LAZAROA 36: 107-129. 2015 http://dx.doi.org/10.5209/rev_LAZA.2015.v36.51255 ISSN: 0210-9778

Validations and descriptions of European syntaxa of vegetation dominated by , bryophytes and algae

Helga Bültmann1, Claude Roux2, José Maria Egea3, Philippe Julve4, Oliver Bricaud5, Giuseppe Giaccone6, Lothar Täuscher7, Marijke Creveld8, Vincenzo Di Martino9, Stjepko Golubiç10 & Nozomu Takeuchi11

Abstract: Bültmann, H., Roux, C., Egea, J.M., Julve, P., Bricaud, O., Giaccone, G., Täuscher, L., Creveld, M., Di Martino, V., Golubiç, S. & Takeuchi, N. Validations and descriptions of European syntaxa of vegetation dominated by lichens, bryophytes and algae. Lazaroa 36: 107-129 (2015).

Fourty-two high-rank syntaxa and seven associations of the thallophyte system of syntaxa are either described as new or validated in this paper. Among those, there are the following nine classes: Aspicilietea candidae, Caulerpetea racemosae, Desmococcetea olivacei, Entophysalidetea deustae, Gloeocapsetea sanguineae, Mesotaenietea berggrenii, Naviculetea gre- gariae, Porpidietea zeoroidis, Roccelletea phycopsis. Eleven orders and ten alliances as well as three associations are des- cribed or validated: the Aspicilietalia verruculosae (incl. Aspicilion mashiginensis and Teloschistion contortuplicati), the Caulerpetalia racemosae (incl. Caulerpion racemosae), the Desmococcetalia olivacei (incl. Desmococcion olivacei), the Dirinetalia massiliensis, the Fucetalia vesiculosi (incl. Ascophyllion nodosi), the Gloeocapsetalia sanguineae, the Lecideetalia confluescentis (incl. Lecideion confluescentis), the Mesotaenietalia berggrenii (incl. Mesotaenion berggrenii, Mesotaenietum berggrenii and Chloromonadetum nivalis), the Naviculetalia gregariae (incl. Oscillatorion limosae and Oscillatorietum limosae), the Porpidietalia zeoroidis (incl. Porpidion zeoroidis), and the Roccelletalia fuciformis (incl. Paralecanographion grumulosae). Further, five orders, seven alliances and four associations, classified in known classes, were described as well. These include: the Bacidinetalia phacodis, the Agonimion octosporae and the Dendrographetalia decolorantis (all in the Arthonio radiatae-Lecidelletea elaeochromae), the Staurothelion solventis (in the Aspicilietea lacus- tris), the Pediastro duplicis-Scenedesmion quadricaudae and the Pediastro duplicis-Scenedesmetum quadricaudae (both in the Asterionelletea formosae), the Peccanion coralloidis and the Peltuletalia euplocae (both in the Collematetea cristati), the Laminarion hyperboreae, the Saccorhizo polyschidi-Laminarietum and the Alario esculenti-Himanthalietum elongatae (all in the Cystoseiretea crinitae), the Delesserietalia sanguinei, the Delesserion sanguinei and the Delesserietum sangui- neae (all in the Lithophylletea soluti), as well as the the Rinodino confragosae-Rusavskietalia elegantis and the Rhizocarpo geographici-Rusavskion elegantis (both in the Rhizocarpetea geographici).

Keywords: algal vegetation, bryophyte vegetation, cryptogams, International Code of Phytosociological Nomenclature, vegetation, phytosociology, syntaxonomy, vegetation of Europe.

1 Michaelweg 40, D-48149 Münster, Germany. Email: [email protected] 2 390 chemin des Vignes-Vieilles, F-84120 Mirabeau, France. Email: [email protected], claude.roux@licheno- logue.org 3 Departamento de Biología Vegetal, Area de Botánica, Facultad de Biología, Campus Universitario de Espinardo, E-30100 Murcia, Spain. Email: [email protected] 4 Laboratoire Ecologie et Biodiversité, FGES - Université Catholique de Lille, 83 boulevard Vauban, Espace Saint-Raphaël, Bâtiment G, F-59016 Lille, France. Email: [email protected], [email protected] 5 57 rue du mas de Pialon, F-13750 Plan-d’Orgon, France; [email protected] 6 Dipartimento di Botanica, Università di Catania, Contrada Rini 10, I-90020 Baucina (PA), Italy. Email: [email protected] 7 Institut für angewandte Gewässerökologie GmbH, Schlunkendorfer Straße 2e, D-14554 Seddiner See, Germany. Email: [email protected] 8 The Netherlands. Email: [email protected] 9 ISAFOM, Consiglio Nazionale Ricerche, OU Catania, Catania, Italy. Email: [email protected] 10 Biological Science Center, Boston University, 5 Cummington str, Boston, MA 02215, USA. Email: [email protected] 11 Department of Earth Sciences, Graduate School of Science, Chiba University, 1-33, Yayoicho, Inage-ku, Chiba 263-8522, Japan. Email: [email protected]

107 LAZAROA 36: 107-129. 2015 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes

Resumen: Bültmann, H., Roux, C., Egea, J.M., Julve, P., Bricaud, O., Giaccone, G., Täuscher, L., Creveld, M., Di Martino, V., Golubiç, S. & Takeuchi, N. Descripción y validación de distintos sintáxones europeos de la vegetación domi- nada por briófitos, líquenes y algas. Lazaroa 36: 107-129 (2015).

Se describen por primera vez o se validan un total de 42 sintáxones de alto rango y 7 asociaciones correspondientes a distintos tipos de vegetación talofítica y briofítica. Entre ellos se incluyen las 9 clases siguientes: Aspicilietea candidae, Caulerpetea racemosae, Desmococcetea olivacei, Entophysalidetea deustae, Gloeocapsetea sanguineae, Mesotaenietea berggrenii, Naviculetea gregariae, Porpidietea zeoroidis, Roccelletea phycopsis. Once órdenes y diez alianzas, además de tres asociaciones pertenecientes a estas clases nuevas, se describen por primera vez o se validan: Aspicilietalia verruculo- sae (incl. Aspicilion mashiginensis y Teloschistion contortuplicati), Caulerpetalia racemosae (incl. Caulerpion racemosae), Desmococcetalia olivacei (incl. Desmococcion olivacei), Dirinetalia massiliensis, Fucetalia vesiculosi (incl. Ascophyllion nodosi), Gloeocapsetalia sanguineae, Lecideetalia confluescentis (incl. Lecideion confluescentis), Mesotaenietalia berg- grenii (incl. Mesotaenion berggrenii, Mesotaenietum berggrenii y Chloromonadetum nivalis), Naviculetalia gregariae (incl. Oscillatorion limosae and Oscillatorietum limosae), Porpidietalia zeoroidis (incl. Porpidion zeoroidis), y Roccelletalia fuci- formis (incl. Paralecanographion grumulosae). Además, otros cinco órdenes, siete alianzas y cuatro asociaciones pertene- cientes a clases ya descritas se proponen como nuevos: Bacidinetalia phacodis, Agonimion octosporae y Dendrographetalia decolorantis (clase Arthonio radiatae-Lecidelletea elaeochromae); Staurothelion solventis (clase Aspicilietea lacustris); Pediastro duplicis-Scenedesmion quadricaudae y Pediastro duplicis-Scenedesmetum quadricaudae (clase Asterionelletea formosae); Peccanion coralloidis y Peltuletalia euplocae (clase Collematetea cristati); Laminarion hyperboreae, Saccorhizo polyschidi-Laminarietum y Alario esculenti-Himanthalietum elongatae (clase Cystoseiretea crinitae); Delesserietalia sanguinei, Delesserion sanguinei y Delesserietum sanguineae (clase Lithophylletea soluti); y Rinodino con- fragosae-Rusavskietalia elegantis y Rhizocarpo geographici-Rusavskion elegantis (clase Rhizocarpetea geographici).

Palabras clave: vegetación algal, vegetación briofítica, vegetación liquénica, criptógamas, Código Internacional de Nomenclatura Fitosociológica, fitosociología, sintaxonomía, vegetación de Europa.

INTRODUCTION communities remains very rudimentary (e.g. PIRK & TÜXEN, 1955, 1957; MICHAEL & al., 1985). Thallophyte vegetation supposedly had occu- Most monographs deal either with bryophyte, pied terrestrial habitats long before vascular lichen or algae vegetation. Works integrating all land plant vegetation evolved (BOMFLEUR & al., thallophyte groups are quite rare (e.g. 2010). Vegetation of bryophytes, lichens and BARKMAN, 1958). Recent large-scale surveys algae has been subject of description using the exist only for bryophyte syntaxa (MARSTALLER, BRAUN-BLANQUET (1964) approach since the 2006). Still, the thallophyte vegetation is being early days of the method (e.g. FREY, 1923; studied by specialists only, and it has rarely been HILITZER, 1925; MESSIKOMMER, 1927; OCHSNER, recognized in vegetation studies. It is often dis- 1928; earlier syntaxonomic surveys by KLIKA & regarded since it is limited to special habitats HADAC, 1944; KLIKA, 1948, KLEMENT, 1955). and taxonomic expertise is insufficient; usually While the vegetation of bryophytes, lichens and only a limited number of the larger and more macroalgae is reasonably well known, the vege- conspicuous terricolous thallophyte species is tation of microalgae has been studied rarely: included in relevé species lists. MARGALEF (1944) studied the vegetation of The status of the thallophyte communities has freshwater algae in Spanish mountains, been discussed in detail by expert vegetation GOLUBIÇ (1967) investigated the vegetation of ecologists familiar with vegetation at different aerophytic microalgae on calcareous rocks, scales (WILMANNS, 1966; BARKMAN, 1973). In TÄUSCHER (1998) presented a synopsis for bent- particular BARKMAN (1973: 460) analysed different hic and planctonic freshwater algal vegetation approaches used for vegetation description at of the lowlands, and KHAYBULLINA & al. (2004, different scales. In Dutch juniper scrub he distin- 2005, 2005a) described the vegetation of soil guished between microcoena delimited by micro- algae. Even vegetation of macrofungi has been habitat and synusiae that were floristically defined. described that way, however the system of fungal He proposed to describe the syntaxa synusiae by

LAZAROA 36: 107-129. 2015 108 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes applying the Braun-Blanquet-approach and Epiphytic and epixylic communities depend on nomenclature the same way as for the classifica- chemistry and structure of the bark and wood, but tion of phytocoena. The terms synusia, union are rarely specific to a tree species. These com- and federation for community, association and munities are usually excluded from the studies of alliance are still occasionally applied (PAUS, forest vegetation. SCHUHWERK (1986) showed a 1997), however always with the agreement that possibility to include thallophytic communities in such assemblages should be described according tables of forest relevés by treating them as to the rules of the ICPN (WEBER & al., 2000). ‘species’. Unfortunately this useful approach has While the terminology of the units has still not been followed by others. not been fixed, researchers mostly followed the Terricolous thallophytes use the same sub- procedures of the Braun-Blanquet approach: strate as vascular plants, but have no roots and studied relevés and described floristically are poikilohydric. They grow on the surface or delimited units for the thallophyte vegetation occasionally in the pores of the soil (microal- using the rank-indicating terminations of the gae). The thallophytes can grow (1) as single ICPN (-etum, -ion, -alia, -etea). Thallophyte thalli or tiny patches within a phytocoenose, (2) syntaxa based on spatially delimitated relevés, as synusiae of single growth form, and (3) as classified by floristic criteria and using ranks stands of bryophytes, lichens and/or algae in corresponding to those of the phytocoena are spatially delimitated spots with scarce or no governed by the ICPN (WEBER & al., 2000: cover of vascular plants and often in gaps in the Definition I). Independently, authors found it vegetation. Only the latter have been described necessary to distinguish between floristically as thallophyte syntaxa. well-delimited thallophyte coenoses and ‘true’ The thallophyte syntaxa of bryophytes, synusiae of one-layered stands of few species lichens and algae have been compiled for the with identical growth-form occurring within first Europe-wide checklist of high ranked syn- phytocoenoses (e.g. BÜLTMANN, 2005 for rein- taxa including the bryophyte, lichen and algal deer lichen stands in the understorey of pine syntaxa (EuroVegChecklist: MUCINA & al., in forests; MARSTALLER, 2006 for the bryophyte press). This checklist will include 27 classes, 53 vegetation of the Rhytidiadelphion squarrosi orders and 137 alliances of bryophyte and lichen Waldheim 1944). Therefore the use of terms of syntaxa and 13 classes, 24 orders and 53 the synusial system for ‘good’ thallophyte syn- alliances of algal syntaxa (see Table 1 for an taxa seems not advisable anymore. Following overview). Of those syntaxa nine classes, 16 BARKMAN (1973), the term micro-communities, orders and 17 alliances, almost 14%, were not micro-associations etc. might be most suitable. yet validly described according to the ICPN In some cases, micro-communities can be very (WEBER & al., 2000) and therefore are to be extensive though, e.g. in arctic or mountain described here, alongside with seven new asso- regions. ciations. The extensive number of already described thallophyte syntaxa shows that construction of a meaningful syntaxonomic system for the thallo- A NEW ALLIANCE IN THE ASPICILIETEA phytic vegetation is feasible (see also LACUSTRIS WILMANNS, 1998). The thallophyte communities growing on The Aspicilietea lacustris Wirth 1972 includes both natural and artificial rock surfaces are well only one order – the Hydroverrucarietalia Èer- defined often species-rich. Yet even the most nohorský et Hadaè ex Klement 1955. The vege- common types have rarely been addresses in tation on acidic rocks is rather well studied, but syntaxonomic studies because of taxonomic the syntaxa from calcareous substrates are less problems, especially the rather time-consuming known. ASTA & al. (1973a) described the identification of constituent species. Staurotheletum solventis from submerged cal-

109 LAZAROA 36: 107-129. 2015 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes careous rocks of the high-mountain regions of bird rocks in the subalpine to nival belts and in France that cannot be placed within any current- the Arctic, and the thermophilous Physcion ly recognised alliance. The Staurothelion sol- dimidiatae Wirth 1972. ventis is described (see below) to accomodate The validation of the order became necessary the aquatic lichen communities on calcareous because CREVELD (1981) chose the invalidly rocks from high mountains, in addition to the published Rhizocarpo-Xanthorion Creveld 1981 Verrucarion siliceae Wirth 1972, the Aspicilion as the type alliance, hence rendering the order lacustris Klement 1950, and the Porinion lectis- invalidly described as well. simae Wirth 1980 from acidic substrates. The The conspicuous orange stands of Rusavskia Staurothelion solventis includes the type associ- elegans (formerly better known as Xanthoria ele- ation and one provisionally described communi- gans) are typical for sites with a strong influence ty (ASTA & al., 1973a). of bird manure on subacidic and calcareous rocks. Similar communities on calcareous rocks, however, belong in the Verrucarietea nigrescentis. Staurothelion solventis Roux all. nov. hoc loco (Hydroverrucarietalia, Aspicilietea lacustris) Rinodino confragosae-Rusavskietalia elegan- TYPE: Holotypus hoc loco: Staurotheletum sol- tis Creveld ordo nov. hoc loco ventis Asta, Clauzade et Roux 1973 (ASTA & al., (Rhizocarpetea geographici) 1973a: 75-81). CHARACTER TAXA OF THE ALLIANCE: Placynthium VALIDATED NAME: Rinodino confragosae- tantaleum, Polyblastia ardesiaca, Staurothele Xanthorietalia elegantis Creveld 1981 (CREVELD, sapaudica, S. solvens. 1981: 87) [ICPN Art. 17] Xanthoria elegans (Link) Th. Fr. is a synonym of Rusavskia elegans. SYNONYMS: Neuropogonetalia Mattick 1951 A NEW ORDER AND AN ALLIANCE IN [Art. 2b], Physcietalia caesiae Mattick 1951 THE RHIZOCARPETEA GEOGRAPHICI p.p. [Art. 2b], Physcietalia Hadac 1962 [Art. The species diversity of lichens on siliceous 2b], Physcietalia caesiae Mattick ex Creveld rocks is higher than on any other substrate 1981 [Art. 17]. TYPE: Holotypus hoc loco: Rhizocarpo geo- (BÜLTMANN, 2010). The Rhizocarpetea geo- graphici Wirth 1972 is a very large class, com- graphici-Rusavskion elegantis Creveld in prising six orders (Rhizocarpetalia obscurati Bültmann & al., 2015 (see below). Wirth 1980, Rhizocarpetalia Klement 1949 CHARACTER TAXA OF THE ORDER: Caloplaca nom. conserv. propos., Acarosporetalia sinopi- vitellinula, Candelariella vitellina, Myriospora cae Creveld 1981, Umbilicarietalia Oberdorfer scabrida, Rhizoplaca melanophthalma, Rinodina ex Klika et Hadac 1944, Parmelietalia sax- confragosa. atilis Wirth 1972 and Rinodino confragosae- DIFFERENTIAL SPECIES AGAINST THE OTHER ORDERS Rusavskietalia elegantis). The latter is a new ON STRONGLY ACIDIC ROCKS ARE NEUTROPHYTES OR order comprising sub-nitrophilous lichen com- CALCIPHYTES: Candelariella aurella, Lecania munities on subacidic siliceous rocks and on erysibe, Lecanora dispersa, Phaeophyscia orbicu- acidic rocks irrigated by calcium-rich water. laris, Placynthium nigrum. Characteristic is the combination of acido- phytes and nitrophilous neutrophytes, indiffer- ent species and weak basiphytes. Besides the Rhizocarpo geographici-Rusavskion elegantis type alliance, the Rhizocarpo-Rusavskion (see Creveld all. nov. hoc loco below), the order includes also the Lecanorion (Rinodino confragosae-Rusavskietalia elegan- rubinae Frey 1933 on nutrient enriched tops of tis, Rhizocarpetea geographici)

LAZAROA 36: 107-129. 2015 110 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes

VALIDATED NAME: Rhizocarpo-Xanthorion culosae Asta et Roux ex Roux in Bültmann et al. Creveld 1981 (CREVELD, 1981: 88) [Art. 3g]. 2015 (see below). SYNONYMS: Neuropogion melaxathi Mattick CHARACTER TAXA OF THE CLASS: Acarospora 1951 [Art. 2b], Physcion caesiae Mattick 1951 freyi, Aspicilia candida, Lecidea tessellata var. [Art. 2b], Caloplacion elegantis Hadac 1962 caesia, L. umbonata. [Arts. 2b, 3b], Xanthorion elegantis Dubiel et Olech 1990 [Art. 3b]. TYPE: Holotypus hoc loco: Buellio nivalis- Aspicilietalia verruculosae Asta et Roux ex Xanthorietum elegantis Creveld 1981 (CREVELD, Roux ordo nov. hoc loco 1981: 89-92, Table II, relevés 1-16). (Aspicilietea candidae) CHARACTER TAXON OF THE ALLIANCE: Physcia caesia. VALIDATED NAME: Aspicilietalia verruculosae Asta et Roux 1977 (ASTA & ROUX, 1977: 69-70) [Art. 3b]. NEW LICHEN SYNTAXA ON CALCIFEROUS SYNONYM: Aspicilietalia verruculosae Asta et SCHISTS AND DECALCIFIED ROCKS IN Roux in Roux 1981 (ROUX, 1981: 134) [Art. 3b]. THE ALPS TYPE: Holotypus hoc loco: Aspicilion mashigi- nensis Asta et Roux ex Roux in Bültmann et al. ASTA & ROUX (1977) and ROUX (1981) 2015 (see below). described (for the high-rank syntaxa prelimi- CHARACTER TAXA OF THE ORDER: Aspicilia poly- nary, hence invalidly) lichen vegetation of cal- chroma subsp. polychroma chemotype verrucu- careous schists and decalcified rocks at high losa, Caloplaca anchon-phoeniceon, C. insularis. altitudes of the Alps. The Aspicilietea candidae comprises the achionophytic vegetation types with the sub-thermophilous and the sub- Aspicilion mashiginensis Asta et Roux ex xerophilous order Aspicilietalia verruculosae Roux all. nov. hoc loco and the less thermophilous Lecideetalia con- (Aspicilietalia verruculosae, Aspicilietea candidae) fluescentis. The Aspicilietalia verruculosae comprises two alliances that differ in the car- VALIDATED NAME: Aspicilion mastrucatae Asta et bonate content of the rock – the Aspicilion Roux 1977 (ASTA & ROUX, 1977: 69-70) [Art. 3b]. mashiginensis (showing low carbonate content) SYNONYM: Aspicilion mastrucatae Asta et Roux and the Teloschistion contortuplicati (slightly in Roux 1981 (ROUX, 1981: 134) [Art. 3b] A. higher or higher carbonate content). mastrucata auct. non (Wahlenb.) Th. Fr. is a The new class Porpidietea zeoroidis, com- synonym of Aspicilia mashiginensis. prising a new order and alliance, was coined to TYPE: Holotypus hoc loco: Lecanoretum albu- accommodate chionophytic lichen vegetation. lae Asta et Roux 1977 (ASTA & ROUX, 1977: 55, In the sequel we validate these syntaxa: Table 20 on pp. 58-59). CHARACTER TAXA OF THE ALLIANCE: Acarospora hospitans, A. impressula, Aspicilia Aspicilietea candidae Asta et Roux ex Roux mashiginensis, A. permutata. class. nov. hoc loco

VALIDATED NAME: Aspicilietea candidae Asta et Teloschistion contortuplicati Roux all. nov. Roux 1977 (ASTA & ROUX, 1977: 69–70) [Art. hoc loco 3b]. (Aspicilietalia verruculosae, Aspicilietea candidae) SYNONYM: Aspicilietea candidae Asta et Roux in Roux 1981 (ROUX, 1981: 133) [Art. 3b]. TYPE: Holotypus hoc loco: Teloschistetum contor- TYPE: Holotypus hoc loco: Aspicilietalia verru- tuplicati Asta et Roux 1977 (ASTA & ROUX, 1977:

111 LAZAROA 36: 107-129. 2015 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes

39; Table 11 on pp. 42-43). Teloschistes contortu- TYPE: Holotypus hoc loco: Porpidion zeoroidis plicatus (Ach.) Clauzade & Rondon, is often list- Asta et Roux ex Roux all. nov. (see below). ed as Seirophora contortuplicata (Ach.) Frödén, CHARACTER TAXA OF THE ORDER: Adelolecia in Frödén & Lassen, however the separation from kolaensis, Polyblastia cupularis var. cupularis, Teloschistes has not been confirmed yet. Porpidia zeoroides, Rhizocarpon petraeum. CHARACTER TAXA OF THE ALLIANCE: Acarospora freyi, A. scabra, Carbonea atronivea, Diplotomma Porpidion zeoroidis Asta et Roux ex Roux all. dispersum, Lecanora diaboli, Polysporina pusilla, nov. hoc loco Teloschistes contortuplicatus. (Porpidietalia zeoroidis, Porpidietea zeoroidis)

VALIDATED NAME: Huilion macrocarpae- Lecideetalia confluescentis Roux ordo nov. trullisatae Asta et Roux 1977 (ASTA & ROUX, hoc loco 1977: 69-70) [Art. 3b]. (Aspicilietea candidae) SYNONYM: Huilion macrocarpae-trullisatae Asta et Roux in Roux 1981 (ROUX, 1981: 135, TYPE: Holotypus hoc loco: Lecideion conflues- 137-138) [Art. 3b] Huilia macrocarpa (DC.) centis Roux in Bültmann et al. 2015 (see below). Hertel var. trullisata (Arnold) Hertel is a syn- CHARACTER TAXA OF THE ORDER: Bellemerea onym of Porpidia zeoroides. subcandida, Farnoldia micropsis, Thelidium TYPE: Holotypus hoc loco: Stenhammarelletum ungeri, Verrucaria fischeri (syn. V. tristis turgidae Hertel ex Asta, Clauzade et Roux 1973 (Massal.) Krempel.) (ASTA & al., 1973b: 547-563, Table 1, rel. 1-8; subassociation of Stenhamarella turgida = Stenhammarelletum turgidae sensu Asta et Lecideion confluescentis Roux all. nov. hoc Roux 1977) The group of relevés named ‘subas- loco sociation of Lecidea confluescens’ in 1973 was (Lecideetalia confluescentis, Aspicilietea can- reclassified as the Lecideetum confluescentis didae) and classified in the Aspicilietea candidae by ASTA & ROUX (1977). TYPE: Holotypus hoc loco: Lecideetum conflues- CHARACTER TAXA OF THE ALLIANCE: Adelolecia centis Asta et Roux 1977 (ASTA & ROUX, 1977: kolaensis, Polyblastia cupularis var. c., 33; Table 7 on pp. 34-35). Porpidia zeoroides, Rhizocarpon petraeum CHARACTER TAXA OF THE ALLIANCE: Lecanora dispersoareolata, Lecidea confluescens, L. lep- rosolimbata, L. speirodes. VALIDATION OF SYNTAXA OF JELLY LICHENS ON LIMESTONE AND BASE-RICH ROCKS Porpidietea zeoroidis Roux class. nov. hoc loco The Collematetea cristati Wirth 1980 is a class TYPE: Holotypus hoc loco: Porpidietalia of cyanobacterial lichen communities on calcare- zeoroidis Asta et Roux ex Roux in Bültmann et ous or base-rich rocks, dominated by jelly lichens. al. 2015 (see below). This growth form can accumulate the occasional CHARACTER TAXA OF THE CLASS: Adelolecia supply of seepage water and become swollen and kolaensis, Polyblastia cupularis var. cupularis, jelly-like when wet. The Collematetalia cristati Porpidia zeoroides, Rhizocarpon petraeum. Wirth 1980 comprises vegetation on limestone with the thermophilous Peccanion coralloidis in Porpidietalia zeoroidis Asta et Roux ex Roux well illuminated and the Collemation tuniformis ordo nov. hoc loco Klement 1955 corr. Wirth 1980 in shaded habitats. (Porpidietea zeoroidis) The other order, the Peltuletalia euplocae (com-

LAZAROA 36: 107-129. 2015 112 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes prising the Peltulion euplocae Llimona et Egea VALIDATION OF THE ROCCELLETEA 1984) accomodates the jelly lichen syntaxa on PHYCOPSIS, WITH NEW ORDERS AND base-rich siliceous rocks. AN ALLIANCE

The ombrophobic and aerohygrophilous rock Peccanion coralloidis Moreno et Egea ex Egea lichen communities of this class are supported all. nov. hoc loco by calcareous to subacidic substrates, found in (Collematetalia cristati, Collematetea cristati) rather well-illuminated rock overhangs. The class is optimally developed in Southern VALIDATED NAME: Peccanion coralloides Europe, Macaronesia and North Africa. It Moreno et Egea 1991 (MORENO & EGEA, 1991: includes conspicuous stands of large fruticose 74) [Art. 3b]. Roccella species, several of them formerly har- SYNONYMS: Dermatocarpion miniati Mattick vested for the production of dye. Syntaxa from 1951 p.p. [Art. 2b], Psorotichion schaereri carbonate rocks are classified in the Dirinetalia Wirth 1995 [Art. 3b]. massiliensis, with one alliance - the Roccellion TYPE: Holotypus hoc loco: Peccanio coral- phycopsis Egea et Llimona 1984, those from loidis-Thyreetum pulvinatae Nowak 1960 subacidic to subneutral rocks in the order (NOWAK, 1960: 367-370, Table 16). Roccelletalia fuciformis (with the moderately CHARACTER TAXA OF THE ALLIANCE: Anema aero-hygrophilous Paralecanographion grumu- decipiens, A. nummularium, Lichinella iodopul- losae and the highly aero-hygrophilous chra, Metamelanena caesiella, Peccania coral- Roccellion Klement 1965). loides, Placynthium subradiatum, Psorotichia diffracta, P. frustulosa, P. schaereri, Pterygiopsis affinis, Thyrea girardii. Roccelletea phycopsis Egea class. nov. hoc loco

Peltuletalia euplocae Morena et Egea ex Egea VALIDATED NAME: Roccelletea phycopsis Egea ordo nov. hoc loco 1989 (EGEA, 1989: 112) [Art. 3b]. (Collematetea cristati) SYNONYMS: Physcietea Mattick 1951 p.p. [Art. 2b], Physcietea Tomaselli 1956 [Art. 2b], VALIDATED NAME: Peltuletalia euplocae Roccelletea phycopsis Follmann 1993 [Art. 3b]. Llimona et Egea 1985 (LLIMONA & EGEA, 1985: TYPE: Holotypus hoc loco: Dirinetalia massiliensis 443) [Art. 3b]. Egea in Bültmann et al. 2015 (see below). SYNONYM: Peltuletalia euplocae Morena et Egea CHARACTER TAXA OF THE CLASS: Alyxoria subel- 1991 (MORENO & EGEA, 1991: 74) [Art. 3b]. evata, Dirina massiliensis morphotype soredia- TYPE: Holotypus hoc loco: Peltulion euplocae ta, Roccella phycopsis, Thelopsis isiaca. Llimona et Egea 1985 (LLIMONA & EGEA, 1985: 433-443). Dirinetalia massiliensis Egea ordo nov. hoc CHARACTER TAXA OF THE ORDER: Collema rys- loco soleum, Lichinella cribellifera, Peltula euploca, (Roccelletea phycopsis) P. omphaliza, P. placodizans. VALIDATED NAME: Dirinetalia massiliensis Egea The holotype Peltulion euplocae Llimona et 1989 (EGEA, 1989: 112) [Art. 3b]. Egea 1985 is a later homonym of the Peltulion SYNONYMS: Roccelletalia vicentinae Follmann euplocae Llimona et Egea 1984 (LLIMONA & 1993 p.p. [Art. 3b]. EGEA, 1984: 92). We have decided to follow TYPE: Holotypus hoc loco: Roccellion phycopsis MORENO & EGEA (1991) and chose the younger Egea et Llimona 1984 (EGEA & LLIMONA, 1984: homonym as the type for the Peltuletalia. 210).

113 LAZAROA 36: 107-129. 2015 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes

CHARACTER TAXA OF THE ORDER: Aloxyria neutral to moderately acidic bark pH. The mougeotii, A. variiformis, Dirina massiliensis Graphidetalia scriptae Hadac in Klika et Hadac morphotype massiliensis. 1944 (with two alliances) occur on very smooth bark, beech in particular, in moderately humid Roccelletalia fuciformis Egea ordo nov. hoc sites; the Bacidinetalia phacodis (with one loco alliance - the Agonimion octosporae) on more (Roccelletea phycopsis) porose bark of older trees in shaded and humid situations, and the Dendrographetalia decol- VALIDATED NAME: Roccelletalia fuciformis Egea orantis (with the Lecanactidion patellarioidis 1989 (EGEA, 1989: 113) [Art. 3b]. Crespo ex Giralt 1996) occur in rather dry habi- SYNONYMS: Physcietalia caesiae Mattick tats and has southern distribution. 1951 p.p. [Art. 2b], Roccelletalia vicentinae Follmann 1993 p.p. [Art. 3b]. TYPE: Holotypus hoc loco: Roccellion Bacidinetalia phacodis Bricaud et Roux ordo Klement 1965 (KLEMENT, 1965: 516). nov. hoc loco CHARACTER SPECIES OF THE ORDER: Roccella (Arthonio radiatae-Lecidelletea elaeochromae) fuciformis, R. tinctoria, R. tuberculata. VALIDATED NAME: Bacidietalia phacodis Bricaud et Roux 2004 (BRICAUD, 2004: 25, 211) Paralecanographion grumulosae Egea all. [Art. 3b]. nov. hoc loco TYPE: Holotypus hoc loco: Agonimion octospo- (Roccelletalia fuciformis, Roccelletea phycop- rae Bricaud et Roux in Bültmann et al. 2015 sis) (see below). CHARACTER TAXA OF THE ORDER: Bacidia frax- VALIDATED NAME: Lecanactidion monstrosae inea, Bacidina phacodes, non-lichenised fungi: Egea 1989 (EGEA, 1989: 113) [Art. 3b] Aleurodiscus disciformis, Dendrominia dryina, Lecanactis monstrosa Bagl. = Lecanactis gru- Didymosphaeria rubicola, Septobasidium mulosa (Dufour) Fr. var. monstrosa (Bagl.) quercinum. Grummann on acidic substrate had been sepa- rated from the more basiphytic var. grumulosa, but is now included in Paralecanographa gru- Agonimion octosporae Bricaud et Roux all. mulosa. nov. hoc loco TYPE: Holotypus hoc loco: Lecanactino mon- (Bacidietalia phacodis, Arthonio radiatae- strosae-Dirinetum insulanae Egea 1989 (EGEA, Lecidelletea elaeochromae) 1989: 114, 138-140). CHARACTER SPECIES OF THE ALLIANCE: VALIDATED NAME: Agonimion octosporae Lecanographa dialeuca, Opegrapha cesareen- Bricaud et Roux 2004 (BRICAUD, 2004: 43-44, sis, O. lutulenta, Paralecanographa grumulosa, 311) [Art. 3b]. circumscripta. SYNONYM: Bacidion rubellae Wirth 1995 [Art. 3b]. TYPE: Holotypus hoc loco: Hypocenomycetum stoechadianae Abbassi Maaf et Roux 1987 TWO NEW ORDERS AND AN ALLIANCE (ABBASSI MAAF & ROUX, 1987: 219-227, Table IN THE ARTHONIO RADIATAE- on pp. 224-227). LECIDELLETEA ELAEOCHROMAE CHARACTER TAXA OF THE ALLIANCE: Bacidia rubella, Biatoridium monasteriense, The Arthonio-Lecidelletea comprises epi- Coenogonium luteum, Gyalecta derivata, G. flo- phytic communities dominated mostly by crus- towii, G. liguriensis, Navicella pileata (non- tose lichens growing on smooth tree bark with a lichenised ), Thelopsis rubella.

LAZAROA 36: 107-129. 2015 114 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes

Dendrographetalia decolorantis Bricaud et Melosirion variantis Margalef 1951 (syn. Roux ordo nov. hoc loco Meridio circularis-Naviculion gregariae M. (Arthonio radiatae-Lecidelletea elaeochromae) Schlüter 1961) comprising benthic diatom com- munities in eutrophic or brackish lowland VALIDATED NAME: Schismatommetalia decol- waters, and the Cymbello-Synedrion capitatae orantis Bricaud et Roux 2004 (BRICAUD, 2004: M. Schlüter 1961 comprising benthic diatom 43, 91, 311) [Art. 3b] Schismatomma decol- communities of small eutrophic lowland ponds. orans (Turner et Borrer ex Sm.) Clauzade et In addition, at least nine alliances were Vìzda is a synonym of Dendrographa decol- described by MARGALEF (1949, 1950, 1951) for orans. the benthic algae communities in Spanish SYNONYMS: Lecanoretalia sienae Crespo 1981 mountains for a variety of water chemistries. It p.p. [Art. 1], Lecanoretalia sienae Crespo ex may well be that new orders or classes for those Crespo et Bueno 1984 p.p. [Art. 2b], communities should be defined in future. Those Lecanoretalia sienae Crespo ex Giralt 1996 p.p. alliances of MARGALEF (l.c.) were tentatively [Art. 5], Lecanoretalia sienae Crespo ex included in the Naviculetalia (Naviculetea). Boqueras 2000 p.p. [Art. 2b]. TYPE: Holotypus hoc loco: Lecanactidion patel- larioidis Crespo ex Giralt 1996 (GIRALT, 1996: Naviculetea gregariae Pankow ex Täuscher 423, 426-427). class. nov. hoc loco CHARACTER TAXA OF THE ORDER: Bactrospora patellarioides, Dendrographa decolorans. VALIDATED NAME: Naviculetea Pankow 1980 (Lecanactis patellarioides (Nyl.) Vain. is a syn- (PANKOW, 1980: 136) [Art. 3b, 3g]. onym of Bactrospora patellarioides). SYNONYM: Naviculetea Pankow ex Täuscher GIRALT (1996) adopted the concept of the name 1998 (TÄUSCHER, 1998: 631) [Art. 2b, 3g]. by using Lecanactidion patellarioidis Crespo TYPE: Holotypus hoc loco: Naviculetalia gre- 1981 ined. (not effectively published). By gariae Pankow ex Täuscher in Bültmann et al. adopting the name with only one valid associa- 2015 (see below). tion - the Dirinetum ceratonie Klement 1965 CHARACTER TAXA OF THE CLASS: Achnanthes bre- (=type), and by listing characteristic species, the vipes, Amphora coffaeiformis, A. ovalis, name Lecanactidion patellarioidis had been val- Anomoeoneis sphaerophora, Aphanothece stagni- idated by GIRALT (1996). na, Brebissonia boeckii, Caloneis amphisbaena, Calothrix scopulorum, Cocconeis placentula, Coleofasciculus chthonoplastes, Diatoma vul- VALIDATION OF THE NAVICULETEA garis, Ellerbeckia arenaria, Gomphonema parvu- WITH AN ORDER AND ALLIANCE FOR lum, Lyngbya aestuarii, Melosira varians, BENTHIC MICROALGAL COMMUNITIES Meridion circulare, Microcoleus autumnalis, IN FRESH AND BRACKISH WATERS Navicula salinarum, N. tripunctata, Oscillatoria limosa, Planothidium lanceolatum, Rhoicosphenia The Naviculetea was proposed by PANKOW abbreviata, Rivularia nitida, Spirulina subsalsa, (1980) as a provisional class and the order Tryblionella hungarica, Ulnaria ulna. Naviculetalia Pankow 1980 was invalidly pub- lished because it was not possible to establish which of several Navicula species in the original Naviculetalia gregariae Pankow ex Täuscher diagnosis of the syntaxon name served as the ordo nov. hoc loco eponymous taxon name. (Naviculetea gregariae) The order includes the Oscillatorion limosae (cyanobacterial and diatom communities in VALIDATED NAME: Naviculetalia Pankow 1980 eutrophic and brackish lowland waters), the (PANKOW, 1980: 136) [Art. 3g].

115 LAZAROA 36: 107-129. 2015 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes

SYNONYMS: Tribonemetalia Margalef 1960 [Art. CHARACTER TAXA OF THE ASSOCIATION: 2b], Amphipleuretalia Margalef 1960 [Art. 2b], Oscillatoria limosa, O. tenuis, Phormidium Euastretalia Margalef 1960 [Art. 2b], chalybeum, Pseudanabaena limnetica. Naviculetalia Pankow ex Täuscher 1997 (TÄUSCHER, 1997: 13) [Arts. 2b, 3g], Naviculetalia Pankow ex Täuscher 1998 A NEW ALLIANCE AND ASSOCIATION IN (TÄUSCHER, 1998: 631) [Art. 3g]. THE ASTERIONELLETEA FORMOSAE TYPE: Holotypus hoc loco: Meridio-Naviculion gregariae M. Schlüter 1961 (SCHLÜTER, 1961: The Asterionelletea formosae and the 585-586, 606 & Table 2 on pp. 573-579). Asterionelletalia formosae were described in a CHARACTER TAXA OF THE ORDER: the same taxa thesis multiplied by hectograph (TÄUSCHER, as for the Naviculetea gregariae (see above). 1981), which is not effectively published according to ICPN Art. 1 (WEBER & al., 2000). The syntaxon names have been used in several Oscillatorion limosae Täuscher all. nov. hoc publications (e.g. Täuscher, 1995), but pre- loco sumed as valid and never unintentionally vali- (Naviculetalia gregariae, Naviculetea gregariae) dated until Täuscher (1998) described the class and the order by listing only one valid alliance VALIDATED NAME: Oscillatorion Möller 1977 Asterionellion Möller et Pankow 1981. (MÖLLER, 1977: 62) [Art. 1]. MÖLLER & PANKOW (1981) validated the SYNONYMS: Oscillatorion Prát in Klika et Hadac alliance by listing only one valid association 1944 [phantom], Limoseto-Diatomeion (Fragilario crotonensis-Asterionelletum Fetzmann 1956 [Arts. 2c, 3b], Oscillatorion gracillimae Messikommer 1927), which they Möller et Pankow 1981 (MÖLLER & PANKOW, proposed to rename to Fragilario crotonensis- 1981: 321) [Art. 2b], Oscillatorion Täuscher Asterionelletum formosae because Asterionella 1998 (TÄUSCHER, 1998: 631) [Art. 2b]. gracillima (Hantzsch) Heiberg is a synonym of TYPE: Holotypus hoc loco: Oscillatorietum A. formosa. According to MÖLLER & PANKOW limosae Möller et Pankow ex Täuscher in (1981) the epithet of the alliance should read Bültmann et al. 2015 (see below). ‘formosae’. CHARACTER TAXA OF THE ALLIANCE: Microcoleus The class and the order include the three autumnalis, Oscillatoria limosa, O. tenuis, alliances, such as the Asterionellion formosae Phormidium chalybeum, Pseudanabaena lim- Möller et Pankow 1981 (communities dominat- netica, Spirulina subsalsa. ed by diatoms), the Aphanizomeno floris-aquae- Microcystion aeruginosae Täuscher 1995 (dom- inated by cyanobacteria), and the Pediastro Oscillatorietum limosae Möller & Pankow ex duplicis-Scenedesmion quadricaudae (dominat- Täuscher ass. nov. hoc loco ed by green microalgae). (Oscillatorion limosae, Naviculetalia gregariae, Naviculetea gregariae) Pediastro duplicis-Scenedesmion quadricau- VALIDATED NAME: Oscillatorietum limosae dae Täuscher all. nov. hoc loco Möller et Pankow 1981 (MÖLLER & PANKOW, (Asterionelletalia formosae, Asterionelletea for- 1981: 318-319) [Art. 5]. mosae) SYNONYMS: Schlamm-Oscillatorietum Kurz 1922 [Art. 2b], Oscillatorietum limosae Prát ex VALIDATED NAME: Pediastro-Scenedesmion Klika et Hadac 1944 [Art. 2b]. Täuscher 1981 (TÄUSCHER, 1981: 93) [Art. 1]. TYPE: Holotypus hoc loco: Möller & Pankow SYNONYMS: Pediastro-Scenedesmion Täuscher (1981: 318. Table 8, relevé 7). 1995 (TÄUSCHER, 1995: 5) [Arts. 2b, 3g],

LAZAROA 36: 107-129. 2015 116 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes

Pediastro-Scenedesmion Täuscher 1998 Gloeocapsetea sanguineae Bültmann & (TÄUSCHER, 1998: 631) [Arts. 2b, 3g]. Golubiç class. nov. hoc loco TYPE: Holotypus hoc loco: Pediastro duplicis- Scenedesmetum quadricaudae Täuscher in TYPE: Holotypus hoc loco: Gloeocapsetalia san- Bültmann et al. 2015 (see below). guineae Bültmann et Golubiç in Bültmann et al. CHARACTER TAXA OF THE ALLIANCE: Actinastrum 2015 (see below). hantzschii, Closterium limneticum, Micractinium CHARACTER TAXA OF THE CLASS: Gloeocapsa pusillum, Mucidosphaerium pulchellum, biformis, G. compacta, G. kuetzingiana, G. san- Pandorina morum, Pediastrum duplex, guinea, Schizothrix heufleri, Scytonema Scenedesmus quadricauda. myochrous.

Pediastro duplicis-Scenedesmetum quadricau- dae Täuscher ass. nov. hoc loco Gloeocapsetalia sanguineae Bültmann & (Pediastro duplicis-Scenedesmion quadricau- Golubiç ordo nov. hoc loco dae, Asterionelletalia formosae, Asterionelletea (Gloeocapsetea sanguineae) formosae) TYPE: Holotypus hoc loco: Gloeocapsion san- VALIDATED NAME: Pediastro-Scenedesmetum guineae Golubiç 1967 (GOLUBIÇ, 1967: 165- Täuscher 1995 (TÄUSCHER, 1995: 8) [Arts. 2b, 3g]. 166. Tables 13 & 14). TYPE: Holotypus hoc loco: TÄUSCHER (1995: 8. CHARACTER TAXA OF THE ORDER: the same taxa Table 1 consists of one relevé, which is the holo- as for the Gloeocapsetea sanguineae (see type). above). CHARACTER TAXA OF THE ASSOCIATION: Pediastrum duplex, Scenedesmus quadricauda. A NEW CLASS, ORDER AND ALLIANCE FOR THE GREEN ALGAE FILMS ON A NEW CLASS AND ORDER FOR THE BARK AND ACIDIC ROCKS VEGETATION OF AEROPHYTIC MICROAL- GAL FILMS ON CALCAREOUS ROCKS The species composition of biofilms of green algae was studied frequently in context of GOLUBIÇ (1967) described nine associations biodeterioration, however rarely using phytoso- of vegetation of rock algae in the Dinarides in ciological methods. It is therefore, that the the alliance Gloeocapsion sanguineae Golubiç alliance Schizogonion cruenti Ochsner 1928, 1967. Weak floristic relations link those com- containing described three associations munities to the syntaxa of benthic aquatic algae, (Pleurococcetum vulgaris, Schizogonietum but the strong differences made it impossible to crenulatae and Trentepohlietum abietinae; all classify the aerophytic rock algae communities nomina nuda) remained a nomen nudum since it to the Naviculetea, and thus a new class and new had been suggested by OCHSNER (1928). order have to be described. One species common in those biofilms is Gloeocapsa sanguinea is a widely distributed Desmococcus olivaceus, which is often listed as cyanobacterium, which is typical for moist sur- one of its synonyms such as Desmococcus faces of calcareous substrates, e.g. in the black viridis (C. Agardh) P.C. Silva, Desmococcus patina or tintenstrichen on rocks, cement walls vulgaris F. Brand, Pleurococcus viridis (C. and monuments (e.g. SAMAD & ADHIKARY, Agardh) Rabenh. or Pleurococcus vulgaris 2008; GOLUBIç & al., 2015) or in the ‘lampen- Nägeli (LAUNDON, 1985). Apatococcus lobatus flora’ in Karst caves (MULEC & KOSI, 2009). It (Chodat) J.B. Petersen is another common is also known as a photobiont in lichens (BÜDEL species of the green algal films (e.g. BRAND, & HENSSEN, 1988). 1925; BARKMAN, 1958), and GÄRTNER &

117 LAZAROA 36: 107-129. 2015 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes

INGOLIÇ (1989); GÄRTNER (1994) proposed to TYPE: Holotypus hoc loco: Pleurococcetum vul- replace the name Pleurococcetum vulgaris with garis Schorler ex Barkman 1958 (BARKMAN, the Apatococcetum lobati but never supported 1958: 341). their proposal with a publication of relevés SYNONYMS: Schizogonion cruenti Ochsner 1928 showing that Apatococcus lobatus is more fre- (OCHSNER, 1928: 47) [Art. 2b], Schizogonion quent than Desmococcus olivaceus. Other cruenti Ochsner in Klika et Hadac 1944 [Art. authors found Desmococcus in equal or higher 2b]. abundance than Apatococcus (RINDI & GUIRY, CHARACTER TAXA OF THE ALLIANCE: the same 2003; RINDI, 2007). It is possible, that taxa as for the Desmococcetea olivacei (see Desmococcus prefers moister sites and it is above). more common in the Atlantic Europe, while Apatococcus prefers continental areas and drier habitats (GUSTAVS, 2010). The Desmococcion MESOTAENIETEA BERGGRENII - A NEW olivacei comprises the two validly published CLASS AND SUBORDINATE SYNTAXA associations, such as the Pleurococcetum vul- FOR SNOW ALGAE garis Schorler ex Barkman 1958 dominated by water-repellent coccoid green algae and the KLIKA & HADAC (1944) proposed the Prasiloletum crispae Knebel ex Barkman 1958, Sphaerellion nivalis Hadaç in Klika & Hadac dominated by filamentous algae. (Schizogonium 1944 for the snow algae syntaxa, yet this name crenulatum (Kütz.) Gay is a synonym of remained a nomen nudum. The vegetation and Prasiola crispa (Lightfoot) Kützing.) habitat of the snow algae is so unique, that the vegetation could not be placed in any existing class. Communities of snow and ice algae have Desmococcetea olivacei Bültmann class. nov. been studied in Antarctica (e.g. FOGG, 1967) to hoc loco Alaska (e.g. TAKEUCHI, 2001). Besides strong floristic difference depending on the growth on TYPE: Holotypus hoc loco: Desmococcetalia oliva- permanent snow or ice as the substrate (e.g. cei Bültmann in Bültmann et al. 2015 (see below). KOL, 1942), further variations have been CHARACTER TAXA OF THE CLASS: Apatococcus loba- tus, Desmococcus olivaceus, Klebsormidium flac- observed with altitude and ecoregion cidum, Porphyridium purpureum, Prasiola crispa, (TAKEUCHI, 2013). The evaluation of samples, Rosenvingiella radicans, Schizogonium murale, which are equivalent to relevés (KOL, 1942; Trentepohlia abietina, Trentepohlia umbrina. FOGG, 1967; TAKEUCHI, 2001), suggests at least two associations, such as the Mesotaenietum Desmococcetalia olivacei Bültmann ordo nov. berggrenii for ice algae and the hoc loco Chloromonadetum nivalis for snow algae, both (Desmococcetea olivacei) are described here based on samples from Alaska. TYPE: Holotypus hoc loco: Desmococcion oliva- Chlamydomonas nivalis is the collective cei Bültmann in Bültmann et al. 2015 (see below). name for algae responsible for the red snow CHARACTER TAXA OF THE ORDER: the same taxa and this taxonomic concept was used to as for the Desmococcetea olivacei (see above). describe (invalidly) the Chlamydomonadetum nivalis (PANKOW & al., 1991). However since the of this ‘species’ is not clear (e.g. Desmococcion olivacei Bültmann all. nov. hoc REMIAS & al., 2013) another characteristic loco species, the ice alga Mesotaenium berggrenii, (Desmococcetalia olivacei, Desmococcetea oli- was chosen to give the name to class, order vacei) and alliance.

LAZAROA 36: 107-129. 2015 118 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes

Mesotaenietea berggrenii Bültmann et TYPE: Holotypus hoc loco: TAKEUCHI (2001: Takeuchi class. nov. hoc loco 3452. Table I, sample 5, Alaska): Mesotaenium berggrenii 20371 cells, Ancylonema nordenski- TYPE: Holotypus hoc loco: Mesotaenietalia oldii 6439 cells/ml, Chlamydomonas nivalis 487 berggrenii Bültmann et Takeuchi in Bültmann et cells/ml, Cylindrocystis brebissonii 243 al. 2015 (see below). cells/ml, Raphidonema sp. 54 cells/ml, CHARACTER TAXA OF THE CLASS: Ancylonema Oscillatoriaceae spec. 1 34330 cells/ml, Osc. nordenskioldii, Chlamydomonas nivalis, spec. 2 16177 cells/ml. Chloromonas nivalis, Cryocystis brevispina, CHARACTER TAXA OF THE ASSOCIATION: Desmotetra antarctica, Mesotaenium Ancylonema nordenskioldii, Mesotaenium berggrenii, Raphidonema nivale. berggrenii.

Mesotaenietalia berggrenii Bültmann et Chloromonadetum nivalis Bültmann et Takeuchi ordo nov. hoc loco Takeuchi ass. nov. hoc loco (Mesotaenietea berggrenii) (Mesotaenion berggrenii, Mesotaenietalia berggrenii, Mesotaenietea berggrenii)

TYPE: Holotypus hoc loco: Mesotaenion berggrenii Bültmann et Takeuchi in Bültmann et TYPE: Holotypus hoc loco: KOL (1942: Sample al. 2015 (see below). 35, Alaska, Thompson Pass, red snow, abundance and species data on pages 17, 19, 20, 21, 23, 27): CHARACTER TAXA OF THE ORDER: Ancylonema Smithsonimonas abbotii very abundant, nordenskioldii, Chlamydomonas nivalis, Chlamydomonas nivalis abundant, Chloromonas Chloromonas nivalis, Mesotaenium berggrenii, nivalis abundant, Gloeocapsa sanguinea (incl. G. Raphidonema nivale. ralfsii (Harvey) Lemmermann) not rare, Mycacanthococcus cellaris f. antarcticus rare, Protoderma cohaerens rare, Raphidonema brevi- Mesotaenion berggrenii Bültmann et rostre rare, Raphidonema nivale rare. Takeuchi all. nov. hoc loco SYNONYMS: Chlamydomonadetum nivalis Gams (Mesotaenietalia berggrenii, Mesotaenietea 1927 [Art. 2b], Chlamydomonadetum nivalis berggrenii) Gams 1936 [Art. 2b], Chlamydomonadetum nivalis Pankow et al. 1991 [Art. 2b]. YPE T : Holotypus hoc loco: Mesotaenietum CHARACTER TAXA OF THE ASSOCIATION: berggrenii Bültmann et Takeuchi in Bültmann et Chlamydomonas nivalis, Chloromonas nivalis. al. 2015 (see below). SYNONYMS: Sphaerellion nivalis Hadac in Klika et Hadac 1944 [2b], Sphaerellion nivalis Hadac VALIDATIONS AND TYPIFICATIONS IN ex Klika 1948 [Art. 2b], Sphaerellion nivalis THE ENTOPHYSALIDETEA DEUSTAE Hadac 1962 [Art. 2b]. CHARACTER TAXA OF THE ALLIANCE: the same The Entophysalidetea comprises the marine taxa as for the Mesotaenietalia berggrenii (see photophytic algal vegetation on hard substrates above). in lower supralittoral and upper parts of eu-lit- toral level. Unfortunately in the description expressis verbis, the association Lithophylletum Mesotaenietum berggrenii Bültmann et lichenoidi, was chosen as type of the class Takeuchi ass. nov. hoc loco (GIACCONE & al., 1993) hence rendering the (Mesotaenion berggrenii, Mesotaenietalia class invalid according to the ICPN. Here we berggrenii, Mesotaenietea berggrenii) provide an ICPN-conform typification.

119 LAZAROA 36: 107-129. 2015 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes

This class includes the Pleurocapsetalia SYNONYMS: Fucetalia Hadac in Klika 1948 [Art. gloeocapsoidis Ercegovic 1932 in the supralit- 2b], Fucetalia Golub et al. 2003 [Arts. 2b, 5], toral zone of the Atlantic and the Mediterranean Ascophyllo nodosi-Fucetalia serrati Julve et (with two alliances), the Bangietalia atropur- Manneville 2006 [Art. 3i]. pureae Giaccone in Giaccone et al. 1993 in the TYPE: Holotypus hoc loco: Ascophyllion nodosi upper eu-littoral and the Neogoniolitho Julve in Bültmann et al. 2015 (see below). notarisii-Nemodermetalia tingitani Molinier CHARACTER TAXA OF THE ORDER: Cladophora 1960 in the lower eu-littoral of the rupestris and the character taxa of the alliances, Mediterranean (one alliance each), the Fucetalia e.g. Ascophyllum nodosum, Fucus serrratus, F. vesiculosi with two new alliances for the upper vesiculosus, Pelvetia canaliculata. and lower eu-littoral of the Atlantic and, finally the Dalmatelletalia polyformis Ercegovic 1932 and the Hyelletalia caespitosae Ercegovic 1932 Ascophyllion nodosi Julve all. nov. hoc loco described from limestone coasts of the Eastern (Fucetalia vesiculosi, Entophysalidetea deustae) Mediterranean (with two alliances each). Several of the characteristic species of the two VALIDATED NAME: Ascophyllion nodosi Julve et orders by ERCEGOVIC (1932) are also known Manneville 2006 (JULVE & MANNEVILLE, 2006: from the Atlantic limestone coasts and it is pos- 222-223. Table 1) [Art. 5]. sible that these poorly-known syntaxa are not SYNONYMS: Dictyosiphonion foeniculacei Du exclusively Mediterranean. Rietz 1941 [Art. 2b], Dictyosiphonion Du Rietz ex Klika 1948 [Art. 2b], Pelvetion canaliculatae Hadac in Klika 1948 [Art. 2b], Fucion Golub et Entophysalidetea deustae Giaccone class. nov. al. 2003 [Arts. 2b, 5]. hoc loco TYPE: Holotypus hoc loco: Ascophylletum nodosi den Hartog 1959 (DEN HARTOG, 1959: VALIDATED NAME: Entophysalidetea deustae 199-204). Giaccone in Giaccone et al. 1993 (GIACCONE & CHARACTER TAXA OF THE ALLIANCE: al., 1993: 262, 270) [Art. 5]. Ascophyllum nodosum, Bostrychia scorpioides, SYNONYMS: Chthamaletea Giaccone 1965 [Art. Catenella caespitosa, Fucus ceranoides, F. spi- 2c], Melarphetea neritoidis Giaccone 1965 p.p. ralis, F. vesiculosus, Pelvetia canaliculata, [Art. 2c], Dictyoto dichotomae-Laurencietea Vertebrata lanosa. pinnatifidae Julve 1992 p.p. [Art. 5], Peyssonnelio dubyi-Lithophylletea incrustantis Julve 1992 p.p. [Art. 2b], Fucetea Golub et al. VALIDATIONS IN THE CYSTOSEIRETEA 2003 [Arts. 2b, 5], Dictyoto dichotomae- Osmundeetea pinnatifidae Julve ex Julve et The Cystoseiretea Giaccone 1965 comprises Manneville 2006 p.p. [Art. 3i]. the Mediterranean Cystoseiretalia Molinier TYPE: Holotypus hoc loco: Ralfsietalia verru- 1960, the Atlantic Laminarietalia hyperboreae cosae Giaccone 1993 (GIACCONE & al., 1993: Julve 1992, and the thio-nitrophytic Ulvetalia 269, 281). lactucae Molinier 1960 occurring in in both CHARACTER TAXON OF THE CLASS: Brachytrichia Atlantic Ocean and the Mediterranean Sea. quoyi. The Laminarietalia include the Laminarion saccharinae Julve 1992 in protected coastal habi- Fucetalia vesiculosi Julve ordo nov. hoc loco tats and the Laminarion hyperboreae in more (Entophysalidetea deustae) dynamic habitats. JULVE & MANNEVILLE (2006) used the name Cystoseirion tamariscifoliae, but as VALIDATED NAME: Fucetalia vesiculosi Julve no such species occurs in the relevés of the origi- 1992 (JULVE, 1992: 566-567) [Art. 2b]. nal diagnosis, the name Laminarion hyperboreae,

LAZAROA 36: 107-129. 2015 120 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes was chosen instead. JULVE & MANNEVILLE (2006) 1, Dilsea carnosa 1, Furcellaria lumbricalis 1, described the Alario esculenti-Himanthalietum Halurus flosculosus 1, Membranoptera alata 1, elongatae and the Saccorhizo polyschidi- Osmundea pinnatifida 1, Calliblepharis jubata Laminarietum hyperborea by presenting only a +, Callithamnion tetragonum +, Ceramium vir- synoptic table. The alliance and both associations gatum +, Chondracanthus acicularis +, are validated here: Claviclonium ovatum +, Codium tomentosum +, Desmarestia aculeata +, Gelidium pulchellum +, Halurus equisetifolius +, Heterosiphonia Laminarion hyperboreae Julve all. nov. hoc plumosa +, Plocamium cartilagineum +. loco CHARACTER TAXA OF THE ASSOCIATION: (Laminarietalia hyperboreae, Cystoseiretea) Calliblepharis jubata, Dilsea carnosa, Laminaria hyperborea, Saccorhiza polyschides. VALIDATED NAME: Cystoseirion tamariscifoliae Julve 1992 (JULVE, 1992: 568) [Art. 2b] SYNONYM: Cystoseirion baccatae Julve 1992 Alario esculenti-Himanthalietum elongatae [Art. 2b]. Julve ass. nov. hoc loco. TYPE: Holotypus hoc loco: Saccorhizo polyschi- (Laminarion hyperboreae, Laminarietalia di-Laminarietum hyperboreae Julve in hyperboreae, Cystoseiretea) Bültmann et al. 2015 (see below). CHARACTER TAXA OF THE ALLIANCE: Alaria escu- VALIDATED NAME: Alario esculenti- lenta, Calliblepharis jubata, Chylocladia verti- Himanthalietum elongatae Julve & Manneville cillata, Dilsea carnosa, Halopteris scoparia, 2006 (JULVE & MANNEVILLE, 2006: 225. Table Himanthalia elongata, Laminaria digitata, L. 1) [Art. 2b]. hyperborea, Lomentaria clavellosa, Saccorhiza TYPE: Holotypus hoc loco: JULVE & MANNEVILLE polyschides. (2006) include a column of a synoptic table, but the full table is given online only. Here the relevé 78 is selected from from that source as the holo- Saccorhizo polyschidi-Laminarietum hyper- type: France, Bretagne, eu- to infralittoral, grow- boreae Julve ass. nov. hoc loco ing on rock exposed to strong wave action (cover ( Laminarion hyperboreae, Laminarietalia scale: + to 5): Himanthalia elongata 4, Corallina hyperboreae, Cystoseiretea) elongata 3, Mastocarpus stellatus 3, Osmundea pinnatifida 2, Alaria esculenta 1, Bifurcaria bifur- VALIDATED NAME: Saccorhizo polyschidi- cata 1, Boergeseniella thuyoides 1, Chondrus Laminarietum hyperboreae Julve et Manneville crispus 1, Chylocladia verticillata 1, Laminaria 2006 (JULVE & MANNEVILLE, 2006: 225. Table digitata 1, Cladophora rupestris +, Codium 1) [Art. 2b]. tomentosum +, Lomentaria articulata +, TYPE: Holotypus hoc loco: JULVE & MANNEVILLE Lomentaria clavellosa +, Polysiphonia stricta +. (2006) included a column of a synoptic table, CHARACTER TAXA OF THE ASSOCIATION: Alaria but the full table was presented online. Here the esculenta, Chylocladia verticillata, Halopteris relevé 235 had been extracted from that source scoparia, Himanthalia elongata, Lomentaria is presented as the holotype: clavellosa. France, Bretagne, upper infralittoral, growing on rock face, exposed to wave action (cover scale: + to 5): Laminaria hyperborea 4, A NEW ORDER AND ALLIANCE IN THE Chondrus crispus 2, Ellisolandia elongata 2, LITHOPHYLLETEA SOLUTI Laminaria digitata 2, Palmaria palmata 2, Saccorhiza polyschides 2, Callophyllis laciniata The class Lithophylletea soluti Giaccone 1, Corallina officinalis 1, Cryptopleura ramosa 1965 includes two Mediterranean orders

121 LAZAROA 36: 107-129. 2015 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes

(Rhodymenietalia ardissonei Augier et VALIDATED NAME: Heterosiphonio plumosi- Boudouresque 1975 and Lithophylletalia soluti Delesserietum sanguineae Julve et Manneville Giaccone 1965) and the Atlantic Delesserietalia 2006 (JULVE & MANNEVILLE, 2006: 226. Table sanguinei. The validation of the latter is neces- 1) [Art. 2b]. sary because in JULVE (1992), the class, the TYPE: Holotypus hoc loco: JULVE & order and the associations were listed without MANNEVILLE (2006) included a column of a syn- the name of an alliance and JULVE & optic table; full relevé table was presented MANNEVILLE (2006) described the alliance with online and here we select the relevé 133 from one invalid association only. this source and present as the holotype: France, Bretagne, eu-littoral, on rock in a shaded pool, exposed to wave action (cover Delesserietalia sanguinei Julve ordo nov. hoc scale: + to 5): Phyllophora crispa 4, Plumaria loco plumosa 3, Corallina officinalis 2, Ceramium (Lithophylletea soluti) virgatum 1, Laminaria digitata 1, Delesseria sanguinea +. ALIDATED NAME V : Delessertietalia sanguinei CHARACTER TAXA OF THE ASSOCIATION: Julve 1992 (JULVE, 1992: 569-570) [Art. 2b]. Apoglossum ruscifolium, Delesseria sanguinea, TYPE: Holotypus hoc loco: Delesserion san- Halurus equisetifolius, Heterosiphonia plumosa, guineae Julve in Bültmann et al. 2015 (see below). Phyllophora crispa, Pleonosporium borreri, CHARACTER TAXA OF THE ORDER: Agarum clathra- Plumaria plumose. tum, Alaria esculenta, Bonnemaisonia hamifera, Calliblepharis ciliata, Callithamnion tetricum, Chondrus crispus, Coccotylus truncatus, VALIDATION IN THE CAULERPETEA Delesseria sanguinea, Fimbrifolium dichotomum, RACEMOSAE Gigartina pistillata, Heterosiphonia plumosa, Membranoptera alata, Odonthalia dentata, The Caulerpetea racemosae includes one Phycodrys rubens, Phyllophora pseudocera- order and one alliance with five associations noïdes, Plumaria plumosa, Polysiphonia arctica, Ptilota gunneri, Stenogramma interruptum. dominated by species of Caulerpa (Caulerpetum racemosae Giaccone et Di Martino 1995, Caulerpetum scalpeIIiformis Delesserion sanguinei Julve all. nov. hoc loco Mayhoub ex Giaccone & Di Martino 1995, (Delesserietalia sanguinei, Lithophylletea soluti) Caulerpetum mexicanae Giaccone et Di Martino 1995, Caulerpetum taxifoliae Di VALIDATED NAME: Delesserion sanguineae Julve Martino et Giaccone 1997 and Caulerpetum et Manneville 2006 (JULVE & MANNEVILLE, proliferae Giaccone et Di Martino 1997). In the 2006: 225-226. Table 1) [Art. 2b]. original description of the higher syntaxa, the SYNONYM: Polysiphonion arcticae Hadac in name was used without epithet and hence Klika 1948 [Art. 2b]. is not possible to decide, which of several TYPE: Holotypus hoc loco: Delesserietum san- Caulerpa were the name-giving species. guineae Julve in Bültmann et al. 2015 (see below). CHARACTER TAXA OF THE ALLIANCE: the same as for the order (see above). Caulerpetea racemosae Giaccone et Di Martino class. nov. hoc loco Delesserietum sanguineae Julve ass. nov. hoc loco VALIDATED NAME: Caulerpeteae Giaccone et Di (Delesserion sanguinei, Delesserietalia san- Martino 1997 (GIACCONE & DI MARTINO, 1997: guinei, Lithophylletea soluti) 6) [Art. 3g].

LAZAROA 36: 107-129. 2015 122 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes

TYPE: Holotypus hoc loco: Caulerpetalia race- VALIDATED NAME: Caulerpion Giaccone et Di mosae Giaccone et Di Martino in Bültmann et Martino 1997 (GIACCONE & DI MARTINO, 1997: al. 2015 (see below). 6) [Art. 3g]. CHARACTER TAXA OF THE CLASS: Caulerpa TYPE: Holotypus hoc loco: Caulerpetum race- prolifera, C. racemosa. mosae Giaccone et Di Martino 1995 (GIACCONE & DI MARTINO, 1995: 66, 68-70). Caulerpetalia racemosae Giaccone et Di CHARACTER TAXA OF THE ALLIANCE: the same as Martino ordo nov. hoc loco for the Caulerpetea racemosae (see above). (Caulerpetea racemosae)

VALIDATED NAME: Caulerpetalia Giaccone et Di ACKNOWLEDGEMENTS Martino 1997 (GIACCONE & DI MARTINO, 1997: 6) [Art. 3g]. First of all, the authors want to thank Laco TYPE: Holotypus hoc loco: Caulerpion race- Mucina, Perth, for his long-standing support. mosae Giaccone et Di Martino in Bültmann et Particular thanks are also due to Jean-Paul al. 2015 (see below). Theurillat, Chambésy and Jens Pallas, Münster, CHARACTER TAXA OF THE ORDER: the same as for for nomenclatural advice and Federico the Caulerpetea racemosae (see above). Fernández-González, Toledo, for the Spanish translations. Caulerpion racemosae Giaccone et Di Martino all. nov. hoc loco (Caulerpetalia racemosae, Caulerpetea race- mosae)

REFERENCES

Abbassi Maaf, L. & Roux, C. —1987— Les peuplements Brand, F. —1925— Analyse der aerophilen lichéniques corticoles de la chênaie verte: étude com- Grünalgenanflüge, insbesondere der proto-pleurococ- parée de la gardiole de Rians et de l’île de Port-Cros coiden Formen — Arch. Protist. 52: 265-354. (Var) — Bull. Soc. Linn. Provence 38: 189-245. Braun-Blanquet, J. —1964— Pflanzensoziologie. Asta, J. & Roux, C. —1977— Étude écologique et phy- Grundzüge der Vegetationskunde 3. Aufl. — Springer- tosociologique de la végétation lichénique des roches Verlag, Wien. plus ou moins décalcifiées en surface aux étages sub- Bricaud, O. —2004— Les peuplements lichéniques corti- alpin et alpin des Alpes françaises — Bull. Mus. Hist. coles sciaphiles et foliicoles méditerranéens de la Nat. Marseille 37: 23-81. France méridionale — Bull. Soc. Linn. Provence, Num. Asta, J., Clauzade, G. & Roux, C. —1973a— Étude de Spéc. 12: 1-324. quelques groupements lichéniques saxicoles-calcicoles Büdel, B. & Henssen, A. —1988— Trebouxia aggregata und du parc national de la Vanoise — Trav. Sci. Parc Nat. Gloeocapsa sanguinea, Phycobionten von Euopsis gra- Vanoise 3: 73-104. natina (Lichinaceae) — Plant Syst. Evol. 158: 235-241. Asta, J., Clauzade, G. & Roux, C. —1973b— Bültmann, H. —2005— Syntaxonomy of arctic terricolous Stenhammarelletum turgidae Hertel ass. nov. — Rev. lichen vegetation, including a case study from Southeast Fac. Ci. Lisboa 2. Ser. C 17(2): 543-567. Greenland — Phytocoenologia 36: 909-949. Barkman, J.J. —1958— Phytosociology and ecology of Bültmann, H. —2010— Diversity and similarity of lichen cryptogamic epiphytes — Van Gorcum, Assen. floras of countries along a south-north gradient from Barkman, J.J. —1973— Synusial approach to classifica- Italy to Greenland — Ann. Bot. S.N. 1(1): 1-9. tion — In: Whittaker, R.H. (Ed.). Ordination and classi- Creveld, M. —1981— Epilithic lichen communites in the fication of vegetation. Pp. 436–491. Dr W. Junk alpine zone of southern Norway — Bibl. Lichenol. 17: Publishers, The Hague. 1-288. Bomfleur, B., Krings, M. & Kerp, H. —2010— Thalloid Den Hartog, C. —1959— The epilithic algal communities organisms and the fossil record. New perspectives from the occuring along the coast of the Netherlands — Wentia 1: Transantarctic Mountains — Plant Sign. Behav. 5: 293-295. 1-241.

123 LAZAROA 36: 107-129. 2015 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes

Egea, J.M. —1989— Las comunidades liquénicas saxíco- Hilitzer, A. —1925— Étude sur la végétation epiphyte de las, ombrofobas, litorales, del suroeste de Europa y la Bohême — Spisy vidávané Prirodovdeckou fakultou Norte de Africa (Roccelletea phycopsis classis prov.) — Karlovy Univ. 41: 1-202. Stud. Geobot. 9: 73-152. Julve, P. —1992— Classification phytosociologique des Egea, J.M. & Llimona, X. —1984— Las comunidades végétations d´algues marines benthiques de France — liquénicas saxícolas, ombrofobas de la costa del SE. de Bull. Soc. Bot. Centre-Ouest N.S. 23: 565-576. España comprendidas entre el Penyal d’Ifach (Alicante) Julve, P. & Manneville O. —2006— Contribution à l’étude y Almería — Collect. Bot. 15: 205-219. synusiale des végétations littorales de macroalgues Ercegovic, A. —1932— Ecološke i sociološke studije î marines des côtes atlantiques françaises — Acta Bot. litofitskim cijanoficejama sa Jugoslavenske obale Gallica 153: 219-234. Jadrana (Ecological and sociological studies on litho- Khaybullina, L.S., Sukhanova, N.V., Kabirov, R.R. & phytic cyanophytes on the Yugoslavian coast of the Solomeshch, A.I. —2004— Sintaksonomija soobšcestv Adriatic Sea) — Rad Jugoslav. Akad. Znanosti i pocvennych vodoroslej juznogo Urala. Cast 1. Sojuz Umjetnosti 244: 129-220. (in Croatian). Amphoro-Phormidion all. nov. hoc loco. (Syntaxonomy Fogg, G. —1967— Observation on the snow algae of the of soil algae communities in South Ural. I. Alliance South Orkney Islands — Phil. Trans. Royal Soc. Amphoro-Phormidion all. nova hoc loco.) — Algologia London, Series B 252: 279-287. 14: 261-276. Frey, E. —1923— Die Berücksichtigung der Lichenen in Khaybullina, L.S., Sukhanova, N.V., Kabirov, R.R. & der soziologischen Pflanzengeographie, speziell der Solomeshch, A.I. —2005— Sintaksonomija soobšcestv Alpen — Verh. Nat. Ges. Basel 35: 303-320. pocvennych vodoroslej juznogo Urala. Cast 2. Sojuz Gärtner, G. & Ingoliç, E. —1989— Ein Beitrag zur Klebsormidio flaccidi-Myrmecion biatorellae all. nova. Kenntnis von Apatococcus lobatus (Chlorophyta, (Syntaxonomy of soil algae communities in South Ural. Chaetophorales, Leptosiroideae) — Plant Syst. Evol. II. Alliance Klebsormidio flaccidi-Myrmeñion biàtorel- 164: 133-143. lae all. nova.) — Algologia 15: 86-100. Gärtner, G. —1994— Zur Taxonomie aerophiler grüner Khaybullina, L.S., Sukhanova, N.V., Kabirov, R.R. & Algenanflüge an Baumrinden — Ber. Nat.-Med. Ver. Solomeshch, A.I. —2005a— Syntaxonomy of commu- Innsbruck 81: 51-59. nities of soil algae in the Southern Ural. 3. Class Giaccone, G. & Di Martino, V. —1995— La vegetazione a Bracteacocco-Hantzschietea cl. nov. — Int. J. Algae 7: Caulerpa racemosa (Forsskäl) C. Agardh nella Baia di S. 281-298. Panagia (Sicilia Sud-Orientale) — Boll. Accad. Gioenia Klement, O. —1955— Prodromus der mitteleuropäischen Sci. Nat. 28: 59-73. Flechtengesellschaften — Feddes Repert. Spec. Nov. Giaccone, G. & Di Martino, V. —1997— Inquadramento Regni Veg., Beit. Veg. 1: 1-194. fitosociologico ed ecologia della vegetazione a Caulerpe Klement, O. —1965— Zur Kenntnis der in Mediterraneo — In: Cossul, A. & Meloni, M.M. Flechtenvegetation der Kanarischen Inseln — Nova (Eds.). Atti del Convegno S.O.S. Caulerpa? Hedwigia 9: 503-582. Introduzione di nuove specie nel Mediterraneo e com- Klika, J. —1948— Rostlinná sociologie. (Plant sociology.) patibilità con quelle presenti. Cagliari, 25.11.1996. Pp. — Melantrich, Praha. (in Czech). 69–86. Poseidon Publ., Cagliari. Klika, J. & Hadac, E. —1944— Rostlinná spolecenstrední Giaccone, G., Alongi, G., Cossu, A., Di Geronimo, R. & Evropy. (Plant communities of Central Europe.) — Serio, D. —1993— La vegetazione marina bentonica Príroda (Praha) 36: 249-259. (in Czech). del Mediterraneo: I. Sopralitorale e Mesolitorale. Kol, E. —1942— The snow and ice algae of Alaska — Proposte di aggiornamento — Boll. Accad. Gioenia Sci. Smithson. Misc. Collect. 101(16): 1-36. Nat. 26: 245-291. Laundon, J.R. —1985— Desmococcus olivaceus. The name Giralt, M. —1996— Líquens epífits i contaminació atmos- of the common subaerial green alga — Taxon 34: 671-672. férica a la plana i les serrallades litorals tarragonines — Llimona, X. & Egea, J.M. —1984— La vegetación Arx. Sec. Ci., Sec. Ci. Biol. 113: 1-528. liquénica saxicola de los volcanes del Mar Menor Golubiç, S. —1967— Algenvegetation der Felsen. Eine (Murcia, SE de España) — Butll. Inst. Cat. Hist. Nat. ökologische Algenstudie im dinarischen Karstgebiet — 51, Sec. Bot. 5: 77-99. Binnengewässer 23: 1-183. Llimona, X. & Egea, J.M. —1985— Las comunidades Golubiç, S., Pietrini, A.M. & Ricci, S. —2015— Euendolithic liquenicas de las superficies de escorrentia de las rocas sil- activity of the cyanobacterium Chroococcus lithophilus iceas mediterraneas — An. Jard. Bot. Madrid 41: 429-444. Erc. In biodeterioration of the Pyramid of Caius Cestius, Margalef, R. —1944— Datos para la flora algológica de Rome, Italy — Int. Biodet. Biodegr. 100: 7-16. nuestras aguas dulces — Publ. Inst. Bot. 4(1): 1-130. Gustavs, L. —2010— Biodiversity and ecophysiology of Margalef, R. —1949— Las asociaciones de algas ei las aeroterrestrial green algae (Trebouxiophyceae, aguas dulces de pequeño volumen del noreste de España Chlorophyta) — Thesis, Univ. Rostock, Rostock. — Vegetatio 1: 258-284.

LAZAROA 36: 107-129. 2015 124 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes

Margalef, R. —1950— Datos para la hidrobiología de la Rindi, F. —2007— Diversity, distribution and ecology of cordillera cantábrica, especialmente del macizo de los green algae and cyanobacteria in urban habitats — In: Picos de Europa — Publ. Inst. Biol. Apl. 7: 37-76. Seckbach, J. (Ed.). Algae and cyanobacteria in extreme Margalef, R. —1951— Regiones limnológicas de Cataluña environments. Pp. 619-638. Springer, Dordrecht. y ensayo de sistematización de las asociaciones de algas Rindi, F. & Guiry, M.D. —2003— Composition and distri- — Collect. Bot. 3: 43-67. bution of subaerial algal assemblages in Galway City, Marstaller, R. —2006— Syntaxonomischer Konspekt der western Ireland — Cryptogamie Algol. 24: 245-267. Moosgesellschaften Europas und angrenzender Gebiete Roux, C. —1981— Étude écologique et phytosociologique — Haussknechtia Beiheft 13: 1-192. des peuplement lichéniques saxicoles-calcicoles du sud- Messikommer, E. —1927— Biologische Studien im est de France — Bibl. Lichenol. 15: 1-557. Torfmoor von Robenhausen unter besonderer Samad, L.K. & Adhikary, S.P. —2008— Diversity of Berücksichtigung der Algenvegetation — Mitt. Bot. Micro-algae and Cyanobacteria on Building Facades Mus. Univ. Zürich 122: 1-171. and Monuments in India — Algae 23: 91-114. Michael, E., Henning, B. & Kreisel, H. —1985— Schlüter, M. —1961— Die Diatomeen-Gesellschaften des Handbuch für Pilzfreunde. IV. Band. Blätterpilze- Naturschutzgebietes Strausberg bei Berlin — Int. Rev. Dunkelblättler — G. Fischer Verlag, Stuttgart. Ges. Hydrobiol. 46: 562-609. Möller, B. & Pankow, H. —1981— Algensoziologische und saprobiologische Untersuchungen an Vorflutern der Schuhwerk, F. —1986— Kryptogamengemeinschaften in Elbe — Limnologica 13: 291-350. Waldassoziationen - ein methodischer Vorschlag zur Möller, B. —1977— Algensoziologische und saprobiolo- Synthese — Phytocoenologia 14: 79-108. gische Untersuchungen an Vorflutern der Elbe — Täuscher, L. —1981— Untersuchungen zur Art- und Thesis, Univ. Rostock, Rostock. Biozönosestruktur des Phytoplanktons des Großen Moreno, P.P. & Egea, J.M. —1991— Biología y taxonomía Müggelsees (Berlin) unter Berücksichtigung produk- de la familia Lichinaceae, con especial referencia a las tions- und saprobiologischer Aspekte — Thesis, especies del S.E. español y Norte de África — Col. Humboldt Univ., Berlin. Blanca 19: 1-87. Täuscher, L. —1995— Hydrobotanische und ökologische Mulec, J. & Kosi, G. —2009— Lampenflora algae and meth- Untersuchungen an und in Gewässern des nördlichen ods of growth control — J. Cave Karst Stud. 71: 109-115. Elb-Havel-Winkels II. Garzer See und naturnahes Nowak, J. —1960— Naskalne zespoly porostów Wyxyny Kleingewässer — Untere Havel – Nat. Ber. 4: 3-11. Krakowsko-Cz stochowskiej. (Saxicolous associations Täuscher, L. —1997— Hydrobotanische und ökologische of lichens of the Cracow-Czestochowa Upland.) — Untersuchungen an und in Gewässern des nördlichen Fragmenta Flor. Geobot. 6: 323-392. (in Polish). Elb-Havel-Winkels IV. Die planktische und benthische Ochsner, F. —1928— Studien über die Epiphyten- Algenbesiedlung der unteren Havel (incl. Gnevsdorfer Vegetation der Schweiz — Jahrb. St. Gallischen Nat. Vorfluter) - Untere Havel —Nat. Ber. 6-7: 9-15. Ges. 63: 1-108. Täuscher, L. —1998— Mikroalgengesellschaften der Pankow, H. —1980— Die benthischen Gewässer Nordostdeutschlands und ihre Nutzung zur Kieselalgengesellschaften der Boddengewässer des Darß Bioindikation — Feddes Repert. 109: 617-638. und des Zingst (südliche Ostsee) — Wiss. Z. Wilhelm- Takeuchi, N. —2001— The altitudinal distribution of snow Pieck-Univ. Rostock, Mat.-nat. Reihe 29: 131-37. algae on an Alaska glacier (Gulkana Glacier in the Pankow, H., Haendel, D. & Richter, W. —1991— Die Alaska Range) — Hydrol. Proc. 15: 3447-3459. Algenflora der Schirmacheroase (Ostantarktika) — Takeuchi, N. —2013— Seasonal and altitudinal variations Nova Hedw. Beih. 103: 1-197. in snow algal communities on an Alaskan glacier Paus, S. M. —1997— Die Erdflechtenvegetation (Gulkana glacier in the Alaska range) — Environ. Res. Nordwestdeutschlands und einiger Randgebiete — Bibl. Lett. 8: 35002-35011. Lichenol. 66: 1-222. Pirk, W. & Tüxen, R. —1955— Das Coprinetum Von der Dunk, K. —1978— Beobachtungen an epiphyllen ephemeroidis, eine Pilzgesellschaft auf frischem Mist Moosen — Hoppea 37: 161-178. der Weiden im mittleren Wesertal— Mitt. Flor.-soziol. Weber, H.E., Moravec, J. & Theurillat, J.-P. —2000— Arbeit. N.F. 1: 1-7. International Code of Phytosociological Nomenclature. Pirk, W. & Tüxen, R. —1957— Das Trametetum gibbosae, 3rd edition — J. Veg. Sci. 11: 739-768. eine Pilzgesellschaft modernder Buchenstümpfe — Wilmanns, O. —1966— Kryptogamen-Gesellschaften Mitt. Flor.-soziol. Arbeit. N.F. 6-7: 120-126. oder Kryptogamen-Synusien? — In: Tüxen, R. (Ed.). Remias, D., Wastian, H., Lütz, C. & Leya, T. —2013— Gesellschaftsmorphologie. Pp. 1-7. Dr W. Junk Publ., Insights into the biology and phylogeny of Chloromonas The Hague. polyptera (Chlorophyta), an alga causing orange snow Wilmanns, O. —1998— Ökologische Pflanzensoziologie in Maritime Antarctica — Antarct. Sci. 25: 648-656. — Quelle & Meyer, Heidelberg.

125 LAZAROA 36: 107-129. 2015 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes

LAZAROA 36: 107-129. 2015 126 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes

Appendix 1: List of taxa (with authorities) used in this paper.

Lichens Lecanora dispersoareolata (Schaer.) Lamy Lecidea confluescens Nyl. Acarospora freyi H. Magn. Lecidea leprosolimbata (Arnold) Lettau ex Poelt Acarospora hospitans H. Magn. Lecidea speirodes Nyl. Acarospora impressula Th. Fr. Lecidea tessellata Flörke var. caesia (Anzi) Arnold Acarospora scabra (Pers.) Th. Fr. Lecidea umbonata (Hepp) Mudd Adelolecia kolaensis (Nyl.) Hertel et Rambold Lichinella cribellifera (Nyl.) P. P. Moreno et Egea Alyxoria mougeotii (A. Massal.) Ertz, Frisch et G. Thor Lichinella iodopulchra (Couderc ex Croz.) P. P. Moreno et Alyxoria variiformis (Anzi) Ertz Egea Alyxoria subelevata (Nyl.) Ertz et Tehler Metamelanea caesiella (Th. Fr.) Hennssen Anema decipiens (A. Massal.) Forssell Myriospora scabrida (Hedl. ex H. Magn.) K. Knudsen et Anema nummularium (Dufour ex Durieu et Mont.) Nyl. ex L. Arcadia Forssell Opegrapha cesareensis Nyl. Aspicilia candida (Anzi) Hue Opegrapha lutulenta Nyl. Aspicilia mashiginensis (Zahlbr.) Oxner Paralecanographa grumulosa (Dufour) Ertz et Tehler Aspicilia permutata (Zahlbr.) Clauzade et Rondon Peccania coralloides (A. Massal.) A. Massal. Aspicilia polychroma Anzi subsp. polychroma chemotype Peltula euploca (Ach.) Poelt verruculosa Peltula omphaliza (Nyl.) Wetmore Bacidia fraxinea Lönnr. Peltula placodizans (Zahlbr.) Wetmore Bacidia rubella (Hoffm.) A. Massal. Phaeophyscia orbicularis (Neck.) Moberg Bacidina phacodes (Körb.) Vfzda Physcia caesia (Hoffm.) Fürnr. Bactrospora patellarioides (Nyl.) Almq. Placynthium nigrum (Huds.) Gray Bellemerea subcandida (Arnold) Hafellner et Cl. Roux Placynthium subradiatum (Nyl.) Arnold Biatoridium monasteriense J. Lahm ex Körb. Placynthium tantaleum (Hepp) Hue Caloplaca anchon-phoeniceon Poelt et Clauzade Polyblastia ardesiaca (Bagl. et Carestia) Zschacke Caloplaca insularis Poelt Polyblastia cupularis A. Massal. var. cupularis Caloplaca vitellinula (Nyl.) H. Olivier Polysporina pusilla (Anzi) M. Steiner ex Kantvilas Candelariella aurella (Hoffm.) Zahlbr. Porpidia zeoroides (Anzi) Knoph et Hertel Candelariella vitellina (Hoffm.) Müll. Arg. Psorotichia diffracta (Nyl.) Forssell Carbonea atronivea (Arnold) Hertel Psorotichia frustulosa Anzi Coenogonium luteum (Dicks.) Kalb et Lücking Psorotichia schaereri (A. Massal) Arnold Collema ryssoleum (Tuck.) A. Schneid. Pterygiopsis affinis (A. Massal.) Henssen Dendrographa decolorans (Turner et Borrer ex Sm.) Ertz Rhizocarpon petraeum (Wulfen) A. Massal. et Tehler Rhizoplaca melanophthalma (DC.) Leuckert et Poelt Diplotomma dispersum (Kremp.) Arnold Rinodina confragosa (Ach.) Körb. Dirina massiliensis Durieu et Mont. morphotype mas- Roccella fuciformis (L.) DC. siliensis Roccella phycopsis (Ach.) Ach. Dirina massiliensis Durieu et Mont. morphotype sorediata Roccella tinctoria DC. Farnoldia micropsis (A. Massal.) Hertel Roccella tuberculata Vain. Gyalecta derivata (Nyl.) H. Olivier Roccellographa circumscripta (Taylor) Ertz et Tehler Gyalecta flotowii Körb. Rusavskia elegans (Link) S. Y. Kondr. & Kärnefelt Gyalecta liguriensis (Vfzda) Vfzda Teloschistes contortuplicatus (Ach.) Clauzade et Rondon Lecania erysibe (Ach.) Mudd Staurothele sapaudica Asta, Clauzade et Cl. Roux Lecanographa dialeuca (Cromb.) Egea et Torrente Staurothele solvens (Anzi) Zschacke Lecanora diaboli Frey et Poelt Thelidium ungeri (Flot.) Körb. Lecanora dispersa (Pers.) Sommerf. Thelopsis isiaca Stizenb.

127 LAZAROA 36: 107-129. 2015 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes

Thelopsis rubella Nyl. Coleofasciculus chthonoplastes (Thur. ex Gomont) M. Thyrea girardii (Durieu et Mont.) Bagl. et Carestia Siegesmund, J.R. Johans. & Friedl Verrucaria fischeri Müll. Arg. Corallina elongata J. Ellis & Sol. Corallina officinalis L. Non-lichenised fungi: Cryocystis brevispina (F.E. Fritsch) E. Kol ex Komárek & Fott Aleurodiscus disciformis (DC.) Pat. Cryptopleura ramosa (Huds.) L. Newton Dendrominia dryina (Pers.) Ghobad-Nejhad & Duhem Delesseria sanguinea (Huds.) J.V. Lamour. Didymosphaeria rubicola Berl. Desmarestia aculeata (L.) J.V. Lamour. Navicella pileata (Tode) Fabre Desmococcus olivaceus (Pers. ex Ach.) J.R. Laundon Septobasidium quercinum (De Not. & Bagl.) Sacc. Desmotetra antarctica (Fritsch) H.U. Ling Diatoma vulgaris Bory Algae & cyanobacteria: Dilsea carnosa (Schmidel) Kuntze Ellerbeckia arenaria (Moore ex Ralfs) R.M. Crawford Achnanthes brevipes C. Agardh Ellisolandia elongata (J.Ellis & Sol.) K.R. Hind & G.W. Actinastrum hantzschii Lagerh. Saunders Agarum clathratum Dumort. Fimbrifolium dichotomum (Lepech.) G.I. Hansen Alaria esculenta (L.) Grev. Fucus ceranoides L. Amphora coffaeiformis (C. Agardh) Kütz. Fucus serratus L. Amphora ovalis (Kütz.) Kütz. Fucus spiralis L. Ancylonema nordenskioeldii Berggr. Fucus vesiculosus L. Anomoeoneis sphaerophora Pfitzer Furcellaria lumbricalis (Huds.) J.V. Lamour. Gelidium pulchellum (Turner) Kütz. Apatococcus lobatus (Chodat) J.B. Petersen Gigartina pistillata (S.G.Gmel.) Stackh. Aphanothece stagnina (Spreng.) A. Braun Gloeocapsa biformis Erceg. Apoglossum ruscifolium (Turner) J. Agardh Gloeocapsa compacta Kütz. Ascophyllum nodosum (L.) Le Jolis Gloeocapsa kuetzingiana Nägeli ex Kütz. Asterionella formosa Hassall Gloeocapsa sanguinea (C.Agardh) Kütz. Bifurcaria bifurcata R. Ross Gomphonema parvulum (Kütz.) Kütz. Boergeseniella thuyoides (Harv.) Kylin Halopteris scoparia (L.) Sauv. Bonnemaisonia hamifera Har. Halurus equisetifolius (Lightf.) Kütz. Bostrychia scorpioides (Huds.) Mont. Halurus flosculosus (J. Ellis) Maggs & Hommers. Brachytrichia quoyi Bornet & Flahault Heterosiphonia plumosa (J. Ellis) Batters Brebissonia boeckii (Ehrenb.) E. OMeara Himanthalia elongata (L.) S.F. Gray Calliblepharis ciliata (Huds.) Kütz. Klebsormidium flaccidum (Kütz.) P.C. Silva, K.R. Mattox Calliblepharis jubata (Gooden. & Woodw.) Kütz. & W.H. Blackw. Callithamnion tetragonum (With.) Gray Laminaria digitata (Huds.) J.V. Lamour. Callithamnion tetricum (Dillwyn) Gray Laminaria hyperborea (Gunnerus) Foslie Callophyllis laciniata (Huds.) Kütz. Lomentaria articulata (Huds.) Lyngb. Caloneis amphisbaena (Bory) Cleve Lomentaria clavellosa (Lightf. ex Turner) Gaillon Calothrix scopulorum C.Agardh ex Bornet & Flahault Lyngbya aestuarii Liebman ex Gomont Catenella caespitosa (With.) L.M. Irvine Mastocarpus stellatus (Stackh.) Guiry Caulerpa prolifera (Forssk.) J.V. Lamour. Melosira varians C.Agardh Caulerpa racemosa (Forssk.) J. Agardh Membranoptera alata (Huds.) Stackh. Ceramium virgatum Roth Meridion circulare (Grev.) C. Agardh Chlamydomonas nivalis (F.A. Bauer) Wille Mesotaenium berggrenii (Wittr.) Lagerh. Chloromonas nivalis (Chodat) Hoham & Mullet Micractinium pusillum Fresen. Chondracanthus acicularis (Roth) Fredericq Microcoleus autumnalis (Gomont) Strunecky, Komárek & Chondrus crispus Stackh. J.R. Johans. Chylocladia verticillata (Lightf.) Bliding Mucidosphaerium pulchellum (H.C. Wood) C. Bock, Cladophora rupestris (L.) Kütz. Pröschold & Krienitz Claviclonium ovatum (J.V. Lamour.) Kraft & Min-Thein Mycacanthococcus cellaris Hansgirg f. antarcticus Wille Closterium limneticum Lemmerm. Navicula salinarum Grunow Cocconeis placentula Ehrenb. Navicula tripunctata (O.F. Müller) Bory Coccotylus truncatus (Pall.) M.J. Wynne & J.N. Heine Odonthalia dentata (L.) Lyngb. Codium tomentosum Stackh. Oscillatoria limosa C.Agardh ex Gomont

LAZAROA 36: 107-129. 2015 128 Helga Bültmann & al. Validations and descriptions of European syntaxa of vegetation dominated by thallophytes

Oscillatoria tenuis C.Agardh ex Gomont Protoderma cohaerens (Wittrock) Printz Osmundea pinnatifida (Huds.) Stackh. Pseudanabaena limnetica (Lemmerm.) Komárek Palmaria palmata (L.) F. Weber & D. Mohr Ptilota gunneri P.C. Silva, Maggs & L.M. Irvine Pandorina morum (O.F. Müller) Bory Raphidonema brevirostre Scherffel Pediastrum duplex Meyen Raphidonema nivale Lagerh. Pelvetia canaliculata (L.) Decne. & Thur. Rhoicosphenia abbreviata (C. Agardh) Lange-Bert. Phormidium chalybeum (Mert. ex Gomont) Anagn. & Rivularia nitida C. Agardh ex Bornet & Flahault Komárek Rosenvingiella radicans (Kütz.) Rindi, L.McIvor & Guiry Phycodrys rubens (L.) Batters Saccorhiza polyschides (Lightf.) Batters Phyllophora crispa (Huds.) P.S. Dixon Scenedesmus quadricauda (Turpin) Bréb. Phyllophora pseudoceranoïdes (S.G. Gmel.) Newroth & Schizogonium murale Kütz. A.R.A. Taylor Schizothrix heufleri Grunow ex Gomont Planothidium lanceolatum (Brébisson ex Kützing) Lange- Scytonema myochrous C.Agardh ex Bornet & Flahault Bert. Smithsonimonas abbotii Kol Pleonosporium borreri (Smith) Nägeli Spirulina subsalsa Oerst. ex Gomont Plocamium cartilagineum (L.) P.S. Dixon Stenogramma interruptum (C. Agardh) Mont. Plumaria plumosa (Huds.) Kuntze Trentepohlia abietina (Flotow ex Kütz.) Hansg. Polysiphonia arctica J. Agardh Trentepohlia umbrina (Kütz.) Bornet Polysiphonia stricta (Dillwyn) Grev. Tryblionella hungarica (Grunow) Freng. Porphyridium purpureum (Bory) K.M. Drew & R. Ross Ulnaria ulna (Nitzsch) P. Compère Prasiola crispa (Lightfoot) Kütz. Vertebrata lanosa (L.) T.A. Chr.

Received: 16 November 2015 Accepted: 23 November 2015

129 LAZAROA 36: 107-129. 2015