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Syntaxonomical review of theOmphalodion commutataealliance (class Helianthemetea)Francisco Gómez Mercado a , Esther Giménez Luquea & Francisco Valle Tendero ba Departamento de Biología Vegetal y Ecología,Escuela Politécnica Superior , Universidad deAlmería , E-04120 , Almería; E-mail:b Departamento de Biología Vegetal (Botánica),Facultad de Ciencias , Universidad de Granada ,Avda. Severo Ochoa s/n., E-18071 , Granada E-mail:Published online: 26 Apr 2013.

To cite this article: Francisco Gómez Mercado , Esther Giménez Luque & FranciscoValle Tendero (2006) Syntaxonomical review of the Omphalodion commutatae alliance(class Helianthemetea), Acta Botanica Gallica: Botany Letters, 153:3, 285-295, DOI:10.1080/12538078.2006.10515547

To link to this article: http://dx.doi.org/10.1080/12538078.2006.10515547

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Acta Bot. Gallica, 2006, 153 (3), 285-295.

Syntaxonomical review of the Omphalodion commutatae alliance (class H elianthemetea)

by Francisco Gomez Mercado(1), Esther Gimenez Luque(1) and Francisco Valle Tenderoe)

(1) Departamento de Biologia Vegetal y Ecologia, Escuela Politecnica Superior, Universidad de

Almeria, E-04120 Almeria; frgomez@ual.es, egimenez@ual.es

(2) Departamento de Biologia Vegetal (Botanica), Facultad de Ciencias, Universidad de Granada,

Avda. Severo Ochoa sin. E-18071 Granada ;.fvalle@ugr.es

arrive le 10 octobre 2005, accepte le 9 novembre 2005

Abstract.- The syntaxa of Omphalodion commutatae (Trachynietalia dista­chyae, Helianthemetea) are examined as regards to their floristic, ecological and biogeographical characteristics, and the new association Sileno lasiostylae­Arenarietum tenuis is described. A correspondence analysis (CA) has been car­ried out and, as a result of it, the relationships between the different syntaxa in the alliance are reviewed.

Key words : therophytic vegetation - mafic communities - correspondance ana­lysis - phytosociology - Spain.

Resume.- On dresse dans un premier temps un etat des caracteres flo ristiques, ecologiques et biogeographiques des syntaxons appartenant a !'alliance Ompha/odion commutatae (Trachynietalia distachyae, Helianthemetea) puis nous decrivons Ia nouvelle association Sileno lasiostylae-Arenarietum tenuis. Nous en analysons Ia d1sposition (CA) et, en nous basant sur les resultats des analyses, nous interpretons les rapports entre les divers syntaxon qui campo­sent !'alliance.

Mots cles : vegetation therophytique - communautes serpentinicoles - analyse de correspondances - phytosociologie - Espagne.

I. INTRODUCTION

The flora and plants growing on ultramafic substrates are very peculiar and present a high percentage of endemic taxa, something commonly attributed to the particular physical and chemical conditions. However, both the profile of these types of soils and the factors influencing the flora (Proctor & Nagy, 1992) may vary considerably from one place toano­ther (Chiarucci et al., 1998).

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In the Iberian Peninsula, this type of substrate occurs on a number of locations in the Betic ranges. Therophyte communities growing on these soils are included in the Omphalodion commutatae alliance (Trachynietalia distachyae, Helianthemetea), which has been extensively studied in the last decades and in which a number of subordinate syn­taxa have been described. Nowadays, there is a large bibliography dealing with its flora, ecology and distribution. The terminology used in the descriptions is heterogeneous and has inevitably caused a number of duplicities and nomenclature conflicts making analysis and global understanding of the topic even more difficult. Some of the surveys providing descriptive data are Rivas Goday & Esteve (1972), Rivas Martinez eta!. (1973), Izco (1976), Asensi et a!. (1979), Nieto et a!. (1988), Nieto et a!. (1989), Perez Raya et a!. ( 1991 ), etc. More recently, Izco and Fernandez-Gonza!ez (200 l) have recorded the history of the alliance and solved many of the previous disputes and nomenclature problems. After recording the information on the flora, ecology and biogeography of the syntaxa recogni­sed by these authors, we carried out the syntaxonomical analysis of the alliance. By means of a multivariate analysis, we examined the relationships between the different associations and subassociations included in the alliance and, a result of it, the new association Sileno lasiostylae-Arenarietum tenuis is here described.

This syntaxonomical review is one of a number of phytosociological research projects dealing with the ecosystems in the south of the Iberian Peninsula mentioned in the EU Directiva Habitat that we have carried out (Cano et a!., 1997; Diez Garretas & Asensi, 1999; Garcia Fuentes eta!., 200 l; Torres eta!., 2001 ). Thus, we aim at providing the neces­sary tools for a syntaxonomical revision of the class in the Iberian Peninsula, on the same lines as that carried out by Loidi et a!. ( 1997) or, on a European scale, by Valachovich et a!. (1997), de Foucault (1999).

II. METHODS

A. Area under study The Omphalodion commutatae alliance, which includes therophyte communities typi­

cally growing on ultramafic and dolomitic substrates, has its optimum and centre of dis­tribution in the sectors Rondefio and Malacitanto-Almijarense of the Betic province, although it clearly reaches the sector Subbetico (Fig. l). Its characteristic species are (Rivas Martinez et al., 1973):

Arenaria capil/ipes Arenaria retusa lberis fontqueri Jasione penicillata Linaria saturejoides

B. Phytosociology

Narduroides salzmannii Omphalodes commutata Pistorinia breviflora Silene psammitis subsp. lasiostyla

Using the syntaxonomical nomenclature suggested by Izco and Fernandez Gonzalez (200 1 ), but slightly modified, the associations and subassociations so far described are reviewed and examined. The description and nomenclature of the new syntax on have been made accordingly to the International Code of phytosociological nomenclature (Weber et a!., 2000). The nomenclature of the syntaxa included in the original releves and, in gene­ral, of all the syntaxa mentioned, has been homogenised following Castroviejo eta!. (1986-

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Fig. 1.- Biogeographical localization of the plant communities under study. Fig. 1.- Localisation biogeographique des communautes vegetales etudiees.

287

Provincia Betica. 1. Sector Rondeno, 1 a. District Rondense, 1 b. District Bermejense, 1 c. District Anticariense. 2. Sector Malacitano-Aimijarense, 2a. District Malacitano­Axarquiense, 2b. District Almijarense, 2c. District Alfacarino-Granatense. 3. Sector Subbetico, 3a. District Subbetico-Maginense, 3b. District Cazorlense, 3c. District Alcaracense, 3d. District Subbetico-Murciano. 4. Sector Alpujarreiio-Gadorense, 4a. District Alpujarreiio, 4b. District Gadorense. 5. Sector Nevadense, 5a. District Nevadense, 5b. District Filabrico. 6. Sector Guadiciano-Bacense, 6a. District Guadiciano­Baztetano, 6b. District Serrano-Bacense, 6c. District Serrano-Mariense, 6d. District Serrano-Estaciense. Provincia Murciano-Aimeriense. 7. Sector Almeriense, ?a. District Almeriense occidental, 7b. District Almeriense oriental, ?c. District Caridemo. Arenario arundanae-Linarietum salzmannii e; Arenario capillipedis-/beridetum fontqueri •: Arenario modestae-Linarietum angustealatae +; Jasonio penicillatae-Linarietum satu­rejoidis linarietosum saturejoidis +, /inarietosum flavae x, linarietosum huteri *; Sileno lasiostylae-Arenarietum tenuis .A.; Violo demetriae-Jonopsidetum prolongoi violetosum demetriae I, /inarietosum platycalicis D.

2001) or, by default (as yet some families have not been included in this book), Tutin eta/.

(1964-1980). The only exceptions to this rule have been:

L. saturejoides var. angustealata (Willmott) Valdes, Anales Univ. Hispa., Ser. Ci. 7:230.

1970.

Linaria salzmannii var.jlava Boiss., Voy. Bot. Espagne 2: 456. 1839

Omphalodes commutata G. Lopez, Anales Jard. Bot. Madrid 37: 83. 1980.

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The occurrence of Arenaria arundana and A. retusa in the associations involved has been modified according to the criterion of Rivas Martinez eta/. ( 1991 ).

Biogeographical references have been homogenised according to Rivas Martinez eta/. (1997) and bioclimatological references (thermotypes), according to Rivas Martinez & Loidi (1999).

The synoptic table I is the result of both the releves available in the bibliography and those carried out to describe the new association. All the species of the class have been taken into account, whereas those companion taxa, a priori considered scarcely significant, have been omitted. A total of 80 rei eves corresponding to 9 syntaxa whose origin is fully explained in table II have been used.

In order to implement the numerical analysis, the records on cover/abundance were transformed according to Van der Maare1 ( 1979). The original matrix made out of 80 rele­ves and 139 taxa was numerically analysed by means of the Canoco programme (ter Braak, 1988). The multivariate analysis used was a correspondence analysis (CA) with which we obtained the relationships between the types of vegetation under study (releves).

III. RESULTS

On proposing the Arenario modestae-Linarietum angustealatae association (Perez Raya et a/., 1991 ), two subassociations were considered, the typical one and another of transition towards pastures of Erophilo spathulatae-Hornungietum petraeae, Arenario modestae­Linarietum angustea/atae hornungietosum petraeae, both recorded by Izco and Femandez­Gonzalez (2001) after having solved all nomenclature problems. Following the suggestion of Alcaraz ( 1996), we agreed that these situations of ecotone must be dealt with as variants. When typifying, Izco and Femandez-Gonzalez (200 1) also give support to the two subas­sociations suggested by Asensi et a/. ( 1979) for the Rondeiia association Viola demetriae­Jonopsidietum prolongoi (the typical one and linarietosum platyca1icis), but they call the first one (which is obviously the typical one, since the type releve coincides with that of the association) vio/etosum demetriae, inadequately refusing to use both the base name given by the authors (typicum) and ionopsidietosum prolongoi.

The Sileno lasiostylae-Arenarietum tenuis nov. association had already been invalidly suggested by Gomez Mercado (1989) and mentioned (invalidly too) by several authors (Alcaraz eta/., 1991; Peinado eta/., 1992; Sanchez Gomez & Alcaraz, 1993; Cano eta!., 1999). It is a therophyte pasture growing on sandy areas and xeric, dolomitic screes of the supra and oromediterranean belts in the sector Subbetico, where it is endemic. It is cha­racterised by Arenaria modesta subsp. tenuis, an endemic taxon of the Sierras de Cazorla, Segura, Alcaraz, Magina, Ia Sagra, Cabra and surrounding calcareous soils (Lopez, 1990), Silene psammitis subsp. lasiostyla, Narduroides salzmannii and Chaenorhinum macropo­dum subsp. degenii. The association is part of the Berberido-Querceto rotundifoliae S. and Daphno-Pineto sylvestris S. series. Table IV shows eight releves from which we chose number 4 as holotype 4.

IV DISCUSSION

Table III summarises the ecological conditions (biogeography, thermoclimate and substra­te) and both the characteristic and differential species of the nine syntaxa considered within

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-1.0 +1.0

-1.0 +1.0

289

Fig. 2.- CA of the original data matrix. Fig. 2.- CA de Ia matrice des donnees

originales. Arena rio arundanae-Linarietum salzmannii e; Arenario capillipedis-lberidetum fontqueri •; Arenario modestae-Linarietum angustealatae +; Jasonio penicillatae-Linarietum saturejoi­dis linarietosum saturejoidis +. linarietosum f/avae x, linarietosum huteri *; Sileno lasios­tylae-Arenarietum tenuis .A.; Viola demetriae­Jonopsidetum prolongoi violetosum demetriae I, linarietosum platycalicis D.

Fig. 3.- CA of a discrete set of data of the original matrix.

Fig. 3. CA d'un ensemble reduit de donnees appartenant a Ia matrice d'origine. Arenario capillipedis-lberidetum fontqueri •; Arenario modestae-Linarietum angustealatae +; Jasonio penicillatae-Linarietum saturejoi­dis /inarietosum saturejoidis +, linarietosum flavae x, linarietosum huteri *; Arenarietum tenuis .a.; Viola demetriae-Jonopsidetum pro­longoi violetosum demetriae I, linarietosum platycalicis D.

Omphalodion. As can be observed, the main ecological factors determining them are, in each biogeographical unit, the type of substrate and the rich endemic flora linked to each of them (edaphic taxa), thermotypes playing no significant role in this case, as happens in other phytosociological groups of vivacious communities prospering in the same habitats. This circumstance is particularly apparent in the Rondeiio sector, where slates, serpentines, dolomites and limestones are inhabited by a number of specialised associations: Arenario arundanae-Linarietum salzmannii grows on the sandstones and slates of the Sierra de la Pizarra, Arenario capillipedis-Iberidetum fontqueri on the serpentines of the Sierra Bermeja, Jasonio penicillatae-Linarietum saturejoidis on the dolomites and Viola deme­triae-Jonopsidietum prolongoi on dolomitic and oolitic limestones. However, associations absent in the Rondeiio sector always grow on dolomitic soils.

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The correspondence analysis (CA) of the 80 releves and the 139 taxa which they inclu­de (Fig. 2) clearly shows four distinct groups which we have labelled with the letters A, B, C y D. As can be observed, the most decisive factor is again substrate, represented here along the axis y. The releves recorded on limestones appear far to the positive y-values (group A, Violo-Jonopsidietum prolongoi), whereas those recorded on slates and sand­stones (group D, Arenario arundanae-Linarietum salzmannii) appear on the opposite extreme with negative y-values. They are both two rare types of substrates for the alliance. On the other hand, the releves of dolomites and ultramafic substrates (the typical ecology of the alliance) tend to be plotted in the middle of the axis y.

Biogeographical distribution, though less evident, is represented along the axis x. Here, the subbetico rei eves, that is, the most distant from the distribution centre of the alliance, are grouped towards the positive extreme. This same idea is supported by the proximity, in the positive half of the axis x, of the alfacarino-granatense and subbetico releves, coming from neighbouring biogeographical units, while the bermejense releves are plotted in the opposite extreme.

In order to make the relationships of the rei eves of the group B (Fig. 2) clearer, we car­ried out a second analysis of the 49 releves and 84 taxa corresponding to the releves of the associations included in this group. In this second analysis shown in figure 3, the most remarkable feature is the independence of the serpentine-dwelling association (group G, Arenario capillipedis-lberidetum fontqueri), in the positive half of the axes x andy, from both the dolomite-dwelling alfacarino granatense Arenario modestae-Linarietum angus­tea/alae (group E) and the also dolomite-dwelling Rondeiia Jasonio penicillatae­Linarietum saturejoidis (group F). The segregation of the Arenario capillipedis-lberidetum fontqueri association, which, according to Perez Raya et a/. ( 1991 ), has been separated into an independent suballiance (Omphalodenion commutatae), as opposed to the other taxa included in the suballiance Silenenion germanae, can never be clearly observed.

Syntaxonomical scheme Helianthemetea (Br.-Bl. in Br.-Bl., Roussine & Negre 1952) Rivas Goday & Rivas Martinez 1963 em. Rivas Martinez 1978

Trachynietalia distachyae Rivas Martinez 1978 Omphalodion commutatae Rivas Martinez, Izco & Costa ex Izco 1976 corr. Perez Raya, Molero & Lopez Nieto 1991

Omphalodenion commutatae Arena rio capillipedis-lberidetum fontqueri Rivas Martinez, Izco & Costa ex Izco 1976

Silenenion germanae Perez Raya ex Nieto, Cabezudo & Trigo 1989 Arenario arundanae-Linarietum salzmannii Asensi, Diez-Garretas & Esteve 1979 corr. Rivas Martinez, Asensi, Molero Mesa & F. Valle 1991 Arenario modestae-Linarietum angustealatae Perez Raya in Perez Raya, Molero Mesa & Lopez Nieto 1991 Jasiono penicillatae-Linarietum saturejoidis Rivas Martinez, Izco & Costa ex Izco 1976

linarietosum saturejoidis linarietosum flavae Nieto Caldera & Cabezudo ex Izco & Fernandez Gonzalez 2001 linarietosum huteri Nieto Caldera, Perez & Cabezudo ex Izco & Fernandez Gonzalez 2001

Sileno lasiostylae-Arenarietum tenuis ass. nov.

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Viola demetriae-Jonopsidietum prolongoi Asensi, Diez-Garretas & Esteve 1979 violetosum demetriae Asensi, Diez Garretas & Esteve ex Izco & Fernandez Gonzalez 2001 linarietosum platycalicis Asensi, Diez Garretas & Esteve 1979 ex Izco & Fernandez Gonzalez 2001

Acknowledgement - The authors acknowledge Dr. Jesus lzco Sevillano for revising our manuscript.

Table 1.- Synoptic table of the Omphalodion commutatae alliance. The meaning of columns is given in table II.

Tableau 1.- Tableau synoptique de !'alliance Ompha/odion commutatae.

4 6 7 9

Linaria salzmannii ~ Arenaria capillipes [] lberis fo/ltqueri Arenaria retusa

GJ Linaria saturejoides var. angustea/ata

~ [] D Linaria saturejoides Jasione penicillata Linaria salzmannii var.Jlava Linaria huteri Arenaria modesta subsp. tenuis ~ Chaenorhinum macropodum subsp. degenii

m rn Jonopsidium prolongoi I Viola demetria Linaria platycalyx -Ononis saxico/a -Arenaria arundana IV III v III Jasione blepharodon lli IV I IV Silene psammitis subsp. lasiostyla IV III III v Narduroides salzmannii v v v Ill Ill Ompha/odes commutata v IV v v Rumex bucephalophorus III u III v Hornungia petraea II II lli v v A rena ria modes/a subsp. modesta v v IV Tuberaria guttata IV II v II Erophila verna v v Minuartia hybrida Ill Ill III Euphorbia exigua IV II Ill Silene germana IV II u Valantia hispida IV III II Astero/inon linum-stellatum III Leontodon taraxacoides subsp. longirostris III III Polvcarpon tetraphyllum subsp. diphyllum IV III II Linaria viscosa II II II Chaenorhinum macropodum ssp. macropodum III Platycapnos tenui/oba subsp. para/lela II C(vpeola jonthlaspi III II Cerastium gracile Desmazeria rigida Lagunls ovatus II III Prolongoa pectinata III Saxifraga tridactylites II Chaenorhinum rubrifolium subsp. raveyi III Jasione montana subsp. echinata II Pistorinia breviflora III Pistorinia hispanica II

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Table II.- Number and origin of releves used for synoptic table and numerical analysis. Tableau 11.- Origine des inventaires utilises dans Ia realisation du tableau synthetique et de

!'analyse numerique. Arenario arundanae-Linarietum salzmannii. 7 inv: Asensi eta/. 1979, table 4: 1-7.

2 Arenario capillipedis-Iberidetumfontqueri. 8 inv: Asensi eta/. 1979, table 2: 1-8. Arenario modestae-Linarietum angustea/atae. 7 inv: Perez Raya eta/. 1991, table 1: l-6.

4 Jasonio penicillatae-Linarietum saturejoidis linarietosum saturejoides 7 inv: Asensi eta/. 1979, table I: l-5; Mota 1990, table 27: 1-2;

5 Jasonio penicillatae-Linarietum saturejoidis linarietosumflavae. 12 inv: Nieto eta/. 1989, table 4: 1-12. 6 Jasonio penici//atae-Linarietum saturejoidis linarietosum huteri. I I inv: Nieto et a/. 1988, table 5: I-ll. 7 Si/eno /asiosty/ae-Arenarietum tenuis. 12 inv: table 4 (en este articulo):l-8; Arrojo 1994, table 18: 2-5. 8 Violo demetriae-Jonopsidetum prolongoi violetosum demetriae. 4 inv: Asensi et a/. 1979, table 3: l-4. 9 Vio/o demetriae-Jonopsidetum pro/ongoi /inarietosum platyca/icis. 8 inv: Asensi eta/. 1979, table 3: 5-9; Martin Osorio

1993, table 12: 1-3.

Table IV, Tableau IV.- Sileno lasiostylae-Arenarietum tenuis ass. nov. Order number 2 4 5 6 7 8 Altitude (I = 10m) 120 170 145 145 140 140 180 190 Cover(%) 10 10 20 20 30 10 10 10 Slope(%) 10 10 20 30 40 30 Exposure NW E E E s SE

Association and alliance features Anmaria modesta subsp. tenuis 2 2 2 2 2 Si/ene psammitis subsp. /asiosty/a + Narduroides salzmannii 2 Chaenorhinum macropodum subsp. degenii

Order and class features Vu/pia unilateralis 2 Campanula /usitanica Cruciane//a angustifolia + 2 + + Arabis recta + + I + Linaria aernginea + I Minuartia hybrida 2 Cerastium brachypeta/um Hornungia petraea 2 2 Rumex bucepha/ophorus + Alyssum minus + Veronica praecox

Companions Bromus tectorum Poa /igu/ata Cynosurus echinatus +

In addition: lnv. 1: Asterolinon /inum-stellatum +,Linaria viscosa 1, Petrorhagia nanteuilii 1, Scabiosa ste//ata !; lnv. 2: Apera interrupta +, Centaurea calcitrapa +, Erophi/a verna+, Jonopsidium prolongoi 1, Minuartia campestris I, Va/eriane//a pumila I, Viola kitaibeliana +, Ziziphora hispanica +; lnv. 5: Aegilops genicu/ata +, Aegilops triuncia/is +, Bromus madritensis I, Desmazeria rigida I, Trifolium scabrum I; lnv. 8: Odontites /ongif/ora +.

Localities: I. J. Sierra de Cazorla, towards the Parador Nacional "El Adelantado", 30SWG0395; 2. J. Sierra de Segura, near to Cortijo de La Cabrilla, 30SWGI999; 3 a! 6. J. Sierra de Segura, Los Arenales, 30SWGI496;7. GR. Sierra de Castril, Barranco de los Quemaos, 30SWG1588; 8. GR. Sierra de Castril, Los Tejos, 30SWGI588.

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294

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