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New and validated high-rank syntaxa from Europe.

INTRODUCCION

In the course of compilation of a standard checklist for the high-rank syntaxa (alliances, orders, classes) of Europe (Mucina & al., in preparation), it turned out that for several syntaxa no valid or legitimate names according to the International Code of Phytosociological Nomenclature (further ICPN; Weber & al., 2000) were available. Some of these invalid names even are in general usage, probably because authors are not aware of the invalidity. This brief nomenclatural contribution attempts to rectify the situation by validating such names. Further, two new alliances resulting from the subdivision of existing alliances as well as a new association serving as type of one of these new alliances are published. Finally, the illegitimate name of one alliance is replaced by a nomen novum.

Altogether, seven new syntaxa and one new name are presented. They belong to the classes Cisto-Micromerietea julianae, Festuco-Brometea, Quercetea ilicis, Stellarietea mediae, and Trifolio-Geranietea sanguinei. Most of them occur on the Balkan Peninsula, but also one syntaxon of subatlantic-submediterranean Europe and one of northwestern Europe are included.

In the following, the syntaxa are treated in alphabetical order. The structure of the entries mostly follows the proposal of DENGLER & al. (2003). The presentations of the syntaxa are the responsibility of the authors named in the respective subheadings. When quoting them, cite "<author(s)> in Mucina & al. 2009".

VALIDATIONS AND DESCRIPTIONS OF SYNTAXA

Alkanno baeoticae-Pinetum halepensis Mucina & Dimopoulos ass. nova hoc loco

Holotypus: releve LM 6702; sampled by Ladislav Mucina; 21-06-1999; sampling scale: Barkman & al. (1964); Greece, island of Euboea (Evvia), Kroupi (Dim. Kireos), S of village; 38 42' 53.5" N; 23 25' 56.71" E; altitude: 340 m a.s.l.; aspect 13; slope: 10; area: 15 m x15 m; total cover (projection): 80%, E3 (tree layers): 40% (10-12 m; 6-10 m high), E2 (shrub layers): 60% (0.5-2 m; 2-5 m high); E1 (herb layer): 25% (60 cm average; 10-80 cm high); cover of ground litter (decaying pine needles): 90%; substrate: deep skeletal soil over ultramafic peridotite; plant identifications (revisions): Galium: F. Ehrendorfer (Vienna, Austria); Convolvulus, Petrorhagia, Reichardia: G. Bazos (Athens, Greece).

E3-beta (upper tree): Pinus halepensis 2b E3-alpha (lower tree): Pinus halepensis 2b E2-beta (upper shrub): Pinus halepensis 2a E2-alpha (lower shrub): Cotinus coggygria 2b, Arbutus unedo 2a, Cistus creticus 2a, Pinus halepensis 2a, Cistus salviifolius 1, Hypericum empetrifolium 1, Sarcopoterium spinosum 1, Phillyrea latifolia +, Pistacia terebinthus + (sapling), Rubus sanctus +, Smilax aspera +

E1: Brachypodium sylvaticum 2b, Alkanna graeca subsp. baeotica 2m, Carlina graeca 1, Briza minor +, Calystegia sp. +, Convolvulus cantabrica +, Dactylis glomerata subsp. hispanica +, Galium heldreichii +, Hypochaeris achyrophorus +, Lathyrus aphaca +, Muscari sp. +, Petrorhagia armerioides +, Plantago lanceolata +, Prunella laciniata +, Reichardia picroides +, Scorzonera serpentinica +, Viola sp. +, Crataegus azarolus r (seedling), Quercus coccifera r (seedling).

Syntaxonomic position: Quercetea ilicis Br.-Bl. ex A. & O. Bolos 1950; Quercetalia ilicis Br.-Bl. ex Molinier 1934; Alkanno baeoticae-Pinion halepensis Mucina & Dimopoulos in Mucina & al. 2009

Incl.: Erica verticillata-Pinus halepensis-Ges. sensu Krause & al. 1963 [Art. 3c].

This association represents low-altitude dry pine woodlands on serpentines of the island of Euboea (Greece). It has been described from peridotites of this island as "Erica verticillata-Pinus halepensis-Ges." by Krause & al. (1963). Our association matches best the column 1 (Sp. 1) of table 5 on page 361 of Krause & 0al. (1963) despite Erica manipuliflora (called E. verticillata by Krause & al., 1963) does not occur in our releve above. However, we found this species as abundant (often dominating) in the shrub layer of pine forests in other localities sampled on Euboea (Mucina & Dimopoulos, 2000a).

The description of this new association became necessary since the name Erico manipuliflorae-Pinetum halepensis (Brullo & al., 1997) is considered a nomen dubium (for details of the argument, see Mucina & Dimopoulos, 2000b) and thus could not be used as the type of the new alliance Alkanno baeoticae-Pinion halepensis (see below). The eponymous taxon Alkanna graeca Boiss. & Spruner in Boiss. subsp. baeotica (A. DC.) Nyman is distributed in the central eastern part of Greek mainland, Euboea, and in the mountains of the Peloponnese (Oligirtos, Taigetos, Parnonas) (Tan & Iatrou, 2001).

Alkanno baeoticae-Pinion halepensis Mucina & Dimopoulos all. nova hoc loco

Holotypus: Alkanno baeoticae-Pinetum halepensis Mucina & Dimopoulos in Mucina & al. 2009 (for the description of the association, see above).

Diagnostic taxa: Alkanna graeca subsp. baeotica, Alyssum euboeum, Centaurea mantoudi, Jurinea mollis subsp. anatolica, Onosma euboica, Quercus trojana subsp. euboica, Scorzonera serpentinica.

Syntaxonomic position: Quercetea ilicis Br.-Bl. ex A. & O. Bolos 1950; Quercetalia ilicis Br.-Bl. ex Mo linier 1934.

The description of this alliance is necessary for the nomenclatural reasons referred to above as well as for the fact that there was no appropriate high-rank syntaxon described earlier to accommodate the floristically and ecologically peculiar low-altitude dry pine woodlands on ultramafic substrates of the Aegean region.

The name Alyssion euboei was suggested by Brullo & al. (1997) as a new syntaxon to encompass the pine forests and associated scrubs on ultramafic peridotites (Erico manipuliflorae-Pinetum halepensis) as well as phrygana (Stachyo-Alyssetum euboei) and therophyte-rich communities (Malcolmio-Alyssetum densistellati; Thero-Brachypodietea). The Alyssion euboei has been rejected as a nomen dubium since the name of the next subordinate syntaxon supposed to typify the Alyssion euboei is considered a nomen dubium according to Art. 38 ICPN. The circumstances of this suggestion have been elaborated in detail in Mucina & Dimopoulos (2000b).

The Alkanno baeoticae-Pinion halepensis is a new alliance comprising Aegean Pinus halepensis forests on serpentine (ultramafic) substrates. It is, according to our present knowledge, distributed on the Greek island of Euboea (Evvia). However, we presume its occurrence also in the adjacent Greek mainland on appropriate geological substrates.

Artemisio albae-Brometalia erecti Ubaldi ex Denggler & Mucina ordo novo hoc loco

Validated name: Artemisio albae-Brometalia erecti (Biondi & al. 1995) Ubaldi 1997 (Ubaldi, 1997: 246) [Art. 5].

Holotypus: Xero-Bromion erecti Zoller 1954 (Zoller, 1954: 38 et seq.).

Syntaxonomic position: Festuco-Brometea Br.-Bl. & Tx. ex Klika & Hadac 1944.

Synonyms: Brometalia erecti Koch 1926 nom. amb. propos. p.p. [typo excl.]; Brometalia erecti Br.-Bl. 1936 nom. amb. propos. p.p. [typo excl.; Art. 31]; Xero-Brometalia Doing 1963 p.p. [Art. 8]; Brometalia erecti Br.Bl. 1936 nom. amb. propos. sensu Korneck 1974 [typo excl.]; Xero-Brometalia erecti Royer ex Dengler 1994 [Art. 8, 12(2)].

Incl.: Xero-Bromenalia Royer 1991 [Art. 12], Artemisio albae-Bromenalia erecti Biondi & al. 1995 [Art. 5].

Character taxa of the order (and of subordinate syntaxa): Achillea virescens, Argyrolobium zanonii, Allium sphaerocephalon, Arabis collina, Asperula purpurea, Avenula praetutiana, Bromus hordeaceus subsp. ferronii, Carduncellus mitissimus, Carex macrolepis, Carlina acanthifolia, Catananche caerulea, Centaurea ambigua, Chamaecytisus spinescens, Coronilla minima, Crupina vulgaris, Dianthus sylvestris, Dorycnium pentaphyllum, Erysimum pseudorhaeticum, Euphorbia myrsinites, Festuca circummediterranea, F. marginata, F. robustifolia, Fumana ericoides, Galium corrudifolium, G. lucidum, Globularia meridionalis, G. punctata, Helianthemum apenninum, H. canum, H. nummularium subsp. nummularium, Hippocrepis comosa, Inula spiraeifolia, Koeleria splendens, Linum tenuifolium, Muscari atlanticum, Ononis natrix subsp. natrix, O. pusilla, Orobanche teucrii, Petrorhagia saxifraga, Phleum ambiguum, Phyteuma orbiculare subsp. tenerum, Potentilla hirta, P. rigoana, Satureja montana, Seseli montanum, Sesleria nitida, Sideritis hyssopifolia subsp. guillonii, S. italica, Silene italica, Teucrium montanum, Thesium divaricatum, Thymus striatus, Tortella nitida, Trinia dalechampii.

Differential taxa of the order (against the other orders of the class): Artemisia alba, Carex halleriana, Cladonia convoluta, Convolvulus cantabrica, Helichrysum italicum, Koeleria vallesiana, Potentilla tabernaemontani, Teucrium capitatum.

The Artemisio albae-Brometalia erecti comprise the xerophytic basiphilous grasslands of Italy, France, southern England, Belgium, southwestern Germany, and northwestern Switzerland. According to the present knowledge, this order contains the alliances Xero-Bromion Zoller 1954 (Germany, Switzerland, Belgium, England and most of France), Festuco-Bromion Barbero & Loisel 1971 (Provence and Liguria), Artemisio albae-Saturejion montanae Allegrezza et al. 1997, Phleo ambigui-Bromion erecti Biondi et al. 1995, and probably also Alyssion bertolonii E. Pignatti & Pignatti 1977 and Cytiso-Bromion caprini Bonin in Barbero & Bonin 1969 (the last four on the Apennine Peninsula).

Traditionally, the Festuco-Brometea have been subdivided into geographically vicariant orders, mainly the suboceanic Brometalia erecti W. Koch 1926 and the subcontinental Festucetalia valesiacae Br.-Bl. & Tx. ex Br.-Bl. 1950, each of them comprising both xerophytic and meso-xerophytic alliances (e.g. Royer, 1991; Pott, 1995; Theurillat & al., 1995; Bardat & al., 2004; Biondi & al., 2005). More recently, it appeared that the meso-xerophytic alliances of both orders are floristically rather similar so that it does not seem reasonable to arrange them in two different orders. Thus, many recent overviews agree in joining the meso-xerophytic alliances of the Festuco-Brometea throughout Europe in one order Brometalia erecti W. Koch 1926 nom. amb. propos. (Brachypodietalia pinnati Korneck 1974) (e.g. Korneck, 1974, Mucina & Kolbek, 1993; Dengler 1994, 2004a; Rodwell & al., 2002; Dengler & al., 2003). While the meso-xerophytic basiphilous grasslands show a more or less continuous distribution through large parts of Europe, the xerophytic types occur in several widely isolated regions, where they have a clearly distinct floristic composition. Thus, it makes sense to keep them in several xerophytic orders, among others the Festucetalia valesiacae s.str. (subcontinental and continental steppes) and the Stipo pulcherrimae-Festucetalia pallentis Pop 1968 (prae-alpine-circumpannonian xerophytic grasslands on rock outcrops).

However, no order was available to encompass the xerophytic part of the former Brometalia erecti s.l., i.e. the xerophytic alliances occurring in the subatlantic-submediterranean part of the class range. Three previous attempts to establish an order with such content (Korneck, 1974; Dengler, 1994; Ubaldi, 1997) resulted in invalid or non-applicable names only (see list of synonyms above). Similarly, two suborders with the same content have been published only invalidly (Royer, 1991; Biondi & al., 1995). The latter two works as well as Biondi & al. (2005) comprise extensive synoptic tables that show the floristic bipartition between the xerophytic and the meso-xerophytic alliances of the former Brometalia erecti s.l., and thus support our concept to separate them on order level. Thus, we validate here the most suitable of the available names, Artemisio albae-Brometalia erecti. It was first described as a suborder by Biondi & al. (1995), but this description was invalid because the authors did not provide an unambiguous reference to the protologue of the selected type alliance. Neither Ubaldi (1997), who raised the suborder to ordinal level, nor Biondi & al. (2005) did correct this mistake.

The presented list of diagnostic species is mainly based on some large-scale overviews (Royer, 1991; Biondi & al., 1995, 2005; Dengler, unpubl.). It is somehow preliminary because we lack a comprehensive Europe-wide synthesis of dry grassland communities that would allow establishing statistically tested diagnostic species. While the core of the new order seems to be rather clear, its precise delimitation both against other xerophytic orders of the class Festuco-Brometea (Festucetalia valesiacae, Brachypodion phoenicoidis Br.Bl. ex Molinier 1934, Scorzonero-Chrysopogonetalia Horvatic et Horvat 1958) and against "neighbouring" classes (Festuco hystricis-Ononidetea striatae RivasMart. & al. 2002, Rosmarinetea officinalis Rivas-Mart. & al. 2002) needs to be clarified in the future.

Dictamno albi-Ferulagion galbaniferae (van Gils & al. 1975) de Foucault & al. ex Carni & Dengler all. nova hoc loco

Validated name: Dictamno-Ferulagion galbaniferae (van Gils & al. 1975) de Foucault & al. 1983 (de Foucault & al., 1983: 449) [Art. 5].

Basionym: Dictamno albae-Ferulagenion galbaniferae van Gils & al. 1975 (van Gils & al., 1975: 57 et seq.; "Dictamno albae-Ferulagion galbaniferae suball. nov. ").

Lectotypus [Art. 27a, selected by Carni (1997: p. 218)]: Libanotido-Laserpitietum sileris van Gils & al. 1975 (van Gils & al., 1975: 49).

Syntaxonomic position: Trifolio-Geranietea sanguinei T. Muller 1962; Antherico ramosi-Geranietalia sanguinei Julve ex Dengler in Dengler & al. 2003.

Synonyms: Geranion sanguinei Tx. in T. Muller 1961 p.min.p. [Art. 8]; Geranion sanguinei Tx. ex T. Muller 1962 p.min.p. [typo excl.]

Character taxa of the alliance (and of subordinate syntaxa): Betonica officinalis, Chamaecytisus supinus, Clematis recta, Coronilla coronata, Cyanus triumfettii, Dictamnus albus, Ferulago campestris, Galium lucidum, Hippocrepis emerus subsp. emeroides, Inula hirta, Knautia drymeia subsp. tergestina, K. fleischmannii, K. illyrica, Laserpitium siler, Lathyrus latifolius, L. pannonicus, Lilium bulbiferum, Melampyrum velebiticum, Orobanche lutea, Paeonia officinalis, Potentilla alba, Pulmonaria angustifolia, Ruta graveolens, Selinum silaifolium, Trifolium rubens, Veronica barrelieri, V. jacquinii.

Differential taxa of the alliance (particularly against Geranion sanguinei): Anthericum ramosum, Brachypodium rupestre, Bromus erecta, Buphthalmum salicifolium, Carex humilis, Cirsium pannonicum, Filipendula vulgaris, Teucrium chamaedrys.

The Dictamno albi-Ferulagion galbaniferae comprises xerophytic forest-edge communities of base-rich soils mainly on the Balkans, but with some associations reaching the southern-eastern and eastern margins of the Alps. So far, the majority of associations have been described from Slovenia (van Gils & al., 1975; Carni 1997, 1998, 2005). The named diagnostic taxa are based on a comprehensive synthesis of Trifolio-Geranietea sanguinei in Europe (Dengler & al., 2006a). They comprise both geoelements of the Illyric-Dinaric region and widespread forest-edge and dry grassland species that are much more frequent in stands of the Illyric-Dinaric region than in central Europe.

Originally, these communities have been placed in the Geranion sanguinei within the suballiance Dictamno albae-Ferulagenion galbaniferae. However, several large-scale syntheses showed that one alliance Geranion sanguinei s.l. is too less to encompass the large variety of different xerophytic, basiphilous forestedge communities of Europe (e.g. de Foucault & al., 1983; Dengler & al., 2006a). Thus, there have been various proposals to split the Geranion sanguinei s.l. into several vicariant alliances, namely the Geranion sanguinei s.str. of central and western Europe, the Galio littoralis-Geranion sanguinei Gehu & Gehu-Franck in de Foucault & al. 1983 of northern Europe (including Fennoscandia, Baltic countries, British Isles and temperate sea coasts), the Origanion virentis Rivas-Mart. & O. Bolos in Rivas-Mart. & al. 1984 of the Iberian Peninsula and the Dictamno albi-Ferulagion galbaniferae of the Balkan Peninsula (de Foucault & al., 1983; Rodwell & al., 2002; Dengler & al., 2003, 2006a; Dengler, 2004b). As no valid name was available for the last alliance so far, we validate the name proposed by de Foucault & al. (1983) here. To reflect the fact that the traditional Geranion sanguinei s.l. has now been split into four floristically and ecologically similar, vicariant alliances, it has been suggested to place them into a new order Antherico ramosi-Geranietalia sanguinei as opposed to the Origanetalia vulgaris s.str., which correspond to the traditional Trifolion medii T. Muller s.l. and consist of the Trifolion medii s.str. and the Knaution dipsacifoliae Julve ex Dengler & Boch 2008 (Dengler, 2003: 190-192, 2004b; Dengler & al., 2003, 2006a; Dengler & Boch, 2008).

Euphorbio taurinensis-Geranion lucidi Matevski & Carni all. nova hoc loco

Validated name: Euphorbio taurinensis-Geranion lucidi Matevski & Carni 2001 (Matevski & Carni, 2001: 55-57) [Art. 8].

Holotypus: Veronico cymbalariae-Geranietum lucidi Matevski & Carni 2003 (Matevski & Carni, 2003: 77).

Syntaxonomic position: Stellarietea mediae Tx. & al. ex von Rochow 1951; Geranio-Cardaminetalia hirsutae Brullo in Brullo & Marceno 1985.

Character taxa of the alliance: Calamintha vardarensis, Euphorbia taurinensis, Lamium garganicum subsp. laevigatum.

Matevski & Carni (2001) suggested the alliance Euphorbio taurinensis-Geranion lucidi to encompass short-lived nitrophilous fringe vegetation of Macedonia (Republic of Macedonia and Greece), but failed to provide a nomenclatural type, which rendered the description invalid according to Art. 8 ICPN. Later, Matevski & Carni (2003) described the same vegetation type as Veronico cymbalariae-Geranietum lucidi, but assigned it to the west-Mediterranean Geranio-Anthriscion Rivas-Mart. 1978. Here, we validate the name Euphorbio taurinensis-Geranion lucidi as we now came to the conclusion that the therophyte-rich nitrophilous fringe communities of the west-Mediterranean and of the Balkans show floristic and ecological peculiarities that should be recognised at the level of alliance.

Gentianello amarellae-Helictotrichion pratensis Royer ex Dengler all. nova hoc loco

Validated name: Gentianello amarellae-Avenulion pratensis Royer 1991 (Royer, 1991: 88-91 and 199) [Art. 3b].

Holotypus: Helictotricho-Caricetum flaccae Shimwell 1971 (Shimwell, 1971: 39 et seq.) [holoty pus].

Syntaxonomic position: Festuco-Brometea Br.-Bl. & Tx. ex Klika & Hadac 1944; Brometalia erecti W. Koch 1926 nom. amb. propos. [Brachypodietalia pinnati Korneck 1974].

Synonyms: Bromion erecti W. Koch 1926 sensu auct. p.min.p. [typo excl.]; Bromion erecti Br.-Bl. 1936 p.p. [Art. 8, 31]; Meso-Bromion Oberd. 1949 sensu auct. p.min.p. [typo excl.]; Gentiano amarellaeAvenulion pratensis Royer ["(Willems 1982) Royer 1987"] ex Julve 1993 [Art. 5].

Incl.: British Isles subgroup [Meso-Bromion] sensu Willems 1982.

Character taxa of the alliance: Blackstonia perfoliata, Gentianella amarella, Thymus praecox subsp. arcticus.

Differential taxa of the alliance (against all other alliances of the order): Centaurea nigra, Hypochaeris radicata, Scleropodium purum, Succisa pratensis, Trifolium repens.

Differential taxa of the alliance (against Filipendulo vulgaris-Helictotrichion pratensis): Anacamptis pyramidalis, Asperula cynanchica, Ctenidium molluscum, Festuca guestfalica subsp. guestfalica, Helianthemum nummularium subsp. obscurum, Koeleria macrantha, Leontodon hispidus, Polygala vulgaris, Sanguisorba minor, Sesleria albicans.

Differential taxa of the alliance (against Bromion erecti and other southern alliances): Agrostis capillaris, Danthonia decumbens, Dicranum scoparium, Euphrasia nemorosa, Festuca ovina, F. rubra, Helictotrichon pratense, Luzula campestris.

The Gentianello amarellae-Helictotrichion pratensis comprises meso-xerophytic basiphilous grasslands of the British Isles and northern France. Traditionally, these communities have been included in a widely delimited Bromion erecti (Meso-Bromion) (e.g. Shimwell, 1971). However, the meso-xerophytic basiphilous grasslands of the British Isles, Denmark, and Fennoscandia differ significantly in their floristic composition from their counterparts further south (Willems, 1982; Royer, 1991; Dengler & al., 2003, 2006b). In particular, they (largely) lack many otherwise widespread Festuco-Brometea species, such as Eryngium campestre, Euphorbia cyparissias, Koeleria pyramidata, Prunella grandiflora, Teucrium chamaedrys, while several slightly acidophilous graminoids and mosses become more frequent to the north (e.g. Dengler & al., 2006b; see list of differential species above). This pattern prompted Willems (1982) to separate a northwest European group within the MesoBromion, which he further subdivided in a British Isles subgroup and a south Scandinavian subgroup. Royer (1991) was the first to give a formal syntaxon name to the first subgroup, namely Gentianello amarellaeAvenulion pratensis, though only provisionally and thus invalidly. In a comprehensive European synthesis of Festuco-Brometea, I could confirm this pattern of mostly negatively differentiated community types of the meso-xerophytic basiphilous grasslands (i.e. the order Brometalia erecti = Brachypodietalia pinnati) at their northern distributional limits, which can be subdivided in a western group (British Isles and N France) and a eastern group (Fennoscandia, Denmark, N Germany, Baltic countries) (Dengler, unpubl.; see Dengler, 2003: 199-201 and 222-223; Dengler & al., 2003). While the latter has been formally described as Filipendulo vulgaris-Helictotrichion pratensis Dengler & Lobel in Dengler & al. 2003 (Dengler & al., 2003), the first group still lacked a valid name though the alliance concept and name of Royer (1991) was widely in use (e.g. Julve, 1993; Rodwell & al., 2002; Bardat & al., 2004). Accordingly, Royer's name is validated here.

Helichryso barrelieri-Phagnalion graeci (Barbero & Quezel 1989) R. Jahn stat. nov. hoc loco

Basionym: Helichryso barrelieri-Phagnalenion graeci Barbero & Quezel 1989 nom. corr. R. Jahn hoc loco

Corrected name: Helichryso orientali-Phagnalenion graeci Barbero & Quezel 1989 (Barbero & Quezel 1989: 44).

Holotypus [Art. 27a]: Micromerio graecae-Hypericetum empetrifolii Barbero & Quezel 1989 (Barbero & Quezel 1989: 58).

Syntaxonomic position: Cisto-Micromerietea julianae Oberd. 1954; Poterietalia spinosi Eig 1939.

Character taxa of the alliance: Cistus salviifolius, C. creticus, Erica manipuliflora, Genista acanthoclada, Hypericum empetrifolium subsp. empetrifolium, Satureja thymbra, Helichrysum stoechas subsp. barrelieri.

The correction of the suballiance name, which forms the basionym for the new alliance, is necessary, because Helichrysum orientale (L.) DC., a conspicuous, obligate Aegean chasmophyte (Zaffran, 1990), does not occur in phrygana except in ungrazed areas and in the neighbourhood of rocks. It is very unlikely that this species occurred in eight out of 18 phrygana releves, as given by Barbero & Quezel (1989: Table 1). Helichrysum stoechas subsp. barrelieri is a similar, but smaller species, known to be a common phrygana plant, which occurs in 29% of 957 phrygana releves made by the author on Crete. This plant is also given in the tables of Barbero & Quezel (1989), but with lower frequency and only in those releves without "Helichrysum orientale". There is no doubt that the records of the two Helichrysum "species" in Barbero & Quezel (1989) must be united under the name Helichrysum stoechas subsp. barrelieri. According to Art. 43 ICPN, the name of the suballiance must be corrected.

Oberdorfer (1954) established the class Cisto-Micromerietea julianae with three alliances based on releves from northern Greece. The alliance Cistion orientale (Oberdorfer 1954, for nomenclatural implications, see Hyperico olympici-Cistion cretici below) represents the class on non-calcareous substrates, while the Micromerion julianae oberdorfer 1954 (incl. Dorycnio-Coridothymion capitati [Oberd. 1954] Brullo & al. 1997) occurs on calcareous substrates (Oberdorfer 1954).

Due to considerable differences in floristic composition, Barbero & Quezel (1989) separated the phrygana communities of central and southern Greece from those of northern Greece. The more southerly phrygana communities were established as a new alliance Hyperico empetrifolii-Micromerion graecae Barbero & Quezel 1989. Based on an extensive set of new releves (Jahn, unpubl.), it is reasonable to subdivide the southerly communities further into one alliance on noncalcareous substrates and one on limestone. The former is newly described here as Helichryso barrelieri-Phagnalion graeci. It is characterized by the absence of Euphorbia acanthothamnos, Phlomis fruticosa, Ballota acetabulosa, and many other subnitrophilous shrubs and herbs, as well as by the presence of the taxa given above, with low nitrogen indicator values between 1 and 3 according to Bohling & al. (2002).

The new alliance contains two of the three suballiances distinguished by Barbero & Quezel (1989) within the Hyperico empetrifolii-Micromerion graecae, the Helichryso barrelieri-Phagnalenion graeci on marly substrates, and the Hyperico empetrifoliiMicromerienion graecae nom. inval. (Art. 3m, 8 ICPN) on slightly acid, igneous substrates (Barbero & Quezel, 1989). By contrast, the suballiance Phlomido fruticosae-Euphorbienion acanthothamni nom. inval. (Art. 3m ICPN), which holds the communities on limestone, remains in the Hyperico empetrifolii-Micromerion graecae s.str. (holotypus of alliance and suballiance is the Dorycnio hirsuti-Micromerietum graecae Barbero & Quezel 1989).

The Helichryso barrelieri-Phagnalion graeci consists of phrygana communities rich in Cistus, Erica, and Genista species. The alliance occurs on non-calcareous substrates, such as granite, gneiss, and phyllitic schists, on serpentine, on hard and soft marls, on volcanic soils, and on sand. Its distribution range includes the Ionian and the Aegean sides of the Greek mainland, the Central and South Aegean islands and the Aegean coast of Anatolia northward to approx. 39 N (Barbero & Quezel, 1989; Heiselmayer & al., 1995; de Bolos & al., 1996; Brullo & al., 1997), i.e. a region where vegetation linked to the thermomediterranean belt is widely distributed (see map in Quezel & Barbero, 1985). Nevertheless, the communities are not strictly limited to low altitudes. In general, releves of the Helichryso barrelieri-Phagnalion graeci have fewer species than those of the Hyperico empetrifolii-Micromerion graecae s.str.

From the Cistion orientale (Hyperico olympici-Cistion cretici, see below), the Helichryso barrelieri-Phagnalion graeci is differentiated by the absence or the retreat of Chrysopogon gryllus, Festuca spp., Thymus comptus, Centaurea salonitana, and further taxa with lower frequencies (oBERDoRFER, 1954). Releves from the Pilion peninsula (RAuS 1979) contain species from both syntaxa, but the diagnostic taxa of the Helichryso barrelieri-Phagnalion graeci have a distinctly higher constancy and frequency. Thus, they are placed in the new alliance.

REPLACEMENT OF AN ILLEGITIMATE SYNTAXON NAME

Hyperico olympici-Cistion cretici (Oberd. 1954) R. Jahn & Bergmeier nom. nov. hoc loco

Basionym: Cistion orientale oberd. 1954 (oBERDORFER, 1954: p. 92) [Art. 34a].

Synonym: Cisto-Hypericion bithynici (oberd. 1954) Brullo & al. 1997 (Brullo & al., 1997: 34) [Art. 3g, 43].

Lectotypus [selected by Brullo & al. (1997: 34)]: Calicotomo-Cistetum cretici oberd. 1954 (oBERDoRfer, 1954: 94; "Calycotometo-Cistetum villosae").

Syntaxonomic position: Cisto-Micromerietea julianae Oberd. 1954; Poterietalia spinosi Eig 1939.

OBERDORFER (1954), when establishing the class Cisto-Micromerietea julianae, described among others the alliance Cistion orientale, phrygana communities on non-calcareous substrates, based on releves from northern Greece. Cistion orientale is an illegitimate name (Art. 34a ICPN), which was rejected by Brullo & al. (1997) and substituted by Cisto-Hypericion bithynici. The latter name is invalid, though, as the authors did not specify the Cistus species involved (Art. 3g ICPN). Oberdorfer (1954: 95, "Liste III") indicated three species of Cistus, with C. creticus (as C. villosus) being the most constant one. Moreover, despite the fact that the name Hypericum bithynicum appears in Oberdorfer's original releves (Oberdorfer, 1954: 95; see also Horvat & al., 1974: 120), the Cisto-Hypericion bithynici is based on a misidentification (Art. 43 ICPN). H. bithynicum, a species of damp scrub and woodland, occurs only in Turkey and has not been found in Greece (Greuter & al., 1986).

According to Art. 39 ICPN, we propose to replace the illegitimate name Cistion orientale with Hyperico olympici-Cistion cretici as a nomen novum. Hypericum olympicum L., eponymous taxon of our new name, occurs in dry open vegetation on lime-poor, rocky, and sandy soils (e.g. Raus, 1979; Bergmeier, unpubl.) and is mentioned by Oberdorfer (1954: 95) to occur in the Cistion orientale, albeit with low constancy. It is largely confined to the Greek mainland and adjacent areas of the Republic of Macedonia, Serbia, Bulgaria, and Turkey. In Greece, this very variable species is most widespread in the northern parts of the country.

ACKNOWLEDGEMENTS

We wish to thank T. Raus (Berlin) for correcting nomenclature of some taxa. F. Ehrendorfer (Vienna) and G. Bazos (Athens) kindly identified or revised some plant collections from Euboea. This paper is a result of cooperation within the "European Vegetation Survey" (EVS)--a working group of the International Association for Vegetation Science (IAVS).

Recibido: 1 de abril de 2009 Aceptado: 15 de abril de 2009

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Ladislav Mucina (*), Jurgen Dengler (**), Erwin Bergmeier (***), Andraz Carni (****), Panayotis Dimopoulos (*****), Ralf Jahn (******) & Vlado Matevski (*******)

* Department of Environmental and Aquatic Sciences. School of Agriculture and Environment. Curtin University of Technology. GPO Box U1987. Perth, WA 6845. Australia. E-mail: L.Mucina@curtin.edu.au.

** Plant Systematics and Vegetation Ecology. Biocentre Klein Flottbek. University of Hamburg. Ohnhorststr. 18. D-22609 Hamburg. Germany. E-mail: dengler@botanik.uni-hamburg.de.

*** Albrecht von Haller Institute of Plant Sciences. University of Gottingen. Untere Karspule 2. D-37073 Gottingen. Germany. E-mail: erwin.bergmeier@bio.uni-goettingen.de.

**** Institute of Biology. Scientific Research Center of the Slovenian Academy of Sciences and Arts. Novi trg 2. P. Box 306. SI-1001 Ljubljana. Slovenia. E-mail: carni@zrc-sazu.si.

***** Department of Environmental and Natural Resources Management. University of loannina. Seferi 2. GR-30100 Agrinio. Greece. E-mail: pdimopul@cc.uoi.gr.

****** Roschenhohe 8. D-09603 GroBschirma. Germany. E-mail: jahnralf@hotmail.com.

******* institute of Biology. Faculty of Natural Sciences and Mathematics. Gazi baba b/b. p.b. 162. MK-91000 Skopje. Republic of Macedonia. E-mail: vladom@pmf.ukim.mk.
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Author:Mucina, Ladislav; Dengler, Jurgen; Bergmeier, Erwin; Carni, Andraz; Dimopoulos, Panayotis; Jahn, Ral
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Date:Jan 1, 2009
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