SYNTAXONOMIC NOVELTIES ON SALT MARSHES FROM WEST

IBERIAN PENINSULA

José Carlos Costa*, Carlos Neto**, Tiago Monteiro-Henriques*, Pedro Arsénio*, Estevão Portela-Pereira**, Ana Caperta* & Jesus Izco***

* Instituto Superior de Agronomia, Technical University of Lisbon (TU Lisbon)** Institute of Geography & Spatial Planning (IGOT), University of Lisbon***Departamento de Botánica, University of Santiago de Compostela

The frontier between the Mediterranean and the Eurosiberian regions converge to the Portuguese northwest coastline. The biogeographical coastal terrestrial borderline was established on the Ria de Aveiro. Regarding salt marshes particularly, it is difficult to establish this limit sharply, since an evident physical barrier is lacking

European classification of salt marshes from Géhu & Rivas-Martínez (1983)

• Costa et al. (2009) corroborate the Géhu and Rivas-Martínez (1983) suggestion: the authors proposed the Tagus River salt marsh as the boundary between the Eurosiberian and the Mediterranean regions. • As for the Iberian Peninsula, they considered the sub-zone Cantabrian-Atlantic belonging to the Eurosiberian territory, and two other sub-zones (Mediterranean-Atlantic and Mediterranean) separated by the Straits of Gibraltar, although using only community data, confirming that salt marshes eastward and westward are distinct.

Salt marsh physiognomy differs among tropical, mediterranean and temperate climates.

In temperate climates vegetation is generally short, such as rushlands or halophytic grasslands (hemicryptophytes dominate, along with a few chamaephytes)

Mediterranean and desert tropical territories shrubs dominate (vegetation types are essentially nanophanerophytes, microphanerophytes, accompanied by chamaephytes and hemicryptophytes)

Limonio vulgare-Juncetum subulati

[Arthrocnemo glauci-Juncetum subulati sensu RIVAS-MARTÍNEZ et al. (1980), COSTA et al. (1996) non Arthrocnemo glauci-Juncetum subulati Brulo & Funari 1976]

Dominated by Juncus subulatus, in depression or abandoned saltpans, flooded for a more or less long period by brackish water derived from autumn and winter rainfall, completely dried out during summer.

Saline-clayey soils, with a high sandy fraction, being the Iberian Arthrocnemion macrostachyi community that bear the longest period of flood.

Characteristic floristic composition: Juncus subulatus, Halimione portulacoides, Sarcocornia fruticosa, Limonium vulgare, Juncus maritimus, Aster tripolium subsp. pannonicus, Arthrocnemum macrostachyum, Sarcocornia perennis subsp. alpini, Polygonum equisetiforme, Myriolimon ferulaceum, Inula crithmoides, Elytrigia elongata, Triglochin barrelieri, etc.

Table 1 - Limonio vulgaris-Juncetum subulati No. of relevé 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Minimum surface (m2) 2 2 2 2 2 2 4 4 20 4 4 2 20 No. of species/ no. of relevés 3 5 5 6 4 6 7 11 14 8 9 7 12 9 9 Characteristics Juncus subulatus 5 4 3 4 4 5 3 4 5 4 4 3 4 V V Halimione portulacoides . . . + + 1 2 3 1 2 2 2 + . III Sarcocornia fruticosa 1 1 1 . . + . 1 + 2 2 1 . . II Juncus maritimus . . . . . 1 1 + 2 + 3 + 1 . III Limonium vulgare . . + . . . . 1 1 + + 1 1 . II Arthrocnemum macrostachyum . + . 1 2 . . + + . 1 . 1 V V Aster tripolium subsp. pannonicus . . 2 . . . . . 1 3 3 2 1 . II Sarcocornia alpini 1 . . 2 + . . . . . . . . . II Polygonum equisetiforme . . . . . . 2 1 + . . . + . II Inula crithmoides . . . . . . . + 1 . . . + . II Myriolimon ferulaceum . . . . . . + 2 . . . . . . II Elytrigia elongata . + . . . . . . . + + . . . . Triglochin barrelieri . . . . . . . . + . . . + . II Suaeda vera . . . . . + . . . . . . . . III Companions Atriplex hastata . . . . . . . . + 1 1 . + . II Parapholis filiformis . . . + . . 2 1 . . . . . . II Polypogon maritimus . . . + . . . . + . . . + . III Bolboschoenus compactus . + . . . . . . . . . . . II II Salicornia patula . . 2 . . . . . . . . 3 . . . Hordeum marinum . . . . . . . . . . . . . II I Phragmites australis . . . . . + . . + . . . . . . Plantago coronopus . . . . . . + + . . . . . . . More: 2 Spartina patens, +Artemisia caerulescens in 13; II Scirpus littoralis, I Ruppia maritima subsp. drepanensis, I Cotula coronopifolia, I Salicornia ramosissima, I Salsola soda in 14; III Limonium algarvense, II Limoniastrum monopetalum, II Myriolimon diffusum, I Cistanche phelypaea, II Salsola soda, II Frankenia laevis, II Salsola soda, I Spergularia media in 15. Places: 1, 4 Vau Salines, Tagus River (Alcochete); 2, S. Francisco, Tagus River (Alcochete); 3, 11 Museu do Sal, Tagus River (Alcochete); 6 Ponta da Erva, Tagus River; 6, 7, 8, 9,13 Hortas, Tagus River (Alcochete); 10, 12 Samouco, Tagus River (Alcochete); 15 RIVAS-MARTÍNEZ et al. (1980) with de name Arthrocnemo glauci-Juncetum subulati; 16 COSTA et al. (1996) with the name Arthrocnemo glauci-Juncetum subulati juncetosum subulati

Limonium vulgare, Myriolimon ferulaceum, Frankenia laevis are differential species of this new community distinguishing it from Arthrocnemo glauci-Juncetum subulati and Elymo elongati-Juncetum subulati.

Of difficult syntaxonomic positioning, as it contains plant species both form Sarcocornietea fruticosae and from Juncetea maritimi, we decided to assign it to the former, Arthrocnemion macrostachyi, Arthrocnemenion macrostachyi, as the characteristic species of Sarcocornietea are more frequent and because these communities occupy the higher parts of the salt marshes resisting to high salinity contents during summer.

Limonio vulgare-Juncetum subulati

Thermomediterranean dry bioclimate

Within the Tagus and the Guadalquivir rivers (Coastal Lusitanian-Andalusian Province, Mediterranean Region)

Triglochino maritimae-Sarcocornietum alpini

Halophytic community dominated by Sarcocornia perennis subsp. alpini, Triglochin maritima and Halimione portulacoides occurring in salt marshes in areas north of the Tagus River, in sandy-silty zones, occasionally flooded by brackish water, in mesomediterranean bioclimate.

Geovicarious of Halimiono portulacoidis-Sarcocornietum alpini, which belongs to the southeast of the Iberian Peninsula, south of the Tagus Estuary, in thermomediterranean territories. Triglochin maritima and Aster tripolium subsp. pannonicus are the differentials, on the other hand, the Mediterranean species Puccinellia iberica, Cistanche phelypaeae and Artemisia gallica are absent from the floristic composition.

Contacts at lower elevations with Puccinellio maritimae-Sarcocornietum perennis and with Limonio-Juncetum maritimi at higher quotas.

Table 2 - Triglochino martimae-Sarcocornietum alpini No. of relevé 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Minimum surface (m2) 2 6 2 4 4 2 4 1 2 4 4 6 15 6 6 No. of species 4 6 3 7 5 4 4 10 6 7 8 8 7 7 9 Characteristics Sarcocornia alpini 2 3 3 2 3 3 3 2 4 4 2 2 5 2 2 Halimione portulacoides 5 3 4 4 3 3 2 2 1 2 4 + + . . Triglochin maritima . . 2 1 + 2 1 2 2 2 1 2 1 1 2 Juncus maritimus 2 . . 1 . . . 1 2 . 2 1 + 1 + Puccinellia maritima (dif. subass.) . . . . . . . 3 1 + 2 2 + 1 2 Plantago maritima (dif. subass.) . . . . . . . . . . . 2 1 1 + Spergularia media . + . + . . + + . . . 1 . 1 2 Aster tripolium subsp. pannonicus . + . . + . . . . + 2 + . . 2 Inula crithmoides . + . + . 1 . 1 . . + . . . . Limonium vulgare . . . + . . . 1 + . 3 . + . . Sarcocornia fruticosa . . . . . . . 1 . 1 . . . . . Sarcocornia perennis . . . . . . . . . . . . . 1 . Companions Spartina maritima 1 . . . . . . . . + . . . . . Salicornia ramosissima . + . . . . . 1 . . . . . . . More: +Puccinellia tenuifolia in 4, 1Cotula coronopifolia, +Bolboschoenus compactus in 15 Places: 1, 3, 4, 5 Barrosa do Cal River (Óbidos Lagoon, Caldas da Rainha); 2 Mondego River (Figueira da Foz); 6, 7 Salir do Porto, (Tornada River, Caldas da Rainha); 8 Vagueira (Ria de Aveiro); 9 Torreira (Ria de Aveiro); 10 Murtosa (Ria de Aveiro), 11 Costa Nova (Ria de Aveiro); 12 Ofir (Cavado River), 13 Currubedo (Galicia); 14, 15 Viana do Castelo (Lima River)

sarcocornietosum alpini is typical

puccinellietosum maritimae J.C. Costa, Neto & Izco subass. nova hoc loco with Puccinellia maritima and Plantago maritima as differential species

sarcocornietosum alpini Portuguese Divisory Sector (Coastal Lusitanian-Andalusian Province, Mediterranean Region) between Óbidos Lagoon and Mondego River.

puccinellietosum maritimae north of the Ria de Aveiro reaching Galicia in Northern Galician-Portuguese Subsector (Cantabrian-Atlantic Subprovince, Atlantic European Province, Eurosiberian Region),

Triglochino maritimae-Sarcocornietum alpini

Puccinellio tenuifoliae-Limonietum daveaui Chamaephytic association in sandy-brackish soils, only occasionally touched by brackish water during equinoctial tides, although occasionally flooded by fresh water during winter or springtime.

Limonium daveaui, Puccinellia tenuifolia, Halimione portulacoides, Frankenia laevis, Juncus maritimus, Myriolimon ferulaceum are the most reliable species

This association stands between Juncetea maritimi and Sarcocornietea fruticosae;.

Table 3 -Puccinellio tenuifoliae-Limonietum daveaui No. of relevé 1 2 3 4 5 6 7 8 9 10 11 12 13 Minimum surface (m2) 1 2 3 4 4 2 4 2 2 2 4 1 1 No. of species 6 4 5 8 9 11 10 7 10 6 9 4 5 Characteristics Limonium daveaui 3 4 2 3 3 3 3 3 4 4 3 3 3 Puccinellia tenuifolia 1 3 2 1 1 1 1 1 + + . . . Halimione portulacoides . 3 2 1 + 2 1 3 . . + 1 1 Frankenia laevis 2 . . 2 1 2 3 . . 2 1 3 2 Myriolimon ferulaceum 2 2 4 1 3 + . . . 1 . . . Juncus maritimus . . . . + . 2 2 2 . 3 2 2 Artemisia gallica . . . . . + 1 + 1 1 1 . . Juncus acutus . . . . 1 . 1 1 + . 2 . . Inula crithmoides 1 . . 3 1 + . . . . . . . Aster tripolium subsp. pannonicus . . . . . + . . + . . . . Spergularia salina . . . 1 . . . . . . . . . Companions Plantago coronopus . . + + . + + . . + . . + Carpobrotus edulis . . . . . . 1 1 . . 1 . . Polypogon maritimus . . . . . + . 1 2 . . . . Hordeum marinum . . . . . + . + 1 . . . . Gaudinia fragilis . . . . . . + . 2 . . . . Dittrichia viscosa . . . . . . + . . . 1 . . More: 2Elytrigia juncea subsp. boreali-atlantica in 1; 1Paspalum distichum in 4; 1Torilis arvensis in 9; +Aetheorhiza bulbosa in 11

Places: 1-6 Salir do Porto (Tornada River), 6 Alcochete (Tagus River), 7-13 Coina (Tagus River)

between the Tagus River and S. Martinho do Porto (Tornada River)Divisory Portuguese-Sadensean Subprovince (Coastal Lusitanian-Andalusian Province, Mediterranean Region) Thermomediterranean subhumid bioclimate

Puccinellio tenuifoliae-Limonietum daveaui

Puccinellio tenuifoliae-Limonietum daveaui

This association stands between Juncetea maritimi and Sarcocornietea fruticosae; however, we propose its inclusion in the Limonio ovalifolii-Frankenio laevis alliance, from low littoral Cantabrian-Atlantic order Glauco-Puccinellietalia, due to the presence of Puccinellia tenuifolia, Frankenia laevis, Juncus maritimus, Juncus acutus and Aster tripolium subsp. pannonicus. This alliance corresponds to the coastal prostate chamaephytic halo-nitrophilous communities reached by the highest tides.

Myriolimetum ferulacei Rothmaler 1943 nom. mut. [bas. Limonietum ferulacei Rothmaler 1943, sin. Inulo crithmoidis-Limonietum ferulacei Rivas-Martínez, Costa, Castroviejo & E. Valdés 1980]

Dominated by the chamaephyte Myriolimon ferulaceum (=Limonium ferulaceum) accompanied by Inula crithmoides, Halimione portulacoides, Frankenia laevis, Puccinellia iberica, Artemisia gallica, etc. and Limoniastrum monopetalum in the Algarve

In clayey or silty soils from salt marshes only inundated by the equinoctial tides or from calcareous and marly sea cliffs heavily sprinkled by the Atlantic Ocean waters during winter storms.

It is included in the Limonion confusi alliance, Limonietalia order, due to its ecology .

Table 4 - Myriolimetum ferulacei No. of relevé 1 2 3 4 5 6 7 8 9 10 11 12 13 Minimum surface (m2) 4 2 8 1 2 4 4 4 6 2 No. of species / nº of relevées 6 4 12 5 5 8 7 8 9 7 11 9 12 Caractheristics Myriolimon ferulaceum 3 4 3 3 2 3 4 4 3 5 IV V V Inula crithmoides . . . . 1 2 1 1 3 . I V III Halimione portulacoides 3 2 1 4 4 . . . . . . V IV Puccinellia iberica . 1 . 2 + . . . . . . . + Artemisia gallica 3 + . . . . . . . . . I . Limonium vulgare . . + . . . . . . . . I . Limoniastrum monopetalum . . . . . . . . . . . III III Triglochin barrelieri . . . . . . . . . . . III + Suaeda vera . . . . . . . . . . I . + Limonium algarvense . . . . . . . . . . . . II Companions Frankenia laevis . . . 1 . 2 2 1 2 1 V . IV Plantago coronopus . . 1 . . 1 1 1 1 + V . + Beta maritima . . . . . 1 1 1 + + + . I Parapholis incurva . . . . . . + + . 1 II . II Dactylis marina . . . . . + 1 . + + + . . Polygonum equisetiforme . . 3 . . . . . . . I I + Parapholis filiformis . . . 1 . . . . . . III . IV Hordeum marinum . . + . . . . . . . I . II Crithmum maritimum . . . . . + . . + . I Juncus maritimus . . . . . . . . . . + I I Mesembryanthemum nodiflorum . . . . . 2 . . . . II . . Cynodon dactylon . . 1 . . . . . . . II . . Aster tripolium subsp. pannonicus . . + . 1 . . . . . . . . Polypogon marinum . . + . . . . . . . . . III Eryngium dilatatum . . . . . . . + + . . . . Sonchus oleraceus + . . . . . . . + . . . More: 2Bellis annua, +Senecio vulgaris in 1; 1Gaudinia fragilis, +Anacyclus radiatus, +Aster squamatus in 3; +Centaurium spicatum in 8; +Spergularia australis in 10; IV Limonium ovalifolium, II Limonium echioides, I Salsola vermiculata, I Catapodium marinum in 11; II Spartina densiflora, II Juncus subulatus in 12; III Spergularia salina, II Spergularia bocconei, I Beta macrocarpa, I Juncus acutus in 13 Places: 1, 2, 4 Coina (Tagus River); 3 Hortas (Alcochete, Tagus River), 5 Samouco (Alcochete, Tagus River), 6 Ericeira (Mafra); 7, 9 Ribamar (Mafra); 8 Sea cliff of Praia Azul (Torres Vedras); 9 Baleal (Peniche); 10 Cambelas (Torres Vedras); 11 ROTHMALER (1943); 12 RIVAS MARTÍNEZ et al. (1980); 13 COSTA (1991)

Thermomediterranean dry bioclimate

Coastal Lusitanian-Andalusian Province.

Limonietum lanceolati

Hemicryptophytic community, dominated by Limonium lanceolatum and Myriolimon ferulaceum accompanied by several plants of Sarcocornietea fruticosae

In clayey or silty soils rich in schistose debris, only inundated during the highest tides.

It occupies places relatively higher than the Inulo crithmoidis-Arthrocnemetum glauci and the Cistancho phelypaeae-Suadetum verae

Table 4 – Limonietum lanceolati No. of relevé 1 2 3 4 5 6 7 Minimum surface (m2) 4 6 2 6 4 4 No. of species / nº of relevés 8 9 7 10 7 4 6

Characteristics Limonium lanceolatum 3 2 4 2 3 3 V Halimione portulacoides 1 1 + 1 2 2 V Myriolimon ferulaceum 1 2 1 2 + . V Arthrocnemum macrostachyum . . + 1 1 1 V Artemisia gallica . 1 + 1 + . III Inula crithmoides + + . + 1 . II Suaeda vera . + + + . . I Puccinellia iberica 1 . . + . . III Companions Suaeda albescens + + . . . 1 II Spergularia marina . + . + . . II Plantago coronopus + . + . . . I Juncus maritimus . . . 1 1 . I Frankenia laevis + . . . . . I Salsola vermiculata . + . . . . I More characteristics: II Limonium vulgare, II Cistanche phelypaea in 6 Places: 1, 2 Vale D. Sancho (Cerca River, Aljezur); 3 Vila Nova de Milfontes near the bridge (Odemira); 4, 5 South of Monte do Zambujeiro (left bank Mira River); 6 - Monte do Carriçal (right bank Mira River); 7 NETO et al. (2009)

Limonietum lanceolati

Mira River and Cerca River (Aljezur), Coastal Vincentine District (Coastal Lusitanian-Andalusian Province)

Thermomediterrean dry

Limonio algarvensis-Myriolimetum diffusi J.C. Costa, Lousã, Caperta & Neto ass. nova hoc loco

Communities dominated by the chamaephytes Myriolimon diffusum (Pourr.) Lledó, Erben & Crespo (= Limonium diffusum (Pourr.) O. Kunze), Limonium algarvense and Limonium latebracteatum found in the Algarve

Limonium latebracteatum was found for the first time in Portugal during this research.

This phytocoenosis occurs in sandy well-drained soils, inundated only during the highest tides, under thermomediterranean dry bioclimate.

Table 6 - Limonio algarvensis-Myriolimetum diffusi No. of relevé 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 Minimum surface (m2) 4 4 4 6 4 6 2 4 6 8 8 4 4 2 2 6 4 2 2 6 No. of species 6 6 4 7 5 8 7 7 6 9 7 9 7 7 5 4 6 7 6 6 Characteristics Myriolimon diffusum 3 2 2 2 2 3 3 3 1 2 1 1 1 1 1 + 3 + 3 . Limonium algarvense . 3 3 + 1 + 1 2 2 2 2 2 1 2 2 2 2 2 1 1 Limonium latebracteatum 1 1 + 1 1 + 1 1 1 + 1 1 + 1 + + 2 . . 1 Limoniastrum monopetalum . . . . . + + 1 + + + + + . . . + + . . Suaeda vera . + + + . . + . . + . + + . + . . + + . Puccinellia iberica . . . . . . . + . 1 . + . + . . + . . . Arthrocnemum macrostachyum . . . . . . . + + . . + . + . . . . . . Myriolimon ferulaceum . . . . . . . . . + . . . . . . . . + . Polygonum equisetiforme . . . . . . . . . . + . . . . . . . . + Inula crithmoides . . . . . . . . . . . + . . . . . . . . Companions Frankenia laevis + 1 . + . + . . . . + . + . . + . . + + Suaeda albescens + . . . + 1 . . 1 . . . 1 + . . 1 . + Salsola vermiculata + + . + + . + . . . + . + . . . . + . + Sporobolus pungens 1 . . . . . . + 2 . . 1 . . . . . 2 1 . Parapholis filiformis . . . . . + . . . . . . . . . . + . . . More: +Elymus elongatus in 6, +Spergularia salina in 10, +Polygonum maritimum in 20 Places: 1, 2, 3, 5, 7, 11, 16 Forte do Rato (Tavira), 4 Cocos Islet (Olhão), 6 Armona Island (Olhão), 8, 9, 12, 14, 15 Torre de Bias (Olhão), 10 near Faro bridge, 13, 18, 20 between Manta Rota e Cacela (Vila Real de Stº António), 17 Tavira Island, 19 Torre de Aires (Tavira),

Occupies places relatively lower than the occupied by the Polygono equisetiformis-Limoniastretum monopetali association, usually between this and Cistancho phelypaeae-Suaedetum verae

Although it resembles the Myriolimetum ferulacei association, its ecology is quite distinct as the latter occurs in clayey or silty soils both in salt marshes and in sea cliffs

Limonio algarvensis-Myriolimetum diffusi

We propose the inclusion of this association in the Limonion confusi alliance, Limonietalia order due their ecology and floristic composition.

Limonio algarvensis-Myriolimetum diffusi

Its area of distribution, so far, is restricted to the salt marshes of the Algarve (Algarvian District, Coastal Lusitanian-Andalusian Province)

SYNTAXONOMIC SCHEMESARCOCORNIETEA FRUTICOSAE Br.-Bl. & Tüxen ex A. & O. Bolòs 1950 nom. mut.Sarcocornietalia fruticosae Br.-Bl. 1933 nom. mut.Arthrocnemion macrostachyi Rivas-Martínez & Costa 1984 nom. mut.Arthrocnemenion macrostachyiLimonio vulgaris-Juncetum subulati J.C. Costa, Neto, T. Almeida & Lousã in J.C. Costa, Arsénio, Monteiro-Henriques, Neto, E. Pereira, T. Almeida & Izco 2009Sarcocornienion alpini Rivas-Martínez, Lousã, T.E. Díaz, Fernández-González & J.C. Costa 1990Triglochino maritimae-Sarcocornietum alpini J.C. Costa, Neto, Izco in J.C. Costa, Arsénio, Monteiro-Henriques, Neto, E. Pereira, T. Almeida & Izco 2009sarcocornietosum alpinipuccinellietosum maritimae J.C. Costa, Neto, Izco subass. nova hoc locoLimomnietalia Br.-Bl. & O. Bolòs 1958Limonion confusi Rivas-Martínez & Costa 1984Myriolimetum ferulacei Rothmaler 1943Limonietum lanceolati Arsénio, J.C. Costa, Neto & Monteiro-Henriques in Neto, Arsénio & J.C. Costa 2009Limonio algarvensis-Myriolimetum diffusi J.C. Costa, Lousã, Caperta & Neto ass. nova hoc locoJUNCETEA MARITIMI Br.-Bl. in Br.-Bl., Roussine & Négre 1952Glauco-Puccinellietalia Beeftink & Westhoff in Beeftink 1962Limonio ovalifolii-Frankenion laevis Arbesú, Bueno & F. Prieto 2002Puccinellio tenuifoliae-Limonietum daveaui J.C. Costa, Neto & E. Pereira in J.C. Costa, Arsénio, Monteiro-Henriques, Neto, E. Pereira, T. Almeida & Izco 2009