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Minuartio valentinae-Quercetum pyrenaicae: a new Iberian broad-leaved oak forest in the eastern coastal mountains and their seral plant communities/Minuartio valentinae-Quercetum pyrenaicae: una nueva serie de vegetacion de los melojares de las montanas levantinas costeras y sus comunidades seriales.

INTRODUCTION

The distribution of the marcescent Pyrenean oak on the Iberian Peninsula is concentrated mainly in sites with a sub-Mediterranean climate (moderate or compensated summer drought) in the northwest. This, together with its preference for siliceous soils (less common in the east), means that only reduced samples remain in eastern mountain zones, and usually in protected areas surrounded by evergreen (generally basophile) vegetation. They appear over 1000 m asl and have traditionally been included in the supra-mediterranean association Cephalanthero rubrae-Quercetum pyrenaicae. All are at risk of local extinction due to their level of isolation (mainly edaphic) and fragmentation (PEREZ BADIA, 2003). The Prades mountains in Catalonia, Pina, Penyagolosa, Castielfabib, and Tuejar in Valencia are sites for which there is bibliographic evidence of the presence of this oak forest (BRAUN-BLANQUET, 1934; FONT QUER, 1934; BOLOS, 1967; Vigo, 1968; COSTA & al., 1985; HERREROS, 2010; Rosello, 1994; MATEO & AGUILELLA, 1990), and Prades and Penyagolosa are its best representations in the eastern Iberian peninsula. Its composition is high in nemoral herbs, and often includes other phanerophytes in the arboreal stratum with which it is in contact, mainly Pinus sylvestris s.L, in addition to Juniperus communis. Different Cytisus and Erica appear frequently in the shrub layer, also accompanied by the typical spiny border shrubs Crataegus monogyna or Prunus spinosa.

Although the vegetation of the Sierra de Espadan has been studied, as a whole, by various authors (BELTRAN, 1911; RIVAS GODAY & BORJA, 1961; BOLOS, 1967, 1975; BOLOS & VIGO, 1979; MATEO & AGUILELLA, 1990), its oak forests are the least studied of all those on the eastern Iberian Peninsula. This can be explained by its isolated location and relatively difficult access, and partly because historically this has been a managed area, as it is located in cooler locations that are ideal for farming (MERLE & FERRIOL, 2008), reducing the forest area, currently under regeneration.

The Espadan Triassic mountain range, although not very high in altitude (highest peak: La Rapita at 1106 m asl) captures more rain and humidity than nearby territories. Its NW-SE orientation leads to an annual average rainfall of over 600 mm on the lower slopes, in addition to the contribution of the frequent mists in the area. Espadan soils comprise mainly Buntsandstein sandstones and argilites, characterized by neutral pH, and locally by the formation of deep argillic horizons that allow good water reserves. Both these factors are instrumental in making cork oak the main vegetation in the mountain range, with a predominance of maritime pine (Pinuspinaster s.l.) in drier areas. Heathlands are found on the forest fringes; they are the main substitution shrubs for cork oak forests (COSTA & al., 1985) in sites with a cooler microclimate and for Quercus pyrenaica formations. Several authors have highlighted the special floristic character of this site where Temperate, Iberian-Moroccan and endemic species converge (RIVAS GODAY & BORJA, 1961; MATEO & AGUILELLA, 1990); some have also delimited an independent chorologic element inside the Valenciano-Tarraconense sector, supported by endemics such as Minuartia valentina, Centaurea paui, Dianthus multiaffinis or Biscutella calduchii (O. Bolos & Masclans) Mateo & M.B. Crespo (ROSELLO, 1994; COSTA & al., 1985).

The main aim of our paper is to describe a new association of broad-leaved oak forests in the Sierra de Espadan. This objective was developed as part of a larger study of all the Quercus pyrenaica forests on the Iberian Peninsula. The results of this study emphasised the significant floristic difference between the forests described here and the rest, even when they appear at similar altitudes and bioclimatic belts. We have also analysed--from a purely phytosociological point of view--the following main seral plant communities: heathlands appearing as forest fringes or as a substitution plant community when the forest is destroyed; and shrublands representing a more degraded stage.

MATERIALS & METHODS

We compiled a data set of published releves on Quercus pyrenaica forests in eastern areas of the Iberian Peninsula (Table 1). These releves were previously analysed to check the relationships between central and eastern Quercus pyrenaica forests (Vilches & al., 2013). We used Principal Components Analysis (PCA) on the site-by-species matrix containing presence-absence values to display the compositional variation among the Quercus pyrenaica woodlands in the study. Using indicator species analyses with site group combinations ('multipatt' function of the R package 'indicspecies') we determined the indicator species of each massif, in addition to the common species in forests belonging to those different massifs. The number and identity of the indicators suggest how the different geographical locations could be related.

For the seral communities we studied the data published in the Catalano-Valenciano biogeographical subprovince (Valenciano-Tarraconense sector), which includes: BRAUN-BLANQUET (1940)ZELLER (1959), BOLOS (1967) MATEO (1983), MATEO & MANSANET (1982), COSTA & al. (1985), GARCIA-FAYOS (1991), MERLE & FERRIOL (2008).

We have followed the taxonomic nomenclature proposed by the published volumes of Flora Iberica (CASTROVIEJO & al., 1986-2013) if the authority is not indicated; and the compilation of the Euro+Med PlantBase (2006-2013) for the rest. In addition we have followed RIVAS-MARTINEZ & al. (2007, 2011b) proposals for the biogeographical and bioclimatological characterisation of the studied forest.

RESULTS & DISCUSSION

Minuartio valentinae-Quercetum pyrenaicae ass. nova hoc loco

[Holotypus ass. Table 2, rel. 2; (Merle & Ferriol, 2008)]

The first axis of the PCA (Figure 1) separated the pre-coastal releves with the lowest altitudes--Sierra de Espadan (downward triangles)--in the most positive part of the diagram, with the most central releves-Sierra de Guadarrama (solid circles) and Sierra de Ayllon (upward triangles)--in the negative part. The releves assigned to Cephalanthero-Quercetum pyrenaicae O. Bolos & Vigo in O. Bolos 1967 and Luzulo-Quercetum pyrenaicae Rivas-Martinez 1964 in the eastern localities of Cuenca, Teruel, Valencia and Catalonia appeared only in the middle, leaving Penyagolosa, the highest peak in the upper section. Thus the first PCA axis could be interpreted as a continentality-oceanity gradient, with releves from the Sierra de Espadan near the coast, and clearly separated from the releves of the central Iberian Peninsula. The second axis could be related to a Mediterranean (aridity) gradient, with releves with shorter or compensated summer drought located in the most positive part, clearly separated from the Sierra de Espadan and Sistema Central, which undergo higher summer dryness. Most of these forests are located at higher altitudes than the coastal Sierra de Espadan (Figure 1), where retention of clouds formed by evaporation from the Mediterranean Sea becomes a factor of vital importance to compensate aridity. Precipitation and fog increase the atmospheric humidity of these mountains, where it is common to find ferns and other cryptogrammic species such as Asplenium onopteris, Polypodium cambricum or Polypodium vulgare (MATEO & AGUILELLA, 1991).

The Sierra de Espadan generated the maximum number of exclusive indicator species, including a high number of endemisms that separated it from releves ascribed to Cephalanthero-Quercetum pyrenaicae from Penyagolosa and Prades (Vilches & al., 2013). Both mountains have almost three months of frost risk in winter, and both Quercus pyrenaica forests are related to Pinus sylvestris s. l. and Cistion laurifolii seral communities. Only the pre-coastal location of Prades (Catalonia) shares certain species with Espadan--such as Rosa pouzinii, Cistus albidus or Asplenium onopteris--in addition to some thermophilous and calcareous species, although the analysis (Figure 2) confirmed a differentiated floristic composition of the latter. In fact this difference was also apparent when this community was compared to the whole Iberian Peninsula (VILCHES & al., in press).

Minuartio valentinae-Quercetum pyrenaicae corresponds to silicicolous subhumid forests of Quercus pyrenaica, appearing over rodenos in the upper mesomediterranean to lower supramediterranean belts of the Sierra de Espadan (Castellon). It is found in sites with reduced continentality and increased humidity due to crypto-precipitations and also to the deep limy soils. Its distribution is thus reduced and localized, specifically on the northern slope of the highest peak: La Rapita (as indicated by MATEO & AGUILELLA, 1990; ROSELLO, 1994). The territory has been cultivated for centuries, and some individuals of Quercus pyrenaica can be found growing on abandoned terraces. The forests have a strong level of regeneration and usually include evergreen phanerophytes from the forests with which they are in contact, such as Quercus suber, Quercus rotundifolia and Pinus pinaster s. l, in addition to shrubs (Erica arborea, Ulexparviflorus, Prunus spinosa). Quercetea ilicis species are widely represented in these forests, in contrast with the Quercus pyrenaica locations of the supramediterranean Cephalanthero-Quercetum pyrenaicae (Penyagolosa or Prades), which have an increased Eurosiberian species composition (Vilches & al., 2013). Thus the absence of some species such as Astragalus glycyphyllos, Primula veris subsp. columnae, Veronica officinalis, Poa nemoralis, Campanula trachelium and Lapsana communis, together with its chorology and dynamism, differentiate it from Cephalanthero-Quercetum pyrenaicae.

As is known, Pyrenean oak requires a minimum annual rainfall of 600 mm to become established, as well as a site that compensates the Mediterranean summer drought (costa, 1997). The timing of the rainfall in this area (autumnspring-winter-summer) facilitates a faster recovery of water reserves (ROSELLO, 1994), enough for Minuartio-Quercetum pyrenaicae to grow in the most humid sites throughout the entire Sierra de Espadan, specifically sub-humid locations (Vall de Almonacid, 670 mm; Bolos & Vigo, 1979; NINYEROLA & al., 2005). There is an absence of most of the drought tolerant species of the Valenciano-Tarraconense sector such as Phillyrea angustifolia (including the subhumid Ph. latifolia), Rhamnus alaternus or Pistacia lentiscus; but also a lack of thermophilic shrubs frequent in other mesomediterranean Quercus pyrenaica forests (Arbutus unedo, Viburnum tinus).

On southern slopes or in more exposed areas, these forests come into contact with the Valencian cork forests of Asplenio onopteridis-Quercetum suberis, and also with Hedero-Quercetum rotundifoliae in drier areas. Our community shares more species with the former, mainly in the understory, where the rocky silicicolous substrate allows the inclusion of Minuartia valentina and other related Espadan endemisms such as Biscutella calduchii or Scrophularia tanacetifolia; it also shares the heath-land fringe community described below.

Hedero helicis-Ericetum arboreae (Costa, Peris, Figuerola & Stubing 1985) RIVAS-MART. & al. 2011 (=Cytiso villosi-Ericetum arboreae Costa, Peris, Figuerola & Stubing 1985 non Zeller 1959)

We agree with Costa & al. (1985) regarding the floristic composition of seral scrub plant communities that form part of the vegetation series of Minuartio-Quercetumpyrenaicae. Hedero-Ericetum is not only the first degraded plant community of Asplenio-Quercetum suberis in the Sierra de Espadan; it is also present as a seral state of Minuartio-Quercetum, although impoverished in scrubs such as Cytisus villosus or Ruscus aculeatus (Merle & Ferriol, 2008). It is found in shady areas where soils retain enough water, and has a coastal influence in the upper mesomediterranean to lower supramediterranean bioclimatic belt. They are included in Quercetea ilicis and not in Cytisetea as it could be expected (RIVAS-MARTINEZ & al., 2011; GAVILAN & al., 2011).

We followed the criteria of RIVAS-MARTINEZ & al. (2011) to distinguish Espadan heathlands (Cytiso villosi-Ericetum arboreae Costa, Peris, Figuerola & Stubing 1985), from the species reported by ZELLER (1959) to describe a similar plant community in Catalonia dominated by Erica arborea and Cytisus villosus, together with other brooms such as Teline monspessulana and Cytisus scoparius subsp. reverchonii. Espadan heathlands are lacking most of these species, although may include Cytisus villosus. Moreover, as mentioned above, in the territory of Minuartio-Quercetum pyrenaicae there is an absence of drought-tolerant and thermophilic species inside the Hedero-Ericetum, such as Ruscus aculeatus, Smilax aspera, Clematis flammula, etc. (Table 2).

Uliciparviflorae-Cistetum albidi ass. nova hoc loco [Holotypus ass. Table 4, rel. 1; (Merle & Ferriol, 2008)]

They represent rockrose-gorse shrublands that appear in a more degraded stage than Hedero-Ericetum arboreae. Of all the associations described in the area, the releves included in the new association Ulici-Cistetum are close to those of Pino pinastri-Cistetum salvifolii Bolos 1967 given by Costa & al. (1985) and recently synonymized by RIVAS-MARTINEZ & al. (2002) to Calicotomo spinosae-Cistetum crispi Br.-Bl. 1940. The original releves given by BRAUN-BLANQUETIN 1940 showed the dominance of Calicotome spinosa, Cistus crispus, and even Erica cinerea in Catalonia (on granitic soils). Bolos (1967) included maritime pine (Pinus pinaster s. l.), Lavandula stoechas and Erica arborea to the set, and indicated in a previous work the enormous local variability of this association, in which he suggested including gorse bushes with various Cistaceae that appeared in Espadan and neighbouring mountains (Bolos, 1957; RIVAS-MARTINEZ, 1979). In our case, the community lacks these species (Calicotome spinosa, Cytisus villosus or Lavandula stoechas) and is particularly enriched with Rosmarinetea species such as Cistus albidus and Ulex parviflorus.

The absence of thermophilous taxa such as Calicotome spinosa or Cistus crispus in Ulici-Cistetum can be also explained in terms of altitude (900-1106 masl), with the latter found at higher altitudes than taxa of Calicotomo-Cistetum. Frequent fires also favour the dominance of Ulex parviflorus and Cistus albidus, which are quite common in some areas of the Sierra de Espadan such as La Rapita (MERLE & FERRIOL, 2008).

CONCLUSIONS

Minuartio valentinae-Quercetum pyrenaicae can be framed perfectly in the Quercion pyrenaicae alliance, which includes mesophytic Quercus pyrenaica forests from the meso--to supramediterranean bioclimatic belts. Previous studies (Herreros, 2010; Vilches & al., 2013) already reflected a clear distinction between this and the adjacent communities, specifically Luzulo forsteri-Quercetum pyrenaicae and Cephalanthero rubrae-Quercetum pyrenaicae.

The series 'Minuartio valentinae-Querco pyrenaicae sigmetum' corresponds to silicicolous subhumid forests of Quercus pyrenaica, appearing on rodenos in the upper mesomediterranean to lower supramediterranean bioclimatic belts of the Sierra de Espadan (Valenciano-Tarraconense biogeographical sector). Hedero helicis-Ericetum arboreae and Ulici parviflorae-Cistetum albidii are their main seral plant communities.

ACKNOWLEDGEMENTS

We want to thank Pru Brooke-Turner the English revision and also to anonymous reviewers for the comments to the manuscript. This paper has been financed by the Madrid Autonomous Government through the research project REMEDINAL2 (S2009/AMB-1783).

SYNTAXONOMICAL SCHEME

QUERCO-FAGETEA SYLVATICAE Br.-Bl. & Vlieger in Vlieger 1937

Quercetalia roboris Tx. in Barner 1931

Quercion pyrenaicae Rivas Goday ex Rivas-Mart. 1964

Minuartio valentinae-Quercetum pyrenaicae ass. nova

QUERCETEA ILICIS Br.-Bl. ex A. & O. Bolos, 1950

Pistacio lentisci-Rhamnetalia alaterni Rivas-Mart. 1975

Ericion arboreae (Rivas-Mart. ex Rivas-Mart. & al. 1986) Rivas-Mart. 1987

Hedero helicis-Ericetum arboreae Costa & al. 1985 in Rivas-Mart. 2011

CISTO-LAVANDULETEA Br.-Bl. in Br.Bl., Molinier & Wagner 1940

Lavanduletalia stoechadis Br.-Bl. in Br.Bl., Molinier & Wagner 1940 em. Rivas-Martinez 1968

Cistion ladaniferi Br.-Bl. ex A. & O. Bolos 1950

Ulici parviflorae-Cistetum albidi ass. nova

doi: 10.5209/rev_LAZA.2013.v34.n1.43578

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Received: 4 October 2013

Accepted: 5 December 2013

Beatriz Vilches de la Serna (*), Hugo Merle (**), Maria Ferriol (**), Daniel Sanchez-Mata (*) & Rosario G. Gavilan (*)

* Departamento de Biologia Vegetal II. Facultad de Farmacia. Universidad Complutense. E-28040 Madrid, Spain. E-mail: bvilches@ucm.es; dsmata@ucm.es; rgavilan@ucm.es

** Instituto Agroforestal Mediterraneo. Universidad Politecnica de Valencia. Camino de Vera 14. Valencia 46022. Spain. Email: humerfa@upvnet.upv.es; mafermo@upvnet.upv.es

Table 1
Summary of data gathering according to the locations studied

No. Original assignment            Locality      Province      Distance
                                                               to coast

1   Luzulo forsteri-Quercetum      Guadarrama    Madrid        480 km
2   Luzulo forsteri-Quercetum      Ayllon        Guadalajara   430 km
3   Luzulo forsteri-Quercetum      Albarracin    Teruel        235 km
4   Luzulo forsteri-Quercetum      Boniches      Cuenca        170 km
    Luzulo forsteri-Quercetum      Ranera                      120 km
5   Cephalanthero-Quercetum        Penyagolosa   Castellon     58 km
6   Cephalanthero-Quercetum        Prades        Tarragona     27 km
7   Cephalanthero-Quercetum s.l.   Espadan       Castellon     35 km

No. Original assignment            Altitude         References

1   Luzulo forsteri-Quercetum      1170-1500 mosl   FERNANDEZ-GONZALEZ,
                                                    1991
2   Luzulo forsteri-Quercetum       960-1350 mosl   FUENTE, 1985
3   Luzulo forsteri-Quercetum      1500-1700 mosl   BARRERRA, 1983
4   Luzulo forsteri-Quercetum      1000-1200 mosl   LOPEZ, 1976
    Luzulo forsteri-Quercetum      1280-1380 mosl   MATEO, 1983
5   Cephalanthero-Quercetum        1250-1500 mosl   VIGO, 1968
6   Cephalanthero-Quercetum         950-1050 mosl   BRAUN-BLANQUET,
                                                    1934; FONT QUER,
                                                    1934, BOLOS, 1967
7   Cephalanthero-Quercetum s.l.    900-1106 mosl   MERLE & FERRIOL,
                                                    2008

Table 2
Minuartio valentinae-Quercetum pyrenaicae ass. nova
(Quercion pyrenaicae, Quercetalia roboris, Querco-Fagetea)

Cover (%)                      100    100    100    100    100
Slope (%)                       40     30     35     35     35
Exposure                        N      N      N      N      N
Area ([m.sup.2])               100     50     50     50     50
Releve N.                       1      2      3      4      5

Characteristics
Quercus pyrenaica               5      5      5      5      5
Quercus pyrenaica shrub         5      3      2      3      4
Hedera helix                    +      1      +      1      +
Hieracium glaucinum             2      +      +      .      .
Luzula forsteri                 +      1      .      .      .
Moehringia trinervia            .      .      .      .      .
Biscutella calduchii            1      1      1      1      1
Minuartia valentina             5      4      3      5      +
Polypodium cambricum            +      +      .      .      .
Hieracium amplexicaule          +      .      .      .      .
Erica arborea                   2      2      1      2      2
Asplenium onopteris             1      1      1      1      +
Rubia peregrina                 +      +      1      +      1
Galium maritimum                +      +      +      +      +
Quercus suber                   1      1      1      .      1
Quercus rotundifolia            .      .      .      1      1
Daphne gnidium                         .      .      .      .
Lonicera implexa                +      +      .      +      .
Companions
Brachypodium retusum            3      3      3      4      3
Rubus ulmifolius                +      1      +      +      +
Rosa pouzinii                   1      1      1      +      .
Ulex parviflorus                2      1      +      +      +
Helianthemum origanifolium *    +      +      +      +      +
Carex muricata                  +      +      .      1      +
Euphorbia flavicoma             +      +      +      +      +
Prunus spinosa                  1      1      1      1      .
Pinus pinaster s.l.             2      1      2      2      2
Teucrium chamaedrys             +      +      +      +      +
Crataegus monogyna              1      .      1      .      .
Geranium robertianum            .      .      +      .      .
Cistus albidus                  +      +      .      .      +
Phleum phleoides                .      .      .      .      .
Viola alba                      +      .      .      .      .
Festuca durandoi                .      .      .      .      .
Sedum sediforme                 .      .      .      +      +
Vicia tenuifolia                .      .      .      1      .
Silene inaperta                 .      +      .      .      .
Orobanche alba                  .      .      .      .      .
Cephalanthera rubra             .      .      .      .      +
Thapsia villosa                 .      .      .      .      +
Hieracium sabaudum              .      .      .      .      .
Pteridium aquilinum             .      .      .      .      .

Cover (%)                      100    100    100    100    100
Slope (%)                       20     30     30     30     30
Exposure                        N      N      N      N      N
Area ([m.sup.2])                50     50     50     50     50
Releve N.                       6      7      8      9      10

Characteristics
Quercus pyrenaica               5      5      4      5      5
Quercus pyrenaica shrub         4      5      4      5      4
Hedera helix                    1      +      1      .      +
Hieracium glaucinum             .      +      1      .      .
Luzula forsteri                 +      .      1      +      .
Moehringia trinervia            +      +      +      +      .
Biscutella calduchii            .      +      +      +      +
Minuartia valentina             +      +      .      1      2
Polypodium cambricum            .      .      .      +      +
Hieracium amplexicaule          .      +      .      +      .
Erica arborea                   3      2      1      1      2
Asplenium onopteris             +      1      1      1      1
Rubia peregrina                 1      1      1      1      +
Galium maritimum                +      +      1      1      +
Quercus suber                   1      1      2      +      +
Quercus rotundifolia            .      1      2      +      .
Daphne gnidium                  +      .      +      1      1
Lonicera implexa                .      .      .      .      .
Companions
Brachypodium retusum            2      2      2      3      3
Rubus ulmifolius                +      +      +      +      +
Rosa pouzinii                   +      +      +      +      +
Ulex parviflorus                +      .      .      1      1
Helianthemum origanifolium *    +      .      .      +      +
Carex muricata                  1      +      1      +      +
Euphorbia flavicoma             +      .      +      .      +
Prunus spinosa                  3      +      +      .      1
Pinus pinaster s.l.             1      2      1      .      1
Teucrium chamaedrys             +      +      +      .      +
Crataegus monogyna              +      .      +      .      1
Geranium robertianum            .      +      +      +      .
Cistus albidus                  +      .      .      +      +
Phleum phleoides                .      +      +      +      .
Viola alba                      +      .      .      .      +
Festuca durandoi                .      +      +      +      .
Sedum sediforme                 .      .      .      .      .
Vicia tenuifolia                +      .      .      .      .
Silene inaperta                 .      +      .      +      .
Orobanche alba                  .      +      .      +      +
Cephalanthera rubra             +      .      .      .      +
Thapsia villosa                 .      .      .      .      +
Hieracium sabaudum              .      +      .      +      .
Pteridium aquilinum             .      2      .      .      .

Cover (%)                      100    100    100    100    100
Slope (%)                       30     30     30     30     30
Exposure                        N      N      N      N      N
Area ([m.sup.2])                50     50     50     50     50
Releve N.                       11     12     13     14     15

Characteristics
Quercus pyrenaica               5      5      3      5      4
Quercus pyrenaica shrub         3      3      5      3      2
Hedera helix                    1      .      2      1      4
Hieracium glaucinum             +      .      .      .      .
Luzula forsteri                1.0     .      +      +      +
Moehringia trinervia            .      .      .      .      .
Biscutella calduchii            +      +      +      +      .
Minuartia valentina             2      3      3      3      2
Polypodium cambricum            .      +      .      +      +
Hieracium amplexicaule          .      .      .      +      +
Erica arborea                   2      1      +      +      +
Asplenium onopteris             1      1      1      1      3
Rubia peregrina                 +      +      +      +      +
Galium maritimum                +1     1      +      +      +
Quercus suber                   .      1      +      .      1
Quercus rotundifolia            .      +      +      .      .
Daphne gnidium                  +      +      .      .      .
Lonicera implexa                .      .      .      +      +
Companions
Brachypodium retusum            1      2      3      .      +
Rubus ulmifolius                +      +      +      .      +
Rosa pouzinii                   +      +      +      .      .
Ulex parviflorus                .      +      +      +      .
Helianthemum origanifolium *    +      +      .      +      .
Carex muricata                  .      .      +      +      .
Euphorbia flavicoma             +      +      .      .      .
Prunus spinosa                  +      +      .      .      .
Pinus pinaster s.l.             2      .      .      .      .
Teucrium chamaedrys             +      .      .      .      .
Crataegus monogyna              +      .      .      +      1
Geranium robertianum            .      .      +      +      +
Cistus albidus                  .      +      .      .      .
Phleum phleoides                .      +      +      .      +
Viola alba                      +      +      +      .      .
Festuca durandoi                .      +      .      +      1
Sedum sediforme                 .      +      .      +      .
Vicia tenuifolia                .      .      +      .      1
Silene inaperta                 .      +      .      .      .
Orobanche alba                  .      .      .      .      .
Cephalanthera rubra             1      .      .      .      .
Thapsia villosa                 .      .      .      .      .
Hieracium sabaudum              .      .      .      .      .
Pteridium aquilinum             +      .      .      .      .

Other species: Asplenium trichomanes, Antirrhinum barrelieri, Cistus
salviifolius + in 9; Dactylis glomerata + in 12; Galium idubedae + in
13 and Scrophularia sciophila + in 14.

Localities: All releves from Sierra de Espadan between 900-1106 m asl.
Holotypuss ass. rel. 2

* Helianthemum origanifolium: H. marifolium subsp. origanifolium

Table 3

Hedero helicis-Ericetum arboreae (Costa, Peris,
Figuerola & Stubing 1985) Rivas-Mart. & al. 2011
(Pistacio lentisci-Rhamnetalia alaterni,
Ericenion arboreae, Quercetea ilicis)

Cover (%)                     100   100   100   100   100   100   100

Slope (%)                      30    30    30    30    30    30    30
Exposure                       N     N     N     N     N     N     N
Area ([m.sup.2])               50    50    50    50    50    50    50
Releve N.                      1     2     3     4     5     6     7

Characteristics
Erica arborea                  4     5     5     5     4     4     4
Asplenium onopteris            +     +     +     +     +     +     +
Rubia peregrina                .     +     +     +     +     1     +
Quercus rotundifolia           .     1     +     1     1     .     1
Quercus suber                  1     .     1     +     1     +     1
Galium maritimum               1     +     +     .     +     +     .
Lonicera implexa               .     .     +     1     .     2     .
Daphne gnidium                 1     1     .     .     .     .     .
Companions
Brachypodium retusum           3     4     4     3     3     +     1
Rosa pouzinii                  1     +     +     +     +     1     +
Biscutella calduchii           +     +     +     +     +     +     +
Crataegus monogyna             +     .     +     1     +     1     +
Euphorbia flavicoma            +     +     +     +     +     .     +
Minuartia valentina            .     +     1     1     .     +     +
Rubus ulmifolius               2     .     .     1     +     +     1
Cistus albidus                 1     1     .     1     .     1     +
Ulex parviflorus               2     2     1     .     +     1     .
Prunus spinosa                 .     1     1     1     +     1     .
Helianthemum origanifolium *   .     +     +     +     .     +     .
Hedera helix                   .     .     .     .     1     +     1
Polypodium cambricum           .     .     +     .     .     +     +
Pinus pinaster s.l.            1     .     +     .     2     .     .
Quercus pyrenaica shrub        .     .     +           .     .     +
Vicia tenuifolia               .     .     +     .     .     .     +
Luzula forsteri                .     .     .     .     +     +     .
Hieracium glaucinum            .     .     +     .     +     .     .
Galium idubedae                .     .     +     +     .     .     .
Viola alba                     .     .     +     +     .     .     .
Quercus pyrenaica              .     1     1     .     .     .     .
Phleum phleoides               +     +     .     .     .     .     .

Other species: Sedum sediforme and Thapsia villosa +
in 2; Cistus salviifolius 1 in 4; Asplenium trichomanes,
Scrophularia tanacetifolia, Geranium robertianum, Hieracium
amplexicaule and Antirrhinum barrelieri + in 6.
Localities: All releves from Sierra de Espadan
(Pico La Rapita) between 900-1106 m.a.s.l.

* Helianthemum origanifolium: H. marifolium subsp. origanifolium

Table 4
Ulici parviflorae-Cistetum albidi ass. nova
(Cistion ladaniferi, Lavanduletalia stoechadis,
Cisto-Lavanduletea)

Cover (%)                100    100    100    100    100
Slope                     20     30     30     30     30
Exposure                  N      N      N      N      N
Area ([m.sup.2])         100     50     50     50     50
Releve N.                 1      2      3      4      5

Characteristics
Ulex parviflorus          2      4      4      3      5
Cistus albidus            2      3      3      4      3
Cistus salviifolius       +      1      1      .      1
Companions
Asplenium onopteris       +      +      +      +      +
Biscutella calduchii      .      +      +      +      +
Galium maritimum          .      +      +      +      +
Trifolium campestre       +      .      1      +      +
Sedum sediforme           +      .      +      +      +
Brachypodium retusum      5      4      3      4      3
Euphorbia flavicoma       +      +      +      +      .
Prunus spinosa            1      +      .      1      +
Helianthemum
  origanifolium *         +      +      +      .      +
Trifolium arvense         +      .      +      .      .
Rubus ulmifolius          1      1      1      .      .
Geranium robertianum      1      .      +      .      .
Dactylis glomerata        +      .      +      .      +
Quercus rotundifolia      .      1      +      .      +
Rosa pouzinii             2      +      .      .      +
Thapsia villosa           2      2      1      1      .
Senecio lividus           1      +      +      +      .
Daphne gnidium            1      +      +      +      .
Galium idubedae           .      .      .      .      .
Hedera helix              .      .      .      .      +
Minuartia valentina       .      +      1      .      .
Quercus suber             +      .      .      .      +
Phleum phleoides          .      .      +      .      +
Vicia tenuifolia          .      .      .      .      .
Lonicera implexa          .      .      .      .      .
Silene inaperta           .      .      .      +      .
Rubia peregrina           +      .      .      .      .
Antirrhinum barrelieri    .      .      .      .      .
Erica arborea             .      2      .      .      .
Polypodium cambricum      .      .      +      .      1
Hieracium amplexicaule    +      .      +      .      .

Cover (%)                100    100    100
Slope                     30     30     30
Exposure                  N      N      N
Area ([m.sup.2])          50     50     50
Releve N.                 6      7      8

Characteristics
Ulex parviflorus          4      3      2
Cistus albidus            3      3      4
Cistus salviifolius       .      .      .
Companions
Asplenium onopteris       +      +      +
Biscutella calduchii      +      +      +
Galium maritimum          +      +      +
Trifolium campestre       +      +      +
Sedum sediforme           +      +      +
Brachypodium retusum      4      .      3
Euphorbia flavicoma       .      +      1
Prunus spinosa            1      .      1
Helianthemum
  origanifolium *         +      +      .
Trifolium arvense         +      1      1
Rubus ulmifolius          .      1      1
Geranium robertianum      +      +      +
Dactylis glomerata        +      .      +
Quercus rotundifolia      +      1      .
Rosa pouzinii             1      .      .
Thapsia villosa           .      .      .
Senecio lividus           .      .      .
Daphne gnidium            .      .      .
Galium idubedae           +      +      1
Hedera helix              .      +      2
Minuartia valentina       .      .      1
Quercus suber             1      .      .
Phleum phleoides          +      .      .
Vicia tenuifolia          .      +      2
Lonicera implexa          .      1      +
Silene inaperta           .      .      +
Rubia peregrina           .      .      +
Antirrhinum barrelieri    +      1      .
Erica arborea             +      .      .
Polypodium cambricum      .      .      .
Hieracium amplexicaule    .      .      .

Other species: Crataegus monogyna 1 in 1; Pteridium
aquilinum + in 3; Quercus pyrenaica 1 in 5; Pinus pinaster
1, Asplenium trichomanes + in 6; Festuca durandoi 1
in 7; Scrophularia sciophila and Carex muricata + in 8.
Localities: All releves from Sierra de Espadan (Pico La
Rapita) between 900-1106 m asl. Holotypus ass. rel. 4.

* Helianthemum origanifolium: H. marifolium subsp.
origanifolium
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Title Annotation:articulo en ingles
Author:de la Serna, Beatriz Vilches; Merle, Hugo; Ferriol, Maria; Sanchez-Mata, Daniel; Gavilan, Rosario G.
Publication:Lazaroa
Date:Jan 1, 2013
Words:4320
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