo taurinae-Tilietum, Table 1 (relevé 22), Borhidi & Kevey (1996), Polysticho setiferi-Aceretum, (one relevé), Hettwer (1999), Fraxino-Aceretum typicum, Table 2 (relevé 2), Hettwer (1999), Fraxino-Aceretum lunarietosum, Table 2 (relevé 16), Bergmeier (1990), Rusco hypoglossi-Aesculetum hippocastani, Table 17 (relevé 4), Raus (1980), Aesculus hippocastanum-Tiliaplatyphyllos community, Table 10 (two relevés), Košir & Škvore (unpubl.), Aceri-Fraxinetum s.l. (three relevés), Pirone et al. (2005), Aceretum obtusati-pseudoplatani, Table 2 (relevés 1-8), Angiolini et al. (2005), Ornithogalo sphaerocarpi-Aceretum pseudoplatani, Table 1 (relevés 6, 10, 23, 44, 48), Angiolini et al. (2005), Glechomo hirsutae-Aceretum obtusati, Table 1 (relevé 49), Angiolini et al. (2005), Acer campestre-Tiliaplatyphyllos-Corylus avellana community, Table 1 (relevés 54-56,61,64),Ubaldi (1988), Agropyro-Coryletum, Table 5 (relevé 1), Maiorca et al. (2006), Acerpseudoplatanus-Ostrya carpinifolia community, Table 2 (relevé 1), Pirone (1998), Mesopbilous mixed forest, Table 4 (one relevé), Pedrotti (1987), Aceri-Ulmetum montanae (one relevé), Brullo et al. (2001), Corylo-Aceretum neapolitani, Table 22 (relevé 2), Rizovski & Džekov (1990), Aceri-Fraxinetum excelsioris, Table 10 (one relevé), Lakušič & Redžič (1989), Aceri-Fraxinetum montenegrium, Table 6 (relevé 1), Biondi et al. (2004), Aceretum obtusati-pseudoplatani, Table 7 (relevé 1), Catorci et al. (2003), Aceretum obtusati-pseudoplatani, Table 2 (relevé 31), Jankovič & Mišič (1960), Fagetum montanum serbicum tilietosum, Table 1 (relevés 3, 12), Pekanovié (1991), Tilio-Fraxinetum excelsioris, Table 17 (relevés 3-5), Pekanovié (1991), Aceri-Fraxinetum excelsioris, Table 16 (relevés 5, 7, 11, 12), Dakskobler (2006), Veronico sublobatae-Fraxinetum, Table 2 (relevés 1-7), Dakskobler (2006), Saxifrago petraeae-Tilietum, Table 3 (relevé 4), Dakskobler (2006), Veratro nigri-Fraxinetum, Table 3 (relevés 5, 6), Medarevié (2005), Aceri-Fraxinetum, Table 5.17 (relevés 31/03, 20/03, 57/02), Schlüter (1967), Aceri-Ulmo-Fraxinetum, Table 2 (relevés 13-17).
Subcluster 2.2
Košir (1954), Ostrýo-Tilietum,Table (relevés 1,3,7,13,16,18), Dakskobler (1999), Saxifragopetraeae-Tilietumplatyphylli,Table 4 (relevés 1, 4, 8, 10, 12), Accetto (1991), Corydalido ochroleucae-Aceretum, Table 1 (relevés 1, 6, 10, 22, 25), Glavač (1958), Tilio-Taxetum, Table (relevés 1, 2, 5, 7, 12, 13, 15), Blečié & Lakušič (1967), Aceri-Fraxinetum montenegrium, Table 5 (relevés 3, 4), Lasen & Urbinati (1995), Aceri-Fraxinetum with Ostrya, Table 1, group C+D (relevés 3, 2, 82, 83, 62, 34, 17, 29, 16, 60), Lasen & Urbinati (1995), Hacquetio-Fraxinetum, Table 1, group H (relevé 48), Busnardo & Lasen (1994), Aceri-Fraxinetum et Aceri-Tilietum, Table (relevés 6, 5, 18, 15, 14, 9, 21, 20, 12), Košir (2004), Corydalido ochroleucae-Aceretum, Table 11 (relevés 1, 2, 5, 6), Marinček (1990), Hacquetio-Fraxinetum, Table (relevé 16), Poldini & Nardini (1993), Hacquetio-Fraxinetum,
App. 1. Internet supplement to: Kosir, P.; Carni, A. & Di Pietro, R. 2008. »fyopuLUSPRESS
Classification and phytogeographical differentiation of noble hardwood forests in southeastern Europe. ' Journal of Vegetation Science 19: 331-342; doi: 10.3170/2008-8-18372
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Table 1 (relevés 2, 3), Dakskobler (1999), Hacquetio-Fraxinetum, Table 2 (relevé 14), Fischer (2000), Aceri-Tilietum, Table 1 (relevé 8), Peter (1991), Asperulo taurinae-Tilietum, Table 10 (relevés 4, 45), Pfeifer (1992), Asperulo taurinae-Tilietum, Table 1 (relevés 11, 18), Hotter (1999), Arunco-Tilietum cordatae, Table (relevé 4), Strobl (1989), Aceri-Fraxinetum forma Vincetoxicum hirundinaria, Table 2, group g (relevés 68, 285), Schwarz (1991), Aceri-Tilietum cordatae, Table 5 (relevés 3-6, 10), Rauscher (1990), Aceri-Fraxinetum typicum, Table 9 (relevé 27), Reitter-Hebenstreit (1984), Aceri-Tilietum platyphylli, Table 5 (relevés 78,72, 53,76), Antonietti (1968), Cirsio erisithali-Ulmetum, Table 1 (relevés 23,30,33,34,44, 62, 68), Csiky et al. (2001), Roso pendulinae-Tilietum, Table 4 (relevés 2, 3), Dakskobler (unpubl.), Saxifrago petraeae-Tilietum platyphylli, (four relevés), Clot
(1989) , Phyllitido-Aceretum, Table 2 (relevé 47), Keller (1974), Asperulo taurinae-Tilietum, Table 1 (relevés 1,3,4, 8, 14, 17, 20), Moor (1975), Phyllitido-Aceretum, Table (relevés 5, 12, 14), Poldini (1982), Carpino-Fraxinetum cerastietosum sylvaticae, Table 9 (relevés 24,26), Stampfli (1986), Tiliaforest, Table 3 (relevés 9,21,24,37,42), Trepp (1947), Asperulo taurinae-Tilietum, Table 1 (relevés 3, 14, 24, 32, 37, 63), Chytrý & Sádlo (1997), Seslerio albicantis-Tilietum cordatae, Table 1 (relevés 1, 6, 7, 19, 29), Angiolini et al. (2005), Ornithogalo sphaerocarpi-Aceretumpseudoplatani, Table 1 (relevé 39), Pedrotti & Gafta (1999), Salvio glutinosae-Fraxinetum excelsioris, Table 4 (relevés 2,3, 5-9,11, 12), Pedroti & Gafta (1999), Calamintho grandiflorae-Aceretum pseudoplatani, Table 5 (relevés 1, 3, 5).
Subcluster 3.1
Accetto (1991), Corydalido ochroleucae-Aceretum,Table 1 (relevé 13),Fukarek& Stefanovic (1958), Aceri-Fraxinetum illyricum, Table 1 (relevés 1,2), Blečié & Lakušič (1967), Aceri-Fraxinetum montenegrium, Table 5 (relevés 2,5), Poldini & Nardini (1993), Hacquetio-Fraxinetum, Table 1 (relevé 4), Fischer (1998b), Arunco-Aceretum, Table 1 (relevé 8), Fischer (1998b), Phyllitido-Aceretum, Table 2 (relevé 4), Fischer (1997a), Fraxino-Aceretum, Table 12 (relevés 2, 6), Fischer (1997a), Phyllitido-Aceretum, Table 14 (relevé 1), Fischer (1997a), Arunco-Aceretum, Table 13 (relevé 2), Fischer (1998a), Arunco-Aceretum, Table 2 (relevés 11), Fischer (1998a), Phyllitido-Aceretum, Table 3 (relevé 1), Fischer (1997b), Arunco-Aceretum, Table 1 (relevés 3, 20), Fischer (2000), Phyllitido-Aceretum, Table 1 (relevé 3), Fischer (1998c), Aceri-Fraxinetum corydaletosum cavae, Table 1 (relevés 1-3), Peter (1991), Phyllitido-Aceretum, Table 9 (relevé 96), Pfeifer (1992), Aceri-Fraxinetum, Table 2 (relevés 8, 9, 14, 16, 17, 21), Sturm (1978), Arunco-Aceretum, Table 6 (relevés 101, 105), Strobl (1989), Aceri-Fraxinetum, forma Allium ursinum, Table 2, group a (relevé 96), Strobl (1989), Aceri-Fraxinetum, forma Anthriscus nitida, Table 2, group f (relevés 209,300,304), Rauscher
(1990) , Arunco-Aceretum, Table 12 (relevé 1), Rauscher (1990), Phyllitido-Aceretum, Table 13 (relevé 2), Brennsteiner (1984), Aceri-Fraxinetum, Table (relevé 8), Weinmeister (1983), Acereto-Ulmetum, Table 3 (relevé 123), Horánszky (1964), Phyllitidi-Aceretum, Table 4 (relevé 1), Jakucs (1967), Phyllitidi-Aceretum subcarpaticum, Table 3 (relevés 1, 6, 11, 16-18, 20, 22, 23, 25, 28), Willner (1996), Corydalido-Aceretum, Table 1 (relevé G400), Pfadenhauer (1969), Ulmo-Aceretum, Table 22 (relevés 334, 404), Büker (1942), Aceri-Fraxinetum typicum, Table 26 (relevé 4), Hartmann & Jahn (1967), Aceri-Fraxinetum typicum, Table 6 (relevés 11,13), Clot (1989), Phyllitido-Aceretum, Table 2 (relevés 57,60), Table 3 (relevé 86), Clot (1989), Corydalido-Aceretum, Table 4 (relevés 111, 113-115, 117,118, 123), Clot (1989), Arunco-Aceretum, Table 5 (relevé 147), Husová et al. (2003), Lunario-Aceretum lunarietosum, Table 6.6.3 (relevé 4), Husová et al. (2003), Arunco-Aceretum abietetosum, Table 6.6.5 (relevés 3, 5, 8, 9, 11, 12), Keller (1974), Asperulo taurinae-Tilietum, Table 1 (relevé 2), Keller (1974), Phyllitido-Aceretum, Table 2 (relevé 6), Moor (1975), Phyllitido-Aceretum, Table (relevés 6, 7, 8, 19, 27, 40, 46, 47, 58, 63), Moor (1952), Phyllitido-Aceretum, Table 1 (relevés 8, 14), Moor (1952), Arunco-Aceretum, Table 4 (relevés 1,3), Stampfli (1986), Tilia forest, Table 3 (relevés 28,30,31,40, 44), Hettwer (1999), Fraxino-Aceretum typicum, Table 2 (relevés 1,3-7, 9, 10), Hettwer (1999), Fraxino-Aceretum lunarietosum, Table 2 (relevés 22, 24), Hettwer (1999), Fraxino-Aceretum phyllitidetosum, Table 2 (relevés 37, 41, 60), Pfadenhauer (1969), Arunco-Aceretum, Table 18 (relevé 171), Pfadenhauer (1969), Phyllitido-Aceretum, Table 20 (relevé 359), Pfadenhauer (1969), Phyllitido-Aceretum,Table 21 (relevés 407,723), Košir& Škvore (unpubl.), Aceri-Fraxinetum s. lat., (onerelevé), Angiolini et al. (2005), Ornithogalo sphaerocarpi-Aceretum pseudoplatani, Table 1 (relevé 33), Pedroti & Gafta (1999), Calamintho grandiflorae-Aceretum pseudoplatani, Table 5 (relevés 2, 4).
Subcluster 3.2
Pfeifer (1992), Ulmo-Aceretum, Table 5 (relevés 1,3,5, 6), Pfeifer (1992), Aceri-Fraxinetum, Table 2 (relevés 2,5,7, 20), Pfeifer (1992), Phyllitido-Aceretum, Table 6 (relevés 1-5), Pfeifer (1992), Asperulo taurinae-Aceretum, Table 3 (relevés 1, 2,7, 9, 10, 12, 14-17), Hotter (1999), Arunco-Tilietum cordatae, Table (relevés 1-3,6), Strobl (1989), Aceri-Fraxinetum forma typica, Table 2, group c (relevé 22), Strobl (1989), Aceri-Fraxinetum forma Aruncus dioicus, Table 2, group d (relevé 9), Strobl (1989), Aceri-Fraxinetum forma Vincetoxicum hirundinaria, Table 2, group g (relevés 48, 291, 228, 215, 216,290), Feldner (1978), Aceri-Fraxinetum, Table 3 (relevé 533), Clot (1989), Corydalido-Aceretum, Table 4 (relevé 122), Clot (1989), Arunco-Aceretum, Table 5 (relevés 152,155), Clot (1989), Sorbo-Aceretum, Table 9 (relevés 270, 273, 277, 282, 287, 290, 297, 301), Moor (1952), Sorbo-Aceretum, Table 2 (relevé 8), Moor (1952), Arunco-Aceretum, Table 4 (relevé 4), Pedrotti & Gafta (1999), Salvio glutinosae-Fraxinetum excelsioris, Table 4 (relevé 4).
Subcluster 3.3
Hotter (1999), Arunco-Tilietum cordatae, Table (relevés 8-12), Strobl (1989), Aceri-Fraxinetum forma Phyllitis scolopendrium, Table 2, group e (relevé 239), Strobl (1989), Aceri-Fraxinetum forma Vincetoxicum hirundinaria, Table 2, group g (relevé 28), Feldner (1978), Phyllitido-Aceretum, Table 3 (relevé 126), Feldner (1978), Adenostylo alliariae-Aceretum, Table 3 (relevés 128,
App. 1. Internet supplement to: Kosir, P.; Carni, A. & Di Pietro, R. 2008. »(«Jopuluspress
Classification and phytogeographical differentiation of noble hardwood forests in southeastern Europe. Journal of Vegetation Science 19: 331-342; doi: 10.3170/2008-8-18372
III
277, 257, 250), Moor (1975), Phyllitido-Aceretum lonchiditetosum, Table (relevés 72-75, 77-80), Pfadenhauer (1969), Ulmo-Aceretum, Table 22 (relevé 753), Clot (1989), Phyllitido-Aceretum, Table 3 (relevé 90), Clot (1989), Arunco-Aceretum, Table 5 (relevés 136, 139, 140), Clot (1989), Ulmo-Aceretum, Table 6 (relevés 175, 187, 191), Table 7 (relevés 202, 205, 214, 217), Table 8 (relevés 247, 258, 269), Clot (1989), Sorbo-Aceretum, Table 9 (relevé 282), Moor (1952), Sorbo-Aceretum, Table 2 (relevé 3), Moor (1952), Arunco-Aceretum, Table 4 (relevé 8), Pfadenhauer (1969), Arunco-Aceretum, Table 18 (relevés 402, 689), Pedrotti & Gafta (1999), Calamintho grandiflorae-Aceretumpseudoplatani, Table 5 (relevés 7-10).
Subcluster 4.1
Zupančič (1997), Corydalido cavae-Aceretum, Table (relevé 1), Zupančič & Zagar (1999), Arunco-Aceretum, Table (relevés 5,7,
9) , Košir (2005a), Lamio orvalae-Aceretum, Table 6 (relevés 1,2,8,9,11,12,19,21,22,27), Košir (2005b), Omphalodo-Aceretum, Table 7 (relevés 1, 2,3, 10, 12, 14, 16, 18, 19, 21), Košir (2005b) Chrysanthemo macrophylli-Aceretum, Table 8 (relevés 6, 11, 15, 18), Table 9 (relevés 3, 9, 20), Table 10 (relevés 2,4, 5), Regula-Bevilacqua (1978), Aceri-Fraxinetum illyricum, Table 39 (relevé 1), Lasen & Urbinati (1995), Hacquetio-Fraxinetum, Table 1, group H (relevés 44,68), Marinček (1995), Hacquetio-Fraxinetum, Table 4 (relevé 14), Marinček (1990), Hacquetio-Fraxinetum,Table (relevé 12), Košir (2002), Hacquetio-Fraxinetum,Table 1 (relevé
10) , Sturm (1978), Arunco-Aceretum,Table 6 (relevés 103,104,106), Franz (1991), Fraxino-Aceretumpseudoplatani lamietosum orvalae, Table 1 (relevé 8), Horvat (1938), Aceri-Fraxinetum croaticum (one relevé), Košir (2005c), Dryopterido affinis-Aceretum, Table 5 (relevés 2, 5, 6, 13, 19, 23, 30, 31, 33, 36), Košir (2004) Hacquetio-Fraxinetum, Table 6 (relevé 16), Košir (unpubl.), Hacquetio-Fraxinetum (two relevés), Košir & Škvore (unpubl.), Aceri-Fraxinetum s.l. (four relevés).
Subcluster X
Hierholzer (1957), Aceri-Coryletum avellanae, Table 6 (relevés 1, 2,4, 5, 6,8, 10, 12), Table 7 (relevés 7, 10), Hierholzer (1957), Aceri-Fraxinetum, Table 8 (relevés 5,6,8), Keller (1974), Asperulo taurinae-Tilietum, Table 1 (relevé 11), Antonietti (1968), Cirsio erisithali-Ulmetum, Table 1 (relevés 47, 48), Biondi et al. (2002), Fraxino excelsioris-Aceretum obtusati, Table 5 (relevé 7).
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