107 Geobotanical characterisation of the Trifolion squamosi nom. corr. alliance (class. Juncetea maritimi) in Catalonia and the Western Mediterranean Abstract We present a summary of part of the phytogeographic results described in the author’s PhD thesis. In this case, we characterise and validate several syntaxa of subsaline meadows in the Trifolion squamosi nom. corr. alliance (class. Juncetea maritimi) in Catalonia (incl. NE Spain and SE France) and in other regions of the Western Mediterranean (N Algeria, Languedoc and Provence). In total, we study six syntaxa in geobotanical terms, based on synthetic tables and factor analyses of correspondence published previously in the thesis: one alliance (Trifolion squamosi), two associations and three subassociations (Agropyro-Trifolietum subass. typicum, subass. festucetosum comb. nova, subass. brachypodietosum nova; Festucetum arundinaceae). Iz vleček V članku predstavljamo del rezultatov fitogeografske raziskave avtorjeve doktorske naloge. Nomenklaturno smo ovrednotili subsalina travišča zveze Trifolion squamosi nom. corr. alliance (class. Juncetea maritimi) v Kataloniji (vključno z SV Španijo in JV Francijo) in ostalih delih zahodnega Sredozemlja (severna Alžirija, Languedoc in Provansa). Šest sintaksonov smo preučili geobotanično na podlagi sintetskih tabel in faktorske korespondenčne analize, objavljene v doktorski nalogi: zvezo (Trifolion squamosi), dve asociaciji in tri subasociacije (Agropyro-Trifolietum subass. typicum, subass. festucetosum comb. nova, subass. brachypodietosum nova; Festucetum arundinaceae). Key words: Catalonia, Western Mediterranean, geobotanical characterisation, syntaxonomic validation, subsaline meadows, Trifolion maritimi nom. inept., Trifolion squamosi nom. corr. Ključne besede: Katalonija, zahodno Sredozemlje, geobotanične značilnosti, sintaksonomsko ovrednotenje, subsalina travišča, Trifolion maritimi nom. inept., Trifolion squamosi nom. corr. Received: 30. 7. 2020 Revision received: 1. 9. 2020 Accepted: 8. 9. 2020 1 Herbarium of the University of Girona. Department Environmental Science, Montilivi campus, C/M. Aurèlia Campmany 69, E-17003 Girona, Catalonia, Spain. Email: g.mercadal.corominas@gmail.com Gabriel Mercadal i Corominas1  DOI: 10.2478/hacq-2020-0019 20/1 • 2021, 107–117 20/1 • 2021, 107–117 108 Gabriel Mercadal i Corominas Geobotanical characterisation of the Trifolion squamosi nom. corr. alliance (class. Juncetea maritimi) in Catalonia and the Western Mediterranean Introduction In 2019, at the University of Girona (Catalonia, Spain), the author of this article defended his PhD thesis on hay meadows in the Mediterranean region of Catalonia (Mercadal 2019a, 2019b, 2019c). However, as that docu- ment was a monograph, the new syntaxa described are invalid, in accordance with Article 1 of the International Code of Phytosociological Nomenclature (ICPN) (We- ber et al. 2000, Theurillat et al. 2020). For that reason, herein we validate and geobotanically characterise sever- al syntaxa that include regularly mown subsaline mead- ows of the Trifolion squamosi nom. corr. alliance (class. Juncetea maritimi) in the Western Mediterranean, as we have done previously for other syntaxonomical units (Mercadal 2018, 2020a, 2020b, 2020c, 2020d). Materials and Methods We geobotanically characterised six syntaxa (one alliance, two associations and four subassociations) in the Trifolion squamosi nom. corr. alliance (class. Juncetea maritimi), presented previously in the author’s PhD thesis (Mercadal 2019c: T able 120). The study area covers the Mediterranean Biogeo- graphic Region of NE Catalonia (incl. NE Spain and SE France), more specifically the Ruscinic territory, which includes the regions of Roussillon, Alt Empordà and Baix Empordà (Figures 1a-1b). Detailed geobotanical in- formation on the meadows mapped in each region can be consulted in Mercadal (2019a). We followed the traditional sygmatist phytosociologi- cal method (Braun-Blanquet 1964, 1979), as well as a synthetic classification. We compared our relevés using synthetic tables and factor analyses of correspondence with those of various related plant communities in the Western Mediterranean (N Algeria, Catalonia, Langue- doc and Provence, cf. Mercadal 2019c: Table 120). In this article, we reference these analytical data and other ecological and chorological information by directly cit- ing the thesis. We reproduce only a few maps and one ta- ble that we consider essential for understanding the text (Figures 1 and 2 & Table 1). The syntaxonomic abbreviations used are those pro- posed by Mucina et al. (2016) or, failing that, those pro- posed by Theurillat et al. (2020); all these are listed in Mercadal (2019c). With regard to the nomenclature of the taxa, we follow the system used in Mercadal (2019b), which basically corresponds to that found in Bolòs et al. (2005) or, failing that, the nomenclature used by Tison & Foucault Figure 1: Location of the study area ( ● – incl. NE of Spain and SE of France) in the Western Mediterranean (a-b) and distribution of some syntaxa on the regional map of northeast Catalonia (c-f). Modified from Mercadal (2019c). Dashed line, relevé comparison area; map grid, UTM coordinate system (10 × 10 km) [UTM zone, 31T]. Slika 1: Lokacija preučevanega območja ( ● – skupaj s SV Španijo in JV Francijo) v zahodnem Sredozemlju (a-b) in razširjenost nekaterih sintaksonov na regionalnem zemljevidu SV Katalonije (c-f). Spremenjeno po Mercadal (2019c). Prekinjena črta prikazuje območje primerjave popisov; mreža je UTM koordinatni sistem (10 × 10 km) [UTM zone, 31T]. Regional map Agropyro-Trifolietum AT typicum AT festucetosum AT brachypodietosum (2014) or Castroviejo (1986-2019). Exceptionally, we followed other works, which are indicated in Mercadal (2019b: Table 3). 20/1 • 2021, 107–117 109 Gabriel Mercadal i Corominas Geobotanical characterisation of the Trifolion squamosi nom. corr. alliance (class. Juncetea maritimi) in Catalonia and the Western Mediterranean Results and Discussion Class. Juncetea maritimi Br.-Bl. in Br.-Bl., Roussine et Nègre 1952 [cf. Mercadal 2019c: 485, 2020c] Ord. Juncetalia maritimi Br.-Bl. ex Horvatić 1934 [cf. Mercadal 2019c: 488, 2020c] 1. All. Trifolion squamosi Br.-Bl. in Br.-Bl., Roussine et Nègre 1952 nom. corr. [cf. Mercadal 2019c: 567] Synonyms: Trifolion maritimi Br.-Bl. 1931 nom. inval. (Art. 2b); Trifolion maritimi Br.-Bl. 1932 nom. inval. (Art. 2b); Trifolion maritimi Br.-Bl. 1950 nom. inval. (Art. 2b); T rifolion maritimi Br.-Bl. ex Molinier et T allon 1950 nom. inval. (Art. 2b); Trifolion maritimi Br.-Bl. in Br-Bl. et al. 1952 nom. inept. (Art. 44); Trifolion squa- mosi Br.-Bl. ex Julve 1993 p. p. nom. inval. (Art. 2b). Homonym: Trifolion maritimi Br.-Bl. ex Simonneau 1952 nom. illeg. (Art. 31). Corresponding name: Tri- folienion squamosi (Br.-Bl. in Br.-Bl. et al. 1952) Rivas Goday et Rivas-Mart. 1963 nom. corr. (Art. 44) Holotypus: Agropyro pycnanthi-Trifolietum maritimi Br.- Bl. in Br.-Bl. et al. 1952 nom. inept. (Art. 44). Physiognomy and floristic composition : subhalophilic and mesohygrophilous meadows, 40 to 80 (150) cm in height, typically with 100% cover. The most abundant life form corresponds to the hemicryptophytes (43%); however, the alliance stands out due to the elevated presence of therophytes (39%). Most of the plants have a Pluriregional (57%) or Mediterranean distribution (28%) (Mercadal 2019c: Figs. 253-254). The dominant taxa are, basically, leguminous plants, most of which are annual (T rifolium squamosum, T . pratense, T . resupinatum, Tetragonolobus maritimus, Melilotus siculus, M. indicus, M. segetalis), and perennial grasses (Elymus athericus, E. campestris, Hordeum secalinum, Festuca arundinacea, F. interrupta) of forage value. These plants are usually accompanied by several different taxa in the class Juncetea maritimi (Carex distans, C. divisa, Limonium narbonense, Alopecurus bulbosus, Juncus compressus subsp. gerardi, J. maritimus) and meadow species with a wide ecological spectrum (Poa pratensis, P. trivialis, Cynodon dactylon, Rumex crispus, etc.) (Mercadal 2019c: Table 138). Characteristic and differential species : Trifolium squa- mosum, T . resupinatum (diff.), T etragonolobus maritimus (diff.), Melilotus siculus, M. indicus, M. segetalis, Medi- cago scutellata (terr. charact. Languedoc), Hordeum se- calinum, Vicia tetrasperma subsp. gracilis, V . lutea subsp. lutea var. hirta, Bellevalia romana (terr. charact. Rous- sillon), Bromus commutatus (diff.), Sonchus asper (diff.), Festuca interrupta (diff.), Tragopogon porrifolius (diff. Languedoc), T . lamottei (diff. Catalonia). Ecology: saline or brackish land near the sea, with an alkaline pH, clay or sandy-clay texture and season- ally flooded. The saline concentration varies over the course of the year: in the rainiest periods, the salinity of the first few centimetres of soil decreases thanks to the precipitation and irrigation. On the other hand, in the summer (and during certain dry periods), the saline concentration increases, but by that time the therophytes of forage value have already wilted or been mown (Adriani 1934, Braun-Blanquet et al. 1952). Generally, the meadows are mown regularly to ob- tain high quality forage, although sometimes they are mown a second time, grazed, irrigated, fertilised and sown again. Geographical distribution: Provence, Languedoc, north- east of Catalonia and North Africa (Algeria) (Figure 2). Most likely, it must also be distributed in other regions of the Western Mediterranean coast. Figure 2: Distribution of the Trifolion squamosi alliance in the Western Mediterranean (Mercadal 2019c: Fig. 255). Dashed line, potential area of the alliance. A, Algeria; C, Catalonia; L, Languedoc, P , Provence. Slika 2: Razširjenost zveze Trifolion squamosi v zahodnem Sredozemlju (Mercadal 2019c: Fig. 255). Prekinjena črta prikazuje potencialno razširjenost zveze. A, Alžirija; C, Katalonija; L, Languedoc, P , Provansa. Syntaxonomy: the alliance was validated simultane- ously by Braun-Blanquet et al. (1952) and Simonneau (1952), resulting in two homonyms being erected with the same date (we do not know the months in which the two papers were published). Applying Article 33 of the ICPN, we choose the correct name as being Trifoli- on maritimi Br.-Bl. in Br.-Bl. et al. 1952 and reject Tri- folion maritimi Br.-Bl. ex Simonneau 1952, which au- tomatically becomes an illegitimate synonym (Art. 31). Later, we correct the name of the alliance according to Article 44, since the correct name of Trifolium mariti- mum Huds. (the name used by Braun- Blanquet to de- 20/1 • 2021, 107–117 110 Gabriel Mercadal i Corominas Geobotanical characterisation of the Trifolion squamosi nom. corr. alliance (class. Juncetea maritimi) in Catalonia and the Western Mediterranean scribe the alliance) currently corresponds, according to all western European floras, to Trifolium squamosum L. (Castroviejo 1986–2019, Bolòs & al. 2005, Lambinon & Verloove 2012, Stace 2014, Tison & Foucault 2014, Pignatti & al. 2017...). We give priority to the authorship of Br.-Bl. in Br.-Bl. et al. 1952, since their type, Agropyro-Trifolietum (Art. 18a) corresponds to the traditional phytosociological concept of the alliance. On the other hand, the type of the rejected name, Trifolion maritimi Br.-Bl. ex Simon- neau 1952, is the association Festucetum arundinaceae, a highly artificialised plant community with a floristic composition very far removed from the classic notion of Trifolion maritimi nom. inept. On the other hand, it should be noted that Julve (1993: 90) made significant amendments to the Trifo- lion maritimi nom. inept. alliance [sub Trifolion squa- mosi Br.-Bl. 1931 em. Julve 1993] and included in it the hygrophilous Atlantic-Mediterranean associations that Foucault & Catteau (2012) would later use to de- scribe the alliance Ranunculo-Oenanthion. Moreover, Julve excluded the alliance Trifolion maritimi nom. inept. from the class Juncetea maritimi and attributed it to the order Eleocharietalia palustris and to the class Agrostio-Arrhenatheretea; however, surprisingly, he did not include Agropyro-Trifolietum. In our opinion, this union of associations and the change of order and class is not very coherent in phytosociological terms, as indicated by Foucault & Catteau (2012). In fact, the change is so great that Mucina et al. (2016) interpret Trifolion squamosi (Br.-Bl. 1931) em. Julve 1993 as an invalid alliance different from Trifolion maritimi nom. inept. [sub Trifolion squamosi Julve 1993]. Later, Géhu (1999) attributed Agropyro-Trifolietum to the alliance Alopecurion utriculati, to which he also added the “Prairies humides thermophiles, d’optimum méditerranéo-(sub)atlantique, souvent à tendance olig- ohaline”, which Julve (1993) had previously included in Trifolion squamosi. Julve (eVeg 2020) currently follows Géhu’s criteria [sub Alopecurion rendlei Zeilder 1954]. Finally, it should be noted that Mucina et al. (2016) maintained Trifolion maritimi nom. inept. as an inde- pendent syntaxonomic unit, but attributed it to the order Potentillo-Polygonetalia and the class Molinio-Ar- rhenatheretea. In our opinion, this syntaxonomic attri- bution is too synthetic, with the intention of grouping together all the regularly mown Atlantic-Mediterrane- an brackish meadows in the same order. Along these lines, Mucina et al. (2016) consider the nitrophilous alliance Trifolio-Cynodontion as a syntaxonomic syno- nym of Trifolion maritimi nom. inept., something that we do not agree with either; this led to Guarino & Pasta (2017) attributing Kickxio-Trifolietum to T ri folion maritimi nom. inept., despite it being a nitrophilous community belonging to Trifolio-Cynodontion. Variability: within the alliance, two associations can be differentiated: Agropyro-Trifolietum [#1.1] and Festuce- tum arundinaceae [#1.2]. However, we also analysed the Lino-Gaudinietum (class. Stipo-Agrostietea) relevés of Franquesa (1995) due to their floristic affinity (Mer- cadal 2019c: Table 138 & Figs. 256-257). 1.1 Ass. Agropyro pycnanthi-Trifolietum maritimi Br.-Bl. in Br.-Bl., Roussine et Nègre 1952 nom. inept. (Art. 44) [cf. Mercadal 2019c: 581] Synonyms: Ass. à Trifolium maritimum et Agropyrum pycnanthum Br.-Bl. 1931 nom. inval. (Art. 2b) (orig. form); Association Trifolium maritimum-Agropyrum pycnanthum Br.-Bl. 1932 nom. inval. (Art. 2b) (orig. form); Trifolietum maritimi Adriani 1934 nom. inval. (Art. 2b); Trifolietum maritimi Soroceanu 1936 nom. inval. (Art. 2b); Asociación Trifolium maritimum-Ag- ropyrum pycnanthum Br.-Bl. 1950 nom. inval. (Art. 2b) (orig. form); Trifolietum maritimi Molinier et Tallon 1950 nom. inval. (Art. 2b); Trifolietum maritimi Br.- Bl. 1951 nom. inval. (Art. 2b); Agropyreto-Trifolietum maritimi Br.-Bl. in Br.-Bl. in al. 1952 (orig. form) [sub Agropyreto-Trifolietum maritimi Br.-Bl. 1931; Ass. à Trifolium maritimum et Agropyrum pycnanthum (acu- tum) Br.-Bl. 1931]. Neotypus hoc loco designatus: Mercadal (2019c), un- published PhD thesis, Table 147, rel. 6, p. 603 (France, Catalonia, Roussillon, Sant Nazari, 1 m, 31TDH9924, 31.04.2017). Plot size, 100 m 2 ; height, 40 cm; total cover, 100%: Hordeum secalinum 1.1, Trifolium squam- osum 3.3, T . pratense 1.1, Festuca interrupta 1.1, Melilo- tus segetalis 1.1, M. siculus 3.3, M. indicus 1.1, Sonchus asper +, Tragopogon lamottei +, Vicia tetrasperma subsp. gracilis +, V. lutea subsp. lutea var. hirta +, V. segetalis +, Bellevalia romana +, Elymus athericus 2.2, E. campestris 1.1, Carex divisa 2.2, C. distans 1.1, C. flacca +, Nar- cissus tazetta subsp. tazetta +, Alopecurus bulbosus 1.2, Scorzonera laciniata +, Plantago crassifolia +, Orchis palustris +, Phragmites australis 1.1, Bromus hordeaceus subsp. hordeaceus +, Gaudinia fragilis +, Linum usitatis- simum subsp. angustifolium +, Picris echioides +, Althaea officinalis +, Bellis perennis +, Galactites tomentosa +, Lathyrus aphaca +, Ophrys apifera +, Poa pratensis subsp. angustifolia +, Sherardia arvensis +. Physiognomy and floristic composition : subhalophilic and mesohygrophilous meadows, 40 to 80 (150) cm in height, dominated by Festuca gr. arundinacea, Trifo- lium squamosum, Tetragonolobus maritimus, Hordeum secalinum, Bromus hordeaceus subsp. hordeaceus or 20/1 • 2021, 107–117 111 Gabriel Mercadal i Corominas Geobotanical characterisation of the Trifolion squamosi nom. corr. alliance (class. Juncetea maritimi) in Catalonia and the Western Mediterranean Melilotus siculus. These plants are accompanied by sev- eral species characteristic of the class Juncetea maritimi (Limonium narbonense, Carex divisa, C. distans, Juncus compressus subsp. gerardi, Alopecurus bulbosus, Festuca interrupta, Elymus athericus, etc.), which are abundant to varying degrees depending on the level of soil mois- ture. Halophytes are more frequent in more natural and saline meadows; on the other hand, in meadows where there is greater human activity (well irrigated and fertilised), different taxa of Molinio-Arrhenathere- tea are more frequent, such as Poa trivialis, Holcus la- natus, Dactylis glomerata, and so on (Mercadal 2019c: Tables 138, 144). Floristically, the association is moderately rich (24 taxa/rel. on average) and has a high percentage of hemicryptophytes (42%) and therophytes (40%). The taxa are mainly Pluriregional (57%) or Mediterranean (28%) (Mercadal 2019c: Figs. 262, 264). Characteristic and differential species : Trifolium squamosum, Melilotus siculus, M. indicus, M. segetalis, Medicago scutellata (terr. charact. Languedoc), Horde- um secalinum, Vicia tetrasperma subsp. gracilis, V. lutea subsp. lutea var. hirta, Bellevalia romana (terr. charact. Roussillon), Tragopogon porrifolius (diff. Languedoc), T. lamottei (diff. Catalonia), Festuca interrupta (diff.), Bromus commutatus (diff.), Elymus athericus (Link) Kerguélen [= Agropyrum pycnanthus (Godr.) Gren. et Godr. and probably A. acutum (DC.) Roem. et Schult. ] (diff.). Ecology: subsaline soils in the Mediterranean coastal region, with a clay or sandy-clay texture, alkaline and seasonally flooded. The meadows are mown and grazed regularly, although sometimes they are also irrigated, fertilised or resown. Geographical distribution: well-known association of the western European Mediterranean: Provence (Cam- argue), Languedoc (around Montpellier) and Catalonia (plains of Roussillon and Empordà). Most likely, the association must also be developed by the wetlands of Narbonne and Sérignan (Languedoc). In Catalonia, it is quite widespread along the Ruscinic coast, between 0–16 (85) m of altitude (Figure 1c & Mercadal 2019c: Fig. 267). Syntaxonomy: association validated in Braun-Blanquet et al. (1952: 121) through the publication of a synthet- ic relevé based on 29 unpublished association relevés from the Languedoc coastal region. For this reason, we designate as a neotype one of our own relevés from Roussillon, a little over 100 km from the association’s locus classicus. Although the name of the association is a nomen ineptum, we did not correct it according to article 44 because we are not 100% sure what the cor- rect name is for Agropyrum pycnanthus (Godr.) Gren. et Godr. nor from A. acutum (DC.) Roem. et Schult. (Tison & Foucault 2014: 232–234, E. Rico pers. com- mun.), the two names that Braun-Blanquet used to describe Agropyro-Trifolietum (cf. synonyms). In this case, the name must be submitted to the Committee for Change and Conservation of Names (CCCN) for a final decision. However, in our opinion, the taxon would correspond to Elymus athericus (cf. Mercadal 2019b: 763) and the correct name of the association would be Elymo atherici-Trifolietum squamosi. Affinities : Agropyro-Trifolietum presents certain floristic affinities with several associations. • With the Oenantho-Gaudinion alliance (Geranio- Festucetum subass. caricetosum and Gaudinio-Arrhe- natheretum subass. lotetosum). Agropyro-Trifolietum constitutes a bridge association between the salt meadows of the order Juncetalia maritimi and the non-saline meadows of the order Trifolio-Hordeeta- lia, and for this reason shares several species with the Oenantho-Gaudinion associations that can withstand slightly saline soils (Trifolium squamosum, Hordeum secalinum, Bromus commutatus, Gaudinia fragilis, Li- num usitatissimum subsp. angustifolium, etc.); how- ever, these are generally easy to differentiate due to an elevated presence of halophytes. • With Lino-Gaudinietum, an association endemic to Cap de Creus (Alt Empordà, NE Catalonia) domi- nated by Gaudinia fragilis and Linum usitatissimum subsp. angustifolium. This slightly brackish small meadow is characterised by several small clover spe- cies (Trifolium nigrescens, T. dubium, T. micranthum, T. strictum) and by Agrostis capillaris s.l. It grows in small despressions in siliceous soils that are season- ally flooded and periodically sprayed by droplets of wind-borne sea water. Its phytosociological attribu- tion is complex, as indicated by Franquesa (1995). Franquesa gave priority to the annual calcifuge taxa and attributed the association to the class Tuberari- etea guttatae, as a transitional community towards Molinio-Arrhenatheretea. However, she also saw af- finities with Agrostion salmanticae (class. Isoëto-Na- nojuncetea) from Extremadura (SW Spain). On the other hand, Rivas-Martínez et al. (2001, 2011) at- tributed Lino-Gaudinietum to the alliance Agrostion castellanae (ord. Agrostietalia castellanae, class. Stipo- Agrostietea). In our opinion, Lino-Gaudinietum is a transitional community between Agrostion castellanae and Trifo- lion squamosi, as shown by Mercadal (2019c: Table 138). However, due to its particular ecology (small calcifuge meadows), the low presence of taxa in the 20/1 • 2021, 107–117 112 Gabriel Mercadal i Corominas Geobotanical characterisation of the Trifolion squamosi nom. corr. alliance (class. Juncetea maritimi) in Catalonia and the Western Mediterranean class Juncetea maritime, and its unique floristic com- position (with several acidophilic taxa), we will keep it within Agrostion castellanae. • With the association Trifolietum nigrescenti-resupi- nati described in the Camargue (Provence) and dominated and characterised by Trifolium resupina- tum, T. nigrescens, T. ornithopodioides and T. camp- estre (Molinier & Tallon 1949, 1950, 1968). These slightly brackish small meadows contain several taxa in the class Juncetea maritimi (Limonium narbon- ense, Atriplex portulacoides, Trifolium squamosum, Elymus athericus), meaning that they are similar in floristic terms to the alliance Trifolion squamosi (Mo- linier & Tallon 1950), constituting a transitional association between Trifolion squamosi and Trifolio- Cynodontion. However, the lack of halophytes typi- cal of Trifolion squamosi leads us to attribute it to Trifolio-Cynodontion, as proposed by Foucault & Catteau (2012). Variability: within the association we distinguish three associations according to the edaphic characteristics and the agricultural practices (Mercadal 2019c: Tables 138, 144 & Fig. 268): subass. typicum [#1.1.1], subass. festucetosum comb. nova [#1.1.2] and subass. brachypo- dietosum subass. nova [#1.1.3]. 1.1.1 Ass. Agropyro-Trifolietum subass. typicum [cf. Mercadal 2019c: 598] Physiognomy and floristic composition : subhalophilic meadow, ranging from mesohygrophilous to hygro- philous, 40 to 80 (150) cm in height, dominated by leguminous plants (Trifolium squamosum, T. resupina- tum, Melilotus siculus, M. indicus, Melilotus segetalis) and grasses of forage value (Elymus athericus, E. camp- estris, Festuca interrupta, F. arundinacea). These plants are generally accompanied by different taxa of the class Juncetea maritime: Limonium narbonense, Scorzonera laciniata, Alopecurus bulbosus, Carex divisa, T etragonolo- bus maritimus, Juncus compressus subsp. gerardi, J. mar- itimus, etc. (Mercadal 2019c: Table 147). Differential species : Melilotus siculus, M. indicus, Medi- cago scutellata (terr. diff. Languedoc), Festuca interrupta, Elymus athericus, Orchis palustris, O. ×lloydiana. Ecology: land next to the sea, often surrounded by marshes. This is the most salinised subassociation (ranging from slightly to moderately saline), which oc- curs on alkaline soils, with a clay loam texture, high humidity and a variable carbonate content (low in the Roussillon region and high in the Languedoc region). The meadows are mown and grazed regularly, and they flood naturally each year (Table 1). Table 1: Differential floristic, ecological and chorological characteristics of the Agropyro-Trifolietum (AT) subassocia- tions. The soil data correspond to the study area. Tabela 1: Razlikovalne floristične, ekološke in horološke značilnosti subasociacij asociacije Agropyro-Trifolietum (AT). Podatki o tleh veljajo za preučevano območje. Characteristics AT typicum AT festucetosum AT brachypodietosum Differential species Melilotus siculus, M. indicus, Medicago scutellata, Festuca interrupta, Elymus athericus, Orchis palustris, O. ×lloydiana Poa trivialis subsp. trivialis, Lotus corniculatus subsp. corniculatus, Agrostis stolonifera, Holcus lanatus, Oenanthe fistulosa, Polypogon monspeliensis, Ranunculus sardous s.l., Lychnis flos-cuculi, Leontodon taraxacoides subsp. taraxacoides Brachypodium phoenicoides, Avena barbata, Inula viscosa, Hypochaeris radicata, Rubus ulmifolius, Cirsium vulgare, Ophrys apifera subsp. apifera, Orobanche crenata, Anacamptis pyramidalis, Barlia robertiana pH 1 : 2.5 H 2 O 7.8-9.9 [8.8] 7.6-9.0 [8.0] 7.7-9.5 [8.4] Ground conductivity 1 : 5 25 °C (µS/cm) No data (> ATf) Slightly saline-moderate saline soils 249.2-1842.7 [741.2] Slightly saline soils 165.1-668.6 [345.8] Non saline-slightly saline soils T otal carbonate content (%) 0.0-3.0 [1.3] (Languedoc: 17.0-40.0%) 0.0-2.7 [0.6] 15.1-32.0 [23.6] Organic matter (%) 1.3-3.6 [2.5] 1.6-5.2 [3.6] 1.3-5.8 [3.2] Soil texture (ISSS) Sandy clay loam Clay Sandy clay Soil moisture (%) No data (> ATf) (hygrophilous to mesohygro philous meadows) 14.0-35.4 [26.6] (mesohygrophilous meadows) 13.7-25.1 [20.9] (mesohygrophilous to mesophilic meadows) Agricultural activities Mowing and grazing Mowing, grazing, fertiliser, sowing and irrigation Mowing and grazing Geographical distribution Languedoc-Roussillon coast Provençal-Catalan coast Empordà plain (Catalonia) ISSS, International Society of Soil Science; in square brackets, the mean. 20/1 • 2021, 107–117 113 Gabriel Mercadal i Corominas Geobotanical characterisation of the Trifolion squamosi nom. corr. alliance (class. Juncetea maritimi) in Catalonia and the Western Mediterranean Geographical distribution: we only know this sub- association around Montpellier (Languedoc) and the Roussillon plain; however, it must be distributed along the coast of the Gulf of Leon, from Provence to Roussillon. In the study area, we only observed it in the lake of Sant Nazari (Roussillon), between 0-1 m of altitude, but it is necessary to prospect more the plain of the Empordà (Figure 1d; Mercadal 2019c: Fig. 272). Syntaxonomy: autonym that we create automatically in order to attribute new subassociations to Agropyro- Trifolietum. The authorship, year of publication and nomenclature type coincide with the data relating to the association. Variability: in the study area, we can distinguish two variants and different facies in a similar manner to Braun-Blanquet et al. (1952) in Languedoc. The var. Trifolium squamosum, the type, (Mercadal 2019c: Ta- ble 147, rel. 1-9), which can represent several facies in accordance with the dominant plant (f. Trifolium squamosum, f. Hordeum secalinum, f. Bromus hordea- ceus s. str., f. Tetragonolobus maritimus); and the var. Melilotus siculus, a grouping that is more salinised and floristically more impoverished (Mercadal 2019c: Ta- ble 147, rel. 10-11). 1.1.2 Ass. Agropyro-Trifolietum subass. festucetosum arundinaceae (Molinier et Tallon 1968) Mercadal comb. nova hoc loco [cf. Mercadal 2019c: 608] Basionym: Arrhenatheretum elatioris Braun 1915 sub- ass. festucetosum arundinaceae Molinier et Tallon 1968 [sub Arrhénathéraie sous-association à Festuca arun- dinacea = Arrhenatheretum Molinier & Tallon 1968 nom. illeg. (Art. 31) subass. festucetosum Molinier et Tallon 1968]. Synonyms: Gaudinio-Arrhenatheretum Br.-Bl. in Br.-Bl. et al. 1952 subass. festucetosum (Moli- nier et Tallon 1968) Gesti 2000 p. max. p. comb. ined. (Art. 1) [sub Gaudinio-Arrhenatheretum Br.-Bl. 1931 subass. festucetosum Molinier et Tallon 1970]; Gaudi- nio-Arrhenatheretum Br.-Bl. in Br.-Bl. et al. 1952 su- bass. festucetosum (Molinier et Tallon 1968) Gesti 2006 p. max. p. comb. inval. (Art. 3i) [sub Gaudinio-Arrhe- natheretum Br.-Bl. 1931 subass. festucetosum Molinier et Tallon 1970]; Agropyro-Trifolietum Br.-Bl. in Br.-Bl. et al. 1952 subass. festucetosum arundinaceae (Molinier et Tallon 1968) Mercadal 2019 comb. ined. (Art. 1) [sub Agropyro-Trifolietum Br.-Bl. ex Br.-Bl. et al. 1952 subass. festucetosum arundinaceae (Molinier et Tallon 1970) Mercadal 2019]. Corresponding names: Lino biennis-Festucetum arundinaceae Dubuis et Simonneau ex Julve 1993 prov. nom. inval. (Art. 3b); Lino bien- nis-Festucetum arundinaceae Dubuis et Simonneau ex de Foucault in de Foucault et Catteau 2012; Dorycnio gracilis-Festucetum arundinaceae (Molinier et Tallon 1968) de Foucault in de Foucault et Catteau 2012 [sub Dorycnio gracilis-Festucetum arundinaceae (Molin. et Devaux 1978) de Foucault in de Foucault et Catteau 2012 nom. illeg. (Art. 31)]. Other denominations: “Prairies à Festuca arundinacea” (Molinier & Tallon 1968); “Groupement à Festuca arundinacea Schreb.” (Dubuis & Simonneau 1968). Lectotypus: Molinier & Tallon (1968), Table VI, rel. 2 (p. unnumbered) [sub Arrhénathéraie sous-associa- tion à Festuca arundinacea]; designated in Foucault et Catteau (2012: 67) [sub Dorycnio–Festucetum (Molin. et Devaux 1978) de Foucault in de Foucault et Cat- teau 2012]. Physiognomy and floristic composition: subhalo- philic and mesohygrophilous meadow, 40 to 120 cm in height, dominated by Festuca arundinacea and with an elevated presence of the association’s characteristic and differential species (Trifolium squamosum, Horde- um secalinum, Bromus commutatus, Vicia tetrasperma, V. lutea, Tragopogon lamottei, Sonchus asper, Oenanthe lachenalii, Bellevalia romana). The only taxa generally absent are the more exclusive ones, typical of more sa- line soils: Melilotus siculus, Medicago scutellata, Elymus athericus, Festuca interrupta, O. palustris, O ×lloydiana. The differential taxa of the subass. festucetosum (cf. diff. species) are, after our amendment, common plants in the order Trifolio-Hordeetalia that withstand brackish soils. Differential species : Poa trivialis subsp. trivialis, Lotus corniculatus subsp. corniculatus, Agrostis stolonifera, Holcus lanatus, Oenanthe fistulosa, Polypogon mon- speliensis, Ranunculus sardous s.l., Lychnis flos-cuculi, Leontodon taraxacoides subsp. taraxacoides. Ecology: soils that are slightly saline, clay and alkaline with a low carbonate content and often seasonally flooded (spring and autumn). During dry periods, the concentration of salt increases, which selects the plant population. For this reason, in the past, most mead- ows were intensively irrigated and fertilised (Dubuis & Simonneau 1968, Voelckel 1977). At present, the meadows are mown, grazed and sometimes remown, irrigated, fertilised and resown (Table 1). Geographical distribution: Camargue (Provence) and the plains of Roussillon and Empordà (Catalonia). Probably also it distributed in the wetlands of Montpel- lier and Narbonne. In the study area, it is quite wide- spread along the Ruscinic coast, from the sea level to an altitude of 16(85) m (Figure 1e & Mercadal 2019c: Fig. 282). Syntaxonomy: we attribute to this new combination the regularly mown subhalophilic and mesohygrophi- 20/1 • 2021, 107–117 114 Gabriel Mercadal i Corominas Geobotanical characterisation of the Trifolion squamosi nom. corr. alliance (class. Juncetea maritimi) in Catalonia and the Western Mediterranean lous meadows dominated by Festuca arundinacea from the Catalan-Provençal coast (Molinier & Tallon 1968, 1970, Dubuis & Simonneau 1968, Molinier & Deveaux 1978, Voelckel 1977, Watt & Vilar 1997, Gesti 2006, Mercadal 2019c). The syntaxon was val- idly described by Molinier & Tallon (1968: Table VI) as "Arrénathéraie sous-association à Festuca arundina- cea", since on page 445 it is clear that the French name "Arrénathéraie" corresponds explicitly to the associa- tion Arrhenatheretum Molinier & Tallon 1968; an il- legitimate homonym (Art. 31) of the Arrhenatheretum Braun 1915, but which also allows to validate the sub- association festucetosum Molinier & Tallon 1968 (Art. 30b). Molinier & Tallon (1968) published in Table VI 10 relevés, but in that case the type was not indicated. Later, after receiving other names (cf. synonyms), Fou- cault & Catteau (2012) characterized two associations: one to include the Camargue relevés by Molinier & T al- lon (1968) [Dorycnio-Festucetum], and another for the Roussillon relevés published by Dubuis & Simonneau (1968) [Lino-Festucetum]. Moreover, for each syntaxon, they chose a type. However, Foucault & Catteau (2012: 67) qualified Dorycnio-Festucetum as a new association, despite the fact that it corresponds to a change in rank (stat. novus). However, according to our results (Mercadal 2019c: Table 144 & Fig. 268), both syntaxa correspond to the subassociation festucetosum Molinier & Tallon 1968, which we transfer from Arrhenatheretum to Agropyro- Trifolietum, and we define several new differential species. Finally, it should be noted that Delbosc et al. (2015) cited the association Lino-Festucetum Dubuis et Simon- neau ex de Foucault in de Foucault et Catteau 2012 [= Agropyro-Trifolietum subass. festucetosum] from the Asco Valley, in northern Corsica. However, the Asco Valley is located at altitudes of between 200-2076 m, and the Agropyro-Trifolietum subass. festucetosum grows near the coast, at altitudes of between 0-85 m. Therefore, it is probably another different syntaxon of Agropyro- Trifolietum subass. festucetosum, and might correspond to a community in the Oenantho-Gaudinion alliance. Nevertheless, Agropyro-Trifolietum could grow on the coasts of Corsica. Variability: in Catalonia, we can distinguish different fa- cies in accordance with the dominant plant that accom- panies Festuca arundinacea, such as: f. Bromus commu- tatus (brackish and very wet areas), f. Bromus hordeaceus (areas that are disturbed, nitrophilous, often grazed or fertilised) or f. Hordeum secalinum (more saline areas, which are often flooded naturally; this is the form clos- est to the subass. typicum). 1.1.3 Ass. Agropyro-Trifolietum subass. brachypodieto- sum phoenicoidis subass. nova hoc loco [cf. Mer- cadal 2019c: 627] Synonyms: Gaudinio-Arrhenatheretum Br.-Bl. in Br.-Bl. et al. 1952 subass. festucetosum (Molinier et Tallon 1968) Gesti 2000 p. min. p. comb. ined. (Art. 1); Gaudinio-Arrhenatheretum Br.-Bl. in Br.-Bl. et al. 1952 subass. festucetosum (Molinier et Tallon 1968) Gesti 2006 p. min. p. comb. inval. (Art. 3i); Agropyro-Trifo- lietum Br.-Bl. in Br.-Bl. et al. 1952 subass. brachypodie- tosum phoenicoidis Mercadal 2019 nom. ined. (Art. 1). Holotypus hoc loco designatus: Mercadal (2019c), unpublished PhD thesis, Table 145, rel. 1, p. 629 (Spain, Catalonia, Alt Empordà, Sant Pere Pescador, 2 m, 31TEG0871, 28.04.2004). Plot size, 100 m 2 ; height, 45 cm; total cover, 100%: Tetragonolobus matirimus +, Tragopogon lamottei +, Vicia tetrasperma subsp. gracilis +, Brachypodium phoenicoides 2.2, Av- ena barbata +, Inula viscosa 1.1, Ophrys apifera subsp. apifera +, O. scolopax subsp. scolopax +, Anacamptis py- ramidalis +, Barlia robertiana +, Elymus campestre 2.2, Centaurium pulchellum subsp. tenuiflorum +, Festuca arundinacea subsp. arundinacea 3.4, Dactylis glomerata subsp. glomerata +, D. glomerata subsp. hispanica +, Lathyrus hirsutus +, Linum usitatissimum subsp. angus- tifolium 1.2, Trifolium pratense 2.2, T. dubium +, T. campestre +, Daucus carota subsp. carota 1.2, Equisetum ramosissimum subsp. ramosissimum 1.1, Hypochaeris radicata +, Plantago lanceolata +, Geranium dissectum +, Bromus hordeaceus subsp. hordeaceus +, Medicago sativa subsp. sativa 1.2, M. polymorpha subsp. polymorpha +, Poa pratensis subsp. pratensis 1.1, P. bulbosa +, Phrag- mites australis +, Aster pilosus +, Sherardia arvensis +, Bellis perennis 1.1, Gaudinia fragilis +, G a l i um v e r um subsp. verum 1.1, Crepis vesicaria subsp. ta r a x a ci f o l i a +, Myosotis ramosissima subsp. ramosissima +, Orobanche minor +, Ranunculus bulbosus +, Aristolochia rotunda +, Bellardia trixago +, Reichardia picroides +, Serapias lin- gua +, Valerianella microcarpa +, Schoenus nigricans +, Carex flacca 1.2, Cerastium glomeratum +, Scabiosa at- ropurpurea 1.1, Urospermum delachampii +. Physiognomy and floristic composition: subhalo- philic meadow, mesohygrophilous to mesophilic, 45 to 120 cm in height, characterised floristically by the low presence of plants related to the association and the upper units. The only species belonging to the association that appear regularly are: Tragopogon lamottei (diff.), Vicia tetrasperma subsp. gracilis, Trifo- lium squamosum and Hordeum secalinum. The diffe- rentials of the subassociation are herbaceous plants from mesophilic meadows or relatively disturbed areas and some calcicolous orchids (cf. diff. species). 20/1 • 2021, 107–117 115 Gabriel Mercadal i Corominas Geobotanical characterisation of the Trifolion squamosi nom. corr. alliance (class. Juncetea maritimi) in Catalonia and the Western Mediterranean Plants typical of Brachypodion phoenicoidis are typical there, since the subass. brachypodietosum comprises a transition towards Brachypodietalia phoenicoidis. Thus, apart from Brachypodium phoenicoides, the following plants are more or less common: Elymus campestris, Urospermum delachampii, Picris hieracioides subsp. hieracioides, Foeniculum vulgare subsp. piperitum and Scabiosa atropurpurea (Mercadal 2019c: Table 155). Differential species: Brachypodium phoenicoides, Avena barbata, Inula viscosa, Hypochaeris radicata, Rubus ul- mifolius, Cirsium vulgare, Ophrys apifera subsp. apifera, Orobanche crenata, Anacamptis pyramidalis, Barlia rob- ertiana. Ecology: land only slightly saline during dry periods, with a sandy-clay texture, alkaline, with a high carbon- ate content and rarely flooded. The meadows are essen - tially only mowed and/or grazed, whilst other agricul- tural activities are rare (Table 1). Geographical distribution: Empordà plain, between 0–4 (16 m) of altitude (Figure 1f & Mercadal 2019c: Fig. 289). 1.2 Ass. Festucetum arundinaceae Simonneau 1952 [cf. Mercadal 2019c: 578] Synonym: Ass. à Festuca arundinaceae Simonneau 1952 (orig. form) [sub Ass. à Festuca arundinacea Simonneau 1950]. Lectotypus hoc loco designatus: Simonneau (1952), Table 23, rel. VI (p. unnumbered). Physiognomy and floristic composition: mesohy- grophilous meadow 100 cm in height clearly domi- nated by Festuca arundinacea, which is usually accom- panied by Juncus maritimus, Trifolium resupinatum, T. squamosum, Melilotus gr. sulcatus (probably M. segeta- lis), Hordeum maritimum, Polypogon monspeliensis and Bolboschoenus maritimus). Floristically, the association is poor (11 taxa/rel. on average) and contains a high percentage of therophytes (71%). All the taxa are Plu- riregional (71%) or Mediterranean (29%) (Mercadal 2019c: Table 138 & Fig. 238). Characteristic and differential species : Trifolium resupi- natum, Medicago intertexta subsp. ciliaris, Centaurium spicatum, Leontodon marrocanus (diff.). Ecology: subsaline soils (sodium-calcareous solonchaks containing 2 to 4 g/kg of salts in the surface horizon) with a silty-clay texture, intensely irrigated, sown, fer- tilised, mowed and grazed (Simonneau 1952, Dubuis & Simonneau 1957). Geographical distribution: in 1950, it occupied several thousand hectares on the Maca plain, near Oran, in northern Algeria (Simonneau 1952). It is probably dis- tributed in other regions of North Africa. Syntaxonomy: Simonneau (1952) published 10 relevés of the association, but he did not indicate the nomen- clature type. Therefore, we specify it in this article. Moreover, the intense agricultural activity makes its phytosociological attribution difficult. The presence of Trifolium squamosum, Melilotus siculus and several halophytes, as well as the ecology of the association, leads us to attribute Festucetum to Trifolion squamosi. In any case, new relevés need to be made, particularly in less heavily worked meadows, to find out how vari- able these are and to enable us to attribute this syn- taxon accurately. Syntaxonomic schema Class. Juncetea maritimi Br.-Bl. in Br.-Bl., Roussine et Nègre 1952 Ord. Juncetalia maritimi Br.-Bl. ex Horvatić 1934 All. Trifolion squamosi Br.-Bl. in Br.-Bl., Roussine et Nègre 1952 nom. corr. [#1] Ass. Agropyro pycnanthi-Trifolietum maritimi Br.-Bl. in Br.-Bl., Roussine et Nègre 1952 nom. inept. [#1.1] subass. typicum [#1.1.1] subass. festucetosum arundinaceae (Moli- nier et Tallon 1968) Mercadal comb. nova hoc loco [#1.1.2] subass. brachypodietosum phoenicoidis sub- ass. nova hoc loco [#1.1.3] Ass. Festucetum arundinaceae Simonneau 1952 [#1.2] Other syntaxa quoted in the text Agrostietalia castellanae Rivas Goday in Rivas-Mart., Costa, Castroviejo et E. Valdés 1980; Agrostio stoloniferae- Arrhenatheretea elatioris (Tüxen 1937 em. 1970) de Fou- cault 1984 nom. ined. (Art. 1) [= Molinio-Arrhenathere- tea]; Agrostion castellanae Rivas Goday 1958 corr. Rivas Goday et Rivas-Mart. 1963; Agrostion salmanticae Rivas Goday 1958; Alopecurion utriculati Zeidler 1954 [= Mo- linio-Hordeion secalini Horvatić 1934]; Brachypodieta- lia phoenicoidis Br.-Bl. ex Molinier 1934; Brachypodion phoenicoidis Br.-Bl. ex Molinier 1934; Eleocharietalia palustris de Foucault 1984 nom. ined. (Art. 1) [= Des- champsietalia cespitosae Horvatić 1958]; Gaudinio fragi- lis-Arrhenatheretum elatioris Br.-Bl. in Br.-Bl. et al. 1952 subass. lotetosum preslii Donker et Stevelink ex Mercadal 2020; Geranio dissecti-Festucetum arundinaceae (O. Bolòs 1959) Mercadal 2020 subass. caricetosum distantis Mer- cadal 2020; Isoëto-Nanojuncetea Br.-Bl. et Tx. in Br.-Bl. et al. 1952; Kickxio commutatae-Trifolietum bocconei Brullo 20/1 • 2021, 107–117 116 Gabriel Mercadal i Corominas Geobotanical characterisation of the Trifolion squamosi nom. corr. alliance (class. Juncetea maritimi) in Catalonia and the Western Mediterranean et Marcenò 1985; Lino biennis-Gaudinietum fragilis Fran- quesa 1995; Molinio-Arrhenatheretea Tx. 1937; Oenantho pimpinelloidis-Gaudinion fragilis Mercadal 2020; Poten- tillo-Polygonetalia avicularis Tx. 1947; Ranunculo ophio- glossifolii-Oenanthion fistulosae de Foucault in de Foucault et Catteau 2012; Stipo giganteae-Agrostietea castellanae Rivas-Mart., Fernández-González et Loidi 1999; Trifo- lietum nigrescenti-resupinati Molinier et Tallon 1968; Tri- folio frafigeri-Hordeetalia secalini Horvatić 1963; Trifolio fragiferi-Cynodontion dactylonis Br.-Bl. et O. Bolòs 1958; Tuberarietea guttatae Rivas Goday et Rivas-Mart. 1963. Acknowledgments To Dr. Jean-Paul Theurillat, professor of the Department of Botany and Plant Biology of the University of Geneva, for reviewing the text. To Dr. Enrique Rico, professor of the Department of Botany of the University of Salamanca, for the nomenclatural clarifications on the genus Elymus. 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