Analysis of the Juniperus oxycedrus L. communities in the centre and south of the Iberian peninsula (Spain and Portugal)

Acta Botanica Gallica ISSN: 1253-8078 (Print) 2166-3408 (Online) Journal homepage: http://www.tandfonline.com/loi/tabg20 Analysis of the Juniperus o...
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Acta Botanica Gallica

ISSN: 1253-8078 (Print) 2166-3408 (Online) Journal homepage: http://www.tandfonline.com/loi/tabg20

Analysis of the Juniperus oxycedrus L. communities in the centre and south of the Iberian peninsula (Spain and Portugal) Eusebio Cano , Alfonso Rodríguez-Torres , Carlos Pinto Gomes , Antonio García-Fuentest , Juan A. Torres , Carlos Salazar , Luis Ruiz-Valenzuela , Ana Cano-Ortiz & Raúl J. Montilla To cite this article: Eusebio Cano , Alfonso Rodríguez-Torres , Carlos Pinto Gomes , Antonio García-Fuentest , Juan A. Torres , Carlos Salazar , Luis Ruiz-Valenzuela , Ana Cano-Ortiz & Raúl J. Montilla (2007) Analysis of the Juniperus oxycedrus L. communities in the centre and south of the Iberian peninsula (Spain and Portugal), Acta Botanica Gallica, 154:1, 79-99, DOI: 10.1080/12538078.2007.10516046 To link to this article: http://dx.doi.org/10.1080/12538078.2007.10516046

Published online: 26 Apr 2013.

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Date: 18 January 2017, At: 14:22

Acta Bot. Gallica, 2007, 154 (1), 79-99.

Analysis of the Juniperus oxycedrus L. communities in the centre and south of the Iberian peninsula (Spain and Portugal) by Eusebio Cano(1), Alfonso Rodríguez-Torres(2), Carlos Pinto Gomes(3), Antonio García-Fuentes(1), Juan A. Torres(1), Carlos Salazar(1), Luis Ruiz-Valenzuela(1), Ana Cano-Ortiz(1) and Raúl J. Montilla(1) (1) Dpto. Biología Animal, Biología Vegetal y Ecología, Facultad de Ciencias Experimentales, Universidad de Jaén, Campus universitario Las Lagunillas, E-23071 Jaén (2) Dirección general de Medio Ambiente Natural, Consejería de Agricultura y Medio Ambiente, Junta de Castilla-La Mancha, C/ Pintor Matías Moreno, E-45071 Toledo (3) Dpto. de Ecología, Universidade de Évora, P-07000 Évora arrivé le 23 mars 2006, accepté le 5 mai 2006 Abstract.- The survey deals with juniper groves of Juniperus oxycedrus, i. e. the edaphoxerophilous communities dominated by J. oxycedrus subsp. oxycedrus and J. oxycedrus subsp. lagunae in the centre and south of the Iberian Peninsula. The study of the vegetation has been carried out using the phytosociological method, complemented by a multivariate analysis to statistically process both bibliographical and field relevés. As a result, a new phytosociological alliance (Juniperion oxycedro-lagunae) encompassing four new associations (Pistacio terebinthi-Juniperetum lagunae, Stipo tenacissimae-Juniperetum lagunae, Echinosparto iberici-Juniperetum lagunae and Cytiso eriocarpi-Juniperetum lagunae) and the community of Phlomis purpurea and J. oxycedrus subsp. lagunae is proposed. These juniper groves are plant formations of general interest for the European Union and exhibit interesting endemic flora and vegetation.

Key words : edaphoxerophilous communities - juniper groves - multivariate analysis - phytosociology - syntaxonomy. Résumé.- L’étude traite des bois de Juniperus oxycedrus, i. e. les communautés édaphoxérophiles dominées par J. oxycedrus subsp. oxycedrus et J. oxycedrus subsp. lagunae dans le centre et le sud de la Péninsule ibérique. L’étude de la végétation a été faite en utilisant la méthode photosociologique, complétée par une analyse multivariée pour classer statistiquement les données bibliographiques et des relevés originaux. On propose une nouvelle alliance phytosociologique (Juniperion oxycedro-lagunae) et quatre nouvelles associations (Pistacio terebinthi-Juniperetum lagunae, Stipo tenacissimae-Juniperetum lagunae, Echinosparto iberici-Juniperetum lagunae, Cytiso eriocarpi-Juniperetum lagunae) et une communauté à Phlomis purpurea - J. oxycedrus subsp. lagunae. Ces bois sont des formations de plantes d’intérêt communautaire (Union européenne) et accueillent une flore endémique et une végétation intéressantes. Mots clés : communautés édaphoxérophiles - bois de Genévrier - analyse multivariée - phytosociologie - syntaxonomie.

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I. INTRODUCTION The Juniperus L. (Cupressaceae) genus encompasses about 50 species with a wide distribution in the northern hemisphere, stretching down to tropical Africa. It is an important genus of gymnosperm shrub-like and tree-like plants with a remarkable occurrence in the Mediterranean region and taxa belonging both to sclerophyllous broad-leaved (class Quercetea ilicis Br.-Bl. ex A. & O. Bolòs 1950) and needle-leaved (class Pino-Juniperetea Rivas-Martínez 1965) vegetation. Among the Mediterranean species, Juniperus oxycedrus L. is most remarkable. This juniper occurs all over the region, reaching, to the west, the east of Portugal, to the south Morocco and to the east the north of Iran (Amaral Franco, 1986). It is considered a characteristic taxon of the Quercetea ilicis class and is included in sclerophyllous plant formations of a climatophilous and edaphoxerophilous character. In the Iberian Peninsula there are several subspecies with clear morphological and ecological differences. The subspecies macrocarpa (Sm.) Ball, with galbuli initially pruinose of up to 1.2-1.5 cm in diameter, has broader leaves (up to 2.5 mm) and tends to occur on dunes and in coastal sandy areas, occasionally even on rocky shores. The Iberian plant formations of this taxon are included in the alliance Juniperion turbinatae Rivas-Martínez 1975 corr. 1987, together with other plant formations dominated by Juniperus navicularis Gand. and Juniperus phoenicea L. subsp. turbinata (Guss.) Nyman, these latter being psamophilous taxa peculiar to sandy areas and dunes in coastal regions. The subspecies oxycedrus and lagunae (Pau ex C. Vicioso) Rivas-Martínez are morphologically and ecologically more akin and occur on hard, either siliceous or carbonated, substrates. The main differences between these two taxa are physiognomical, depending, for example, on the size of the mature pseudofruits or how broad the leaves are. Whereas the oxycedrus subspecies tends to have a shrub-like appearance, with galbuli smaller than 1 cm in diameter and 1-1.5 mm broad leaves, the lagunae subspecies is a tree of good size with a pyramidal profile, galbuli bigger than 1 cm and up to 2 mm broad leaves. As both subspecies can co-exist perfectly well in similar biotopes, a number of authors have frequently confused them. However, the current data on J. oxycedrus subsp. lagunae (Fig. 1) suggest a more continental distribution pattern, with a tendency to colonise the inner regions of the Iberian Peninsula. Its occurrence in the centre, east and south of the Iberian Peninsula is very clear. Here it gives rise to widely spread plant formations, usually on rocky sites and shallow soils on which holm oaks (Quercus rotundifolia Lam.) are no longer dominant or are simply absent because the edaphic requirements for their development are missing. These edaphoxerophilous communities have a permanent character and, although they were originally restricted, they are currently expanding. Frequent forest fires together with deforestation and scrub clearing provoke soil erosion in a growing number of areas. Since these biotopes cannot be colonised by Fagaceae, the forests of Quercus rotundifolia remain restricted to less hostile territories. If the same factors producing this transformation persist, the area colonised by species of the genus Juniperus (Allende et al., 1999) will continue to grow. Since the areas in which these juniper groves occur have no farming value are difficult to reach and a good distance away from urban settlements, the copses of J. oxycedrus subsp. lagunae represent vegetal islands not usually negatively affected by human action. These phytocoenoses appear as reservoirs of interesting endemic taxa of Iberian Peninsula, such as Coincya longirostra (Boiss.) Greuter & Burdet, Cytisus scoparius (L.) Link subsp.

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Fig. 1.- Distribution of Juniperus oxycedrus subsp. lagunae in the Iberian Peninsula. Fig. 1. Distribution de Juniperus oxycedrus subsp. lagunae dans la Péninsule Ibérique.

scoparius, Cytisus striatus (Hill) Rothm. subsp. eriocarpus (Boiss. & Reut.) Rivas Mart., Dianthus crassipes R. de Roemer, Dianthus lusitanus Brot., Digitalis thapsi L., Flueggea tinctoria (L.) G.L. Webster, Lavandula stoechas L. subsp. luisieri (Rozeira) Rozeira, Lavandula stoechas L. subsp. sampaiana Rozeira and Genista hirsuta Vahl. It must also be noted that they occupy areas of great ecological interest declared as SCI habitats (i. e. sites of community importance according to the EU Directive 92/43/EEC), with the occurrence of many Luso-Extremaduran endemic phytocoenoses of chasmocomophytic character such as Coincyo longirostrae-Dianthetum lusitani Melendo in Cano, Melendo & F. Valle 1997, Digitali thapsi-Dianthetum lusitani Rivas-Martínez ex Fuente 1986 and Jasiono marianae-Dianthetum lusitani Rivas Goday (1955) 1964. This reinforces the ecological interest of these plant formations, which need to be studied in order to preserve them.

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By means of the Braun-Blanquet phytosociological method, this paper aims both at analysing those plant communities in the centre and southwest of the Iberian Peninsula which are dominated by Juniperus oxycedrus subsp. oxycedrus and subsp. lagunae and also at suggesting new proposals for the phytocoenoses not yet described. II. MATERIAL AND METHODS A. Study area The study area is in the centre and southwest of the Iberian Peninsula (Spain and Portugal) extending between approximately 3º and 9º west longitude and 37º and 41º north latitude. Biogeographically it represents the Toledan-Taganean and MarianicMonchiquensean sectors (Rivas-Martínez et al., 1997; Rivas-Martínez & Loidi, 1999a) of the Luso-Extremaduran Subprovince (Ibero-Atlantic province). In these Toledan-Taganean territories the juniper groves occupy both the eastern, more continentalised areas and in the Portuguese territories, more oceanic areas extending from Portalegre to Castelo-Branco. In the Marianic-Monchiquensean sector these plant formations tend to occur in the Barranco del Río Viar (Sierra Norte of Seville: Real de la Jara and Cazalla de la Sierra) and in Sierra Morena in Jaen and Ciudad Real (Fig. 2). From a geological point of view, all the sites which can be colonised by these formations of Juniperus oxycedrus subsp. lagunae are always small ranges composed of quartzites, granites, Precambrian slates and even arkoses, with altitudes ranging from 280 m (Barranco del Río Viar, Seville) to 1,257 m (Abulagoso, C. Real) (Direcçao General de Minas and Serviços Geologicos, 1968; I.G.M.E., 1980). In order to bioclimatically define the territories in the centre and south of the Iberian Peninsula with dense populations of Juniperus oxycedrus subsp. lagunae, the subject of this paper, we studied 100 weather stations. According to the criteria of Rivas-Martínez & Loidi (1999b), 29 stations have an ombrothermic index (Io) ranging from 3.6 to 6.3, which indicates therefore a subhumid-humid ombrotype. In the other 71 stations there is an Io ranging from 2.02 to 3.6, which indicates a dry-subhumid ombrotype in the study area. The thermotype ranges from thermomediterranean in the more thermic territories close to the Guadalquivir River valley (south of the Iberian Peninsula) and supramediterranean in the outcrops of the Meseta Central Ibérica, which are more continentalised. However, the average values of Io (3.89), continentality index (Ic = 18.54) and compensated thermicity index (Itc = 284) clearly reveal the territorial dominance of the mesomediterranean thermotype. Continentality values (Ic) range from 10.8 in Santiago Do Cacen (Portugal) to 21.1 in Santa Cruz de Mudela (Ciudad Real, Spain). Thus, in the study area there is a Mediterranean pluviseasonal-oceanic macrobioclimate in the western regions and a Mediterranean pluviseasonal-continental macrobioclimate in the eastern regions (Fig. 3). B. Study of flora and vegetation We have carried out a number of field studies in the area (Spain and Portugal) to make relevés and sample the communities dominated by Juniperus oxycedrus subsp. oxycedrus and subsp. lagunae. These relevés have been made using the Braun-Blanquet phytosociological method (Braun-Blanquet, 1979; Gèhu & Rivas-Martínez, 1981). Our survey also includes phytosociological relevés of previous studies of other authors which mention plant communities with either a physiognomy similar to our samples or a high occurrence

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Biogeographical characterization of the study area Ibero-atlantic province Luso-Extremaduran subprovince 1. Toledan-Taganean sector 2. Marianic-Monchiquensean sector

Symbols n Cytiso eriocarpi-Juniperetum lagunae s Echinosparto iberici-Juniperetum lagunae l Pistacio terebenthi-Juniperetum lagunae + Stipo tenacissimae-Juniperetum lagunae p community of Phlomis purpureaJuniperus oxycedrus subsp. lagunae

Fig. 2.- Map of the study area with the location of the relevés used for the characterisation of the plant formations of Juniperus oxycedrus subsp. lagunae. Fig. 2.- Carte de l’aire d’étude et localisation des relevés utilisés pour la caractérisation des formations à Juniperus oxycedrus subsp. lagunae.

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Fig. 3.- Bioclimatic map of the study area. Fig. 3.- Carte bioclimatique de l’aire d’étude.

of Juniperus oxycedrus in a community. See Appendix 1 to check the data source of each relevé. The taxonomical nomenclature is based on the works in the following preference list: Flora iberica: Castroviejo et al. (eds.) (1986, 1990, 1993a, 1993b, 1997a, 1997b), MuñozGarmendia & Navarro (eds.) (1998), Talavera et al. (eds.) (1999, 2000), Paiva et al. (eds.) (2001), Nieto et al. (eds.) (2003) and Aedo & Herrero (eds.) (2005); Flora de Andalucía Occidental: Valdés et al. (1987); Flora Europaea: Tutin et al. (eds.) (1964-80), with the exception of the taxa: Juniperus oxycedrus subsp. lagunae (Pau ex C. Vicioso) Rivas Mart. in Itinera Geobot. 15 (2): 702 (2002), Cytisus striatus (Hill) Rothm. subsp. eriocarpus (Boiss. & Reut.) Rivas Mart. in Anales Inst. Bot. Cavanilles 34: 540 (1977); Pistacia xsaportae Burnat, Fl. Alp. Marit. 2: 54 (1896). Our syntaxonomical scheme follows the proposal of Rivas-Martínez et al. (2002).

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C. Multivariate analysis The statistical analysis was carried out by means of a data matrix containing the characteristic species of associations and larger units with the companion species of the phytosociological tables. These plant species were taken as floristic variables. The original data matrix consisted of 222 columns (floristic variables) and 154 rows (relevés). The Braun-Blanquet abundance rates were later transformed for statistical analysis according to Van der Maarel (1979). This data matrix was subjected to a preliminary analysis of classification (cluster, Euclid distance, Ward method) in order to check the main groups as defined by the communities rich in Juniperus oxycedrus subsp. lagunae. Later, we implemented two Detrended Correspondence Analyses (DCA) to clarify the groups of relevés which are defined in this survey. This analysis was then applied to the data matrix of relevés with a high occurrence of Juniperus oxycedrus subsp. lagunae. III. RESULTS AND DISCUSSION A. The cluster analysis The cluster analysis classification reveals two large groups: group 1 with 93 relevés and group 2 with 61 relevés (Fig. 4). Group 1 consists of plant formations dominated by different Quercus spp. species. These plant formations represent the climax vegetation or first dynamic stage of the climatophilous series of this territory and therefore their successional dynamics are well established. Group 2 includes all communities with a high occurrence of Juniperus oxycedrus subsp. lagunae which we will deal with later. A closer analysis of group 1 reveals two large subgroups. In the first subgroup (Fig. 4, 1a), which is notably exposed to oceanic influence and subhumid-humid ombrotypes; the most remarkable formations are the thermomediterranean Gaditan juniper groves of Cytiso tribracteolati-Juniperetum oxycedri A.V. Pérez, Galán & Cabezudo in A.V. Pérez, Galán, P. Navas, D. Navas, Y. Gil & Cabezudo 1999 (R35-37) and other climactic formations in which the mesomediterranean Luso-Extremaduran cork oak groves of Poterio agrimonioidis-Quercetum suberis Rivas Goday in Rivas Goday et al. 1960 and facies of holm oak woods rich in gall oaks (Quercus faginea Lam.) and cork oak trees (Quercus suber L.) (R56-64, R124-129, R136-140) are dominant. The second subgroup (Fig. 4, 1b) consists of a number of tree-like and shrub-like plant formations which occur in inner peninsular territories with a more continentalised character and dry-subhumid ombrotypes: they are supramediterranean Carpetan-Leonese juniper groves of Festuco elegantis-Juniperetum oxycedri (Rivas-Martínez & Sánchez Mata in Sánchez Mata 1989) Sánchez Mata 1999 (R38-41), supramediterranean Carpetan-Leonese holm oak woods of Junipero oxycedriQuercetum rotundifoliae Rivas-Martínez 1965 (R65-72), mesomediterranean and LusoExtremaduran kermes oak groves of Hyacinthoido hispanicae-Quercetum cocciferae (Rivas Goday in Rivas Goday et al. 1960) Peinado & Martínez-Parras 1985 (R75-81), and a large number of relevés (group of relevés from R43 to R109) which correspond to the different variants and subassociations of mesomediterranean Luso-Extremaduran holm oak woods (Pyro bourgaeanae-Quercetum rotundifoliae Rivas-Martínez 1987). Group 2 has been defined by the edaphoxerophilous plant formations of Juniperus oxycedrus subsp. lagunae, which acts as the first and foremost distinctive character when compared to the previous group. In all the relevés of group 2, Juniperus oxycedrus subsp. lagunae is, together with Juniperus oxycedrus subsp. oxycedrus, dominant. These communities tend to occur in small ranges with granitic or quartzitic materials, intense sun

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Fig. 4.- Dendrogram of the communities with J. oxycedrus subsp. oxycedrus and J. oxycedrus subsp. lagunae occur. Fig. 4.- Dendrogramme des communautés à J. oxycedrus subsp. oxycedrus et subsp. lagunae.

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exposure and high evapotranspiration. They are edaphoxerophilous communities in contact with biotopes of deep soils occupied by cork, holm and Pyrenean oak groves which cannot compete with the above-mentioned subrupicolous species. Having confirmed the independence of these Juniper groves of Juniperus oxycedrus subsp. lagunae from the other tree-like and shrub-like plant formations considered in the cluster, let us concentrate on the analysis of group 2 in order to assess its diversity in regard to plant associations. B. Study of the permanent communities of J. oxycedrus subsp. lagunae To study the juniper groves of Juniperus oxycedrus subsp. lagunae the original data matrix was reduced to the relevés of group 2. By so doing, a new matrix of 61 relevés and 170 taxa arises and then a multivariate analysis, a DCA-ordination now, is once more implemented. Fig. 5 shows the groups of relevés characteristic of each group of samples.

Fig. 5.- Detrended Correspondence Analysis (DCA) with the groups of relevés characteristic of the associations Echinosparto-Juniperetum lagunae (A1) and Cytiso-Juniperetum lagunae (A2). Fig. 5.- Analyse de correspondance détendancée (DCA) des groupes de relevés caractéristiques des associations Echinosparto-Juniperetum lagunae (A1) et Cytiso-Juniperetum lagunae (A2).

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First we see (Fig. 5, A1) a group of nine relevés concentrated around axis 1 (R143-145, R147, R148, R150, R152-154) and characterised by the occurrence of Echinospartum ibericum Rivas Mart., Sánchez Mata & Sancho and Adenocarpus argyrophyllus (Rivas Goday) Caball., together with other silicicolous species or species peculiar to supramediterranean areas, such as Festuca elegans Boiss. subsp. elegans. Apart from the occurrence of the already mentioned floristic elements, this central group of relevés also has in common the fact that they have all been sampled in the supramediterranean Toledan-Taganean and Marianic-Monchiquensean territories. Consequently, it is a group which characterises a permanent community dominated by Juniperus oxycedrus subsp. lagunae and colonises Luso-Extremaduran silicicolous outcrops in a supramediterranean, subhumid-humid, ombrotype. In addition, it also contacts topologically with the Pyrenean oak groves of Sorbo torminalis-Quercetum pyrenaicae Rivas Goday ex Rivas-Martínez 1987. We propose the name of Echinosparto iberici-Juniperetum lagunae nova (Table I, typus relevé number 4 [R144]) for this plant association. There are five other relevés fluctuating around this group A1 (R141-142, R146, R149, R151). Since they have a wide representation of the characteristic elements of the already mentioned association, we have included them in the descriptive phytosociological table of this phytocoenosis. However, given their somewhat peculiar floristic variables, they are not so clearly grouped with the other relevés of group A1. This is the case, for example, of relevé R149. As can be seen, there are three taxa – Armeria arenaria (Pers.) Schult. subsp. segoviensis (Gand. ex Bernis) Nieto Feliner, Halimium umbellatum (L) Spach and Arenaria querioides Pourr. ex Willk. – which are not present in the rest of the samples of this community. Nevertheless, the original dendrogram (Fig. 4, 2c) reveals that these fluctuating relevés are clearly included in the group which defines this type of juniper groves. In a second group (Fig. 5, A2), which has a core of thirteen relevés (R13-18, R21-27), the taxon Cytisus striatus (Hill) Rothm. subsp. eriocarpus (Boiss & Reut.) Rivas-Mart. is very frequent in all of them. This species occurs in the west of the Iberian Peninsula and in the northwest of Morocco. They are relevés dominated by the taxon Juniperus oxycedrus subsp. lagunae and have a wide distribution in the southwestern territories of the Iberian Peninsula. This formation tends to occur in small ranges with quartzitic geological materials, and mesophytic species are very common in it as a result of the subhumid-humid ombrotype. Relevés R13, R15, R21 and R23, plotted at a lower level on axis 2, show certain differences from the rest of the group due to the presence in these relevés of a cohort of species (Myrtus communis L., Rhamnus alaternus L., Rh. lycioides L., Pistacia xsaportae Burnat, Polypodium cambricum L., Ruscus aculeatus L. and Viburnum tinus L.) with a more thermic and oceanic character. If we consider the original cluster analysis (Fig. 4, 2d), it can be seen that these relevés show an affinity with this group A2, which in turn is quite independent from the rest of the juniper groves of Juniperus oxycedrus subsp. lagunae. Since species such as Teline tribracteolata (Webb) Talavera & P.E. Gibbs, Osyris quadripartita Salzm. ex Decne. and Odontites foliosa Pérez Lara are not present, for this group A2 we propose a new association: Cytiso eriocarpi-Juniperetum lagunae Pinto & Cano ass. nova (Table II, typus relevé number 2 [R14]), clearly differentiated from the juniper groves of Cytiso tribracteolati-Juniperetum oxycedri A.V. Pérez et al. in A.V. Pérez et al. 1999. Another important difference between these two associations concerns the topological contact: whereas in the new association this contact is with cork oak groves of Junipero lagunae-Quercetum suberis Rivas-Martínez et al. 2001 (Rivas-Martínez et al., 2001) occurring on more vigorous soils, the topological contact of the association Cytiso tri-

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bracteolati-Juniperetum oxycedri is with the cork oak grove of Teucrio baetici-Quercetum suberis Rivas-Martínez ex Díez Garretas, Cuenca & Asensi 1988. We must make some comments on the location of three sampling sites (R4, R9 and R12), which in the DCA-analysis appear close to this group A2 (Fig. 5). This is a group of relevés carried out in the Toledan-Taganean sector, in points close to the Serranía de San Vicente, with high abundance rates (4) of Juniperus oxycedrus subsp. lagunae. It is important to note the rare occurrence of taxa characteristic of upper syntaxonomical units (11, 12 and 5 species respectively) together with a whole cohort of species peculiar to different dynamical stages of the climatophilous vegetation of the territory (Quercus rotundifolia Lam., Daphne gnidium L., Cytisus scoparius (L.) Link subsp. scoparius, Retama sphaerocarpa (L.) Boiss., Lavandula stoechas L. subsp. sampaiana Rozeira, etc.). We estimate that these three relevés are, in fact, samples of sites in transition between the climactic holm oak wood and these edaphoxerohilous juniper groves of Juniperus oxycedrus subsp. lagunae, i. e. they are relevés which reveal the intense dynamism and dramatic changes of the vegetal cover. In the following analyses, we have opted for dismissing these relevés. A similar case is that of the relevé identified as R42 in the figure. It can be seen at the top of the figure, near axis 2. This is another relevé with very few species (9 species), two of which, Quercus rotundifolia and Olea sylvestris, have abundance rates of 2 and 3 respectively. Since the relevé is very poor in species and the climatophilous species are very obvious, we have also dismissed it in future analyses. We now have to comment on a numerous group of relevés which in the DCA-analysis lie to the left of axis 2 (Fig. 5). Taking into account the position of axis 1, the presence of two groups, respectively above and below axis 1, can be discerned. However, once they are compared with those established in the original cluster analysis (Fig. 4, 2a-2b), we observe that they do not correspond with these groups of relevés. We then opted for implementing a second DCA-analysis with this group of 30 relevés in order to clearly define their position. This data matrix includes the relevé identified as R103, which in Fig. 5 lies in between this group, to the left of axis 2, and the relevés of group A1. The reason why we opted for including this sample was precisely because of its low rate of climatophilous taxa and the occurrence of calciphilous elements, as happens in other relevés close to it in the figure. Figure 6 shows this set of relevés and reveals once again the presence of two groups, which we have labelled A3 and A4. There is still a small group of four relevés (R19-20, R122-123) which we will comment on later. Group A3 consists of fifteen relevés (R1-3, R5-8, R10-11, R101, R104-108) located on the lower part of axis 2 with a high occurrence of subrupicolous elements such as Pistacia terebinthus L. and a remarkable abundance of Cytisus scoparius (L.) Link subsp. scoparius, Lavandula stoechas (L.) subsp. sampaiana Rozeira, together with other species of a more xerophilous character, such as Asparagus acutifolius L. or Retama sphaerocarpa (L.) Boiss. All of them have the common feature of having been carried out in the more continentalised territories of the area, with a very warm climate and intense xericity in summer and below-zero temperatures and frequent frosts for months in winter. All of them have been carried out in the Toledan-Taganean biogeographical sector, i. e. under a dry-subhumid ombrotype and a mesomediterranean thermotype. We have typified this association as Pistacio terebinthi-Juniperetum lagunae Rodríguez Torres & Cano ass. nova (Table III, typus relevé number 1 [R1]). As we have said, this association occurs in the mesomediterranean dry-subhumid bioclimatic belt and it clearly differs floristically, ecologically, bio-

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Table I, Tableau I.- Echinosparto iberici-Juniperetum lagunae ass. nova (group A1, Fig. 5). Relevé reference Altitude (m) Area (m2) Coverage (%) Aspect Slope (%) Number of order

R141 951 500 60 S 20 1

R142 R143 R144 R145 R146 R147 1035 1025 1257 1257 1069 1078 1000 1000 1000 500 300 300 50 60 60 50 30 30 S S N N S S 30 25 45 40 30 30 2 3 4 5 6 7

R148 1057 400 35 N 50 8

R149 R150 R151 R152 R153 1220 976 1110 1160 1060 200 500 200 200 200 70 40 40 60 50 N S S N 30 45 25 10 9 10 11 12 13

R154 1140 200 50 NE 10 14

Character taxa of association and upper units Juniperus oxycedrus subsp. lagunae 3 Adenocarpus argyrophyllus . . Echinospartum ibericum . Quercus rotundifolia Juniperus oxycedrus subsp. oxycedrus . Phillyrea angustifolia 1 Arbutus unedo + Daphne gnidium + Thapsia villosa + Erica arborea .

2 . . . . . . . . +

3 . . + 1 . . . . .

2 + 3 + . . . . . .

1 + 2 + . . . . . .

2 . 1 . . . . . . .

2 . 1 . . . . . . .

2 2 . + . . . . . .

+ 3 . . . . . . . .

2 . . + . 1 . . . .

2 2 . . . . . . . .

2 3 . + . . . . . .

1 1 . . + . . . . .

1 + . + . . . . . .

Accompanying taxa Cistus ladanifer Dianthus lusitanus Arrhenatherum bulbosum Asphodelus albus Lavandula stoechas subsp. sampaiana Coincya longirostra Erica scoparia Astragalus lusitanicus Halimium ocymoides Urginea maritima Stipa gigantea Jasione tomentosa Erica umbellata Linaria saxatilis Calluna vulgaris Digitalis mariana Quercus pyrenaica Festuca elegans subsp. elegans Erica aragonensis Jasione mariana Arenaria querioides Halimium umbellatum Armeria arenaria subsp. segoviensis

1 . . 1 + + . . + + + 1 + + + + . . . . . . .

1 + 1 1 1 1 . 1 1 + + . . + . + . . . . . . .

. + 1 + . . . + . 1 + . + . . + 1 + . . . .

. + 1 + . . . . . . 1 + . + + . 1 1 . . . . .

+ 1 . . 1 . . . 1 . . . . . . . . . . . . . .

+ 1 . . + . + . 1 . . . . . . + . . + 1 . . .

+ 2 . . . . . . 1 . 1 . . . . . + 1 . 1 . . .

. . . . . . . . . . . . . . . . . . . . 1 2 1

+ + . . + + . . . . + + . + . + . . . . . . .

. . + . 1 . . . . . . . . . 1 . . . . . . .

+ . . . . . . . . . . + . . . + . . . . . . .

+ . . . . . . . . + . . . . . + + + + . . . .

. . . . . . . . + . . . . . . + . . . . . . .

2 . . 1 + + 2 + 1 . . . . . . . . . . . . . .

Localities: Sierra de Navalmanzano (Ciudad Real, Spain) (R141, R142, R143). Abulagoso (Ciudad Real, Spain) (R144, R145). Cuerda Navarrillo (Ciudad Real, Spain) (R146). Collado de los Gavilanes (Ciudad Real, Spain) (R147). Sierra del Rey (Ciudad Real, Spain). (R148). Monte Pelado (Toledo, Spain) (R149). Sierra Dormilleros (Ciudad Real, Spain) (R150). Sierra Quintana (Jaén, Spain) (R151, R152). Peña Malabrigo (Jaén, Spain) (R153). Cresterías Sierra Estrella (Jaén, Spain) (R154).

91

Table II, Tableau II.- Cytiso eriocarpi-Juniperetum lagunae Pinto & Cano ass. nova (group A2, Fig. 5). Relevé reference Altitude (m) Area (m2) Coverage (%) Aspect Slope (%) Number of order

R13 100 200 85 W 25 1

R14 100 200 70 . . 2

R15 R16 R17 R18 R21 R22 100 100 100 100 150 100 200 200 200 300 200 200 85 80 70 60 85 70 SW S S SW O . 25 30 30 15 25 . 3 4 5 6 7 8

R23 R24 R25 R26 R27 200 100 100 100 100 200 200 200 200 200 85 80 70 70 70 SW S S S S 25 30 30 30 20 9 10 11 12 13

Charácter taxa of association and upper units Juniperus oxycedrus subsp. lagunae Juniperus oxycedrus subsp. oxycedrus Cytisus striatus subsp. eriocarpus Erica arborea Phillyrea angustifolia Pistacia terebinthus Olea europaea var. sylvestris Pistacia lentiscus Rhamnus lycioides Myrtus communis Ruscus aculeatus Rhamnus alaternus Viburnum tinus Daphne gnidium Phillyrea latifolia Asparagus aphyllus Ruscus aculeatus Pistacia xsaportae

4 2 2 2 2 1 2 2 1 + 1 2 1 . . . . .

3 2 2 2 2 . . . . . . . . + + . . .

4 2 + 2 2 . . 1 1 + . 2 . . . . . .

4 1 + 2 1 . . . . . . . . . . . . .

4 2 1 1 2 . 1 1 . . . . . . . . . .

4 1 1 2 1 + . + . + . . . . . . . .

4 2 + 2 2 1 2 1 1 + . 2 + + . + 1 +

3 2 + 2 2 . . . + + . . + + + . . .

4 2 . 2 2 . . + 1 + . 1 . . . . . +

4 1 + 2 1 . . . . . . . . . . . . .

3 2 + 1 2 . 1 1 . . . . . . . . . .

4 2 . 1 1 . 1 + + . . + . + + . . +

4 2 1 1 1 . 1 + . + . + + + + + . .

Accompanying taxa Lavandula stoechas subsp. sampaiana Halimium viscosum Cistus salviifolius Dianthus lusitanus Arrhenatherum bulbosum Sedum hirsutum Sedum brevifolium Polypodium cambricum Cistus ladanifer Halimium ocymoides Erica australis Calluna vulgaris Adenocarpus anisochilus Simethis planifolia Conopodium capillifolium Digitalis thapsi Hypericum linarifolium Cheilanthes hispanica Asphodelus aestivus Ruta angustifolia Hyparrhenia hirta var. pubescens Gladiolus illyricus Micropyrum tenellum Sedum rubens

1 1 1 + + + + + . . . . . . . . . . . . . . . .

1 1 . 1 1 + + . + + . . + + + . . . . . . . . .

. 1 + . . . . . + 1 + + . . . . . . . . . . . .

. . . 1 . 1 1 . 1 1 + 1 . . . + + + . . . . . .

+ . . 1 + + 1 . 1 . . + . . . + . . . . . . . .

. . . + . + + . 1 1 + . . + . . . + . . . . . .

1 1 1 + . + + + . 1 . 1 + + . . . . 1 + + . . .

1 + + 1 . + + + + + . . + + + . . . 1 . . . . .

. + + . . . . . + 1 . . . + . . . + 1 . . . . .

. . . 1 . 1 1 . 1 1 + . . + . + + + . . + + + +

. . . 1 + + 1 . 1 . . . . . . + . . + . . + + .

+ + + + + + + . + . . + . . + + . + . + + + + +

+ + + + + + + . + . + + . + + + . + + + + + + 1

Localities: Nisa (Portugal) (R13, R21). Portas de Rodão (Portugal) (R14, R15, R22. R23). Zimbreira (Portugal) (R16, R24, R26). Foz do Cobrao (Portugal) (R17, R25). Castillo del Rey Bamba (Rodao, Portugal) (R18). Porto do Tejo (Portugal) (R27).

92

Table III, Tableau III.- Pistacio terebinthi-Juniperetum lagunae Rodríguez Torres & Cano ass. nova (group A3, Fig. 6). Relevé reference Altitude (m) Area (m2) Coverage (%) Aspect Slope (%) Number of order

R1 R2 R3 640 650 620 500 500 400 90 90 60 SE E S 8 8 10 1 2 3

R5 R6 450 650 500 500 100 90 N SE 10 8 4 5

R7 700 500 90 E 10 6

R8 700 500 60 S 10 7

R10 R11 R101 R104 R105 600 500 725 450 700 500 1000 500 500 500 100 50 60 70 80 N E SW S W 10 50 10 20 6 8 9 10 11 12

R106 R107R108 600 610 600 500 500 500 90 60 60 SW E S 20 15 20 13 14 15

Character taxa of association and upper units 5 Juniperus oxycedrus subsp. lagunae 5 Juniperus oxycedrus subsp. oxycedrus . . 2 2 Pistacia terebinthus Jasminum fruticans 2 1 Quercus rotundifolia + . Daphne gnidium 1 1 + + Olea europaea var. sylvestris . 1 Quercus broteroi Quercus faginea . . Quercus suber . + Osyris alba . 2 Asparagus acutifolius . .

3 . 3 . + 1 . + . . 2 1

5 + . 1 + 1 . . . . . .

5 . 2 2 + 1 + . . + . 1

5 . 2 1 . 1 + + . . 2 .

3 . 3 . + 1 . + . . 2 1

5 . . 1 + . . . . . . .

2 . . . . . . . . . 2 .

1 . 2 . 1 + . . + . 2 1

3 . + 1 + + 2 . . . . .

4 . + . 1 1 . . . . . +

2 . 2 1 1 1 + + . 1 + +

3 . 2 . 1 1 . . . + 2 .

3 . 2 1 + 1 + . . . 2 +

Accompanying taxa Cytisus scoparius subsp. scoparius Retama sphaerocarpa Thymus mastichina Rosmarinus officinalis Helianthemum appeninum Lavandula stoechas subsp. sampaiana Asplenium trichomanes Digitalis thapsi Cytisus striatus subsp. eriocarpus Arrhenatherum bulbosum Thapsia villosa Cistus ladanifer Halimium viscosum Dianthus lusitanus Urginea maritima Tamus communis Hedera helix Helichrysum serotinum Rubus ulmifolius Stipa gigantea Melica ciliata Hypericum perforatum Bryonia cretica subsp. dioica Stipa tenacissima Santolina canescens Thymus zygis Dactylis hispanica Phagnalon saxatile Helichrysum stoechas Cynosurus echinatus Umbilicus rupestris Adenocarpus telonensis Rubia peregrina Lavandula pedunculata Pyrus bourgaeana Genista hirsuta Rhamnus lycioides Santolina rosmarinifolia Lonicera implexa Thapsia maxima Lonicera periclymenum Anthirrhinum graniticum

1 1 + . . 1 . + . + 1 . . 1 + + . . + + + + . . . . . . . . . . . . . . . . . . . .

3 1 1 . . 1 . . . . . + . . . . . + . . . . + . . . . . . . . . . . . . . . . . . .

2 1 1 + 2 1 + + 1 + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

2 1 1 . . 2 + + . + + 1 . . 2 + + + . . . . . . . . . . . . . . . . . . . . . . . .

. 1 + . . 1 . + . + + . . 1 . + . . . + + + . . . . . . . . . . . . . . . . . . . .

3 1 1 . . 1 . . . . . + . . . . . + . . . . . . . . . . . . . . . . . . . . . . . .

1 1 . . . 1 . . . . + . . . . . . . . . . . + 1 2 1 . . . . . . . . . . . . . . . .

1 + . . . + . . . . . . + . . + . . . . . + . . . . . + + . . . . . 1 . . . + + 1 +

. + + . . 2 . . . . . . + . . . . . . + . . . . . . . . + . . . . . . . 2 . . . . .

2 + + . . . . . . 1 + . . . + . . . . . . + . . . . + . . . + 1 . 1 . . . . . . . .

1 + + . . + . . . . . . . . + + . . + . . + . . . . . . 1 . . . . . . . . . . . + +

2 1 . . . 2 . . . 1 + . . . . . . . + . . . . . . . 1 . + + . . + . . . . 2 . . + +

. + . . . 1 . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . 1 2 . . . . .

2 1 1 + 2 1 + + + + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

2 1 1 . . 2 . + . . + + . . . + + + . . . . . . . . . . . . . . . . . . . . . . . .

93 Briza maxima Campanula lusitanica Sedum arenarium Mercurialis annua subsp. ambigua Lupinus angustifolius Brassica barrelieri Aristolochia paucinervis Asplenium ceterach Melica magnolii Eryngium campestre Scorzonera angustifolia Sesamoides canescens Hyparrhenia hirta

. . . . . . . . . . . . .

. . . . . . . . . . . . .

. . . . . . . . . . . . .

. . . . . . . . . . . . .

. . . . . . . . . . . . .

. . . . . . . . . . . . .

. . . . . . . . . . . . .

. . . . . . . . . . . . .

. . . . . . . . . . . . .

. . . . . . . . . . . . .

+ . . . . . . . . . . . .

+ + + + + + + . . . . . .

. . . . . . . + . . . . .

+ . . . . . . . + + . . .

. . . . . . . . . . + + +

Localities: La Enebrada (Castillo de Bayuela, Spain) (R1, R2). Pico San Vicente (Hinojosa de San Vicente, Spain) (R3, R106, R108). Ctra. Navamorcuende a La Iglesuela (Toledo, Spain) (R5, R105). Castillo de Bayuela (Toledo, Spain) (R6). Cerro Garrido (Real de San Vicente, Spain). (R7). Hinojosa de San Vicente (Toledo, Spain) (R8, R107). La Iglesuela (Toledo, Spain) (R10). Ctra. Pelahustan (Toledo, Spain) (R101). Proximidades Talavera de la Reina (Toledo, Spain) (R104). Barranco de Castrejón (Puebla de Montalbán, Spain) (R11).

Fig. 6.- Detrended Correspondence Analysis (DCA) with the groups of relevés characteristic of Pistacio-Juniperetum lagunae (A3) and Stipo-Juniperetum lagunae (A4), together with the community of Phlomis purpurea - J. oxycedrus subsp. lagunae (A5). Fig. 6.- Analyse de correspondance détendancée (DCA) avec les groupes de relevés caractéristiques des Pistacio-Juniperetum lagunae (A3) et Stipo-Juniperetum lagunae (A4), ensemble avec la communauté à Phlomis purpurea - J. oxycedrus subsp. lagunae (A5).

94

climatically and biogeographically from the associations of Junipero oxycedri-Quercetum rotundifoliae Rivas-Martínez 1965 and Festuco elegantis-Juniperetum oxycedri (RivasMartínez & Sánchez Mata in Sánchez Mata 1989) Sánchez Mata 1999, since the optimum of these latter plant formations is in the supramediterranean of Carpetan-Leonese subprovince. However, in the Toledan-Taganean territories the taxon Juniperus oxycedrus subsp. lagunae penetrates into the woods of Pyro-Quercetum rotundifoliae Rivas-Martínez 1987. This is a dynamic feature of the association suggested here, which also topologically Table V.- Community of Phlomis purpurea succeeds to these holm oak woods. and Juniperus oxycedrus subsp. lagunae (group A5, Fig. 6).

The group labelled A4 (Fig. 6), composed Tableau V.- Communité à Phlomis purpurea - Juniperus oxycedrus subsp. lagunae of relevés R28-34, R73-74 and R102-103, is (groupe A5, Fig. 6). characterised by the occurrence of taxa thriRelevé reference R19 R20 R122 R123 ving on carbonated soils (Stipa tenacissima, Altitude (m) 280 300 240 225 Thymus zygis, Staehelina dubia, Ruta chale- Area (m2) 500 500 150 200 60 60 80 80 pensis). This basophilous flora shares its Coverage (%) Aspect NW SW NE NE biotope with other more or less silicicolous Slope (%) 40 25 10 12 taxa, such as Stipa gigantea, Lupinus angus- Number of order 1 2 3 4 tifolius, Cistus ladanifer, Genista hirsuta. Character taxa of the community and upper units These samples were taken in the province of Juniperus oxycedrus subsp. lagunae 4 4 3 4 Toledo and in the north of the province of Pistacia terebinthus 1 2 + + 2 2 2 1 Ciudad Real (Spain). For this community Olea europaea var. sylvestris 1 2 . . we propose the name Stipo tenacissimae- Phlomis purpurea Jasminum fruticans 1 1 1 1 Juniperetum lagunae ass. nova (Table IV, Pistacia lentiscus 1 1 2 2 typus inv. number 10 [R102]). It is a neutro- Myrtus communis 1 1 1 1 1 1 . . basophilous association of eastern Toledan- Asparagus albus aphyllus 1 1 . . Taganean distribution and it occurs in the Asparagus Aristolochia baetica 1 1 . . mesomediterranean thermotype in a dry Rhamnus lycioides + . . . 1 . . . ombrotype. As we have said, the fundamen- Phillyrea latifolia . . + . tal floristic difference of this association is Smilax aspera the co-existence in its relevés of yet mentio- Accompanying taxa ned basophilous species with another set of Cistus albidus + + + + 1 . . . acidophilous species ones. This community Cistus ladanifer gnidium + . . . topologically succeeds to Pyro bourgaea- Daphne Thymus mastichina + . . . nae-Quercetum rotundifoliae in its basophi- Rosmarinus officinalis + + . . Cytisus scoparius subsp. scoparius 1 2 . . lous facies. Micromeria graeca 1 + . + It remains for us to comment on the posi- Rhamnus alaternus 1 . . . tion of a little group of samples (Fig. 6, A5) Flueggea tinctoria + . . . + . . . (R19-20, R122 and R123), all of them carried Digitalis mariana 1 . . . out in sites of intense thermicity (upper ther- Dianthus crassipes Teucrium fruticans . + . . momediterranean and lower mesomediterra- Lavandula stoechas subsp. sampaiana . + . . nean thermotypes), with practically no Dactylis hispanica . . + + . . + + winter frosts and high summer temperatures Umbilicus rupestris Cheilanthes maderensis . . + + (Sierra Norte de Sevilla, Sierra Morena, Bryonia cretica subsp. dioica . . + + Spain). The high abundance rates of thermo- Selaginella denticulata . . + + . . + + philous species such as Olea europaea (L.) Sanguisorba minor var. sylvestris Brot., Jasminum fruticans L., Localities: Garganta del río Viar (Sevilla, Spain) (R19, R20). Phlomis purpurea L., Asparagus aphyllus L. Río Viar (Sevilla, Spain) (R122, R123).

95

or Pistacia lentiscus L. and the position, far away from the two groups described in the DCA-analysis, suggest that these relevés could be considered as an independent association. However, in the absence of more samples from other sites, we prefer to consider it as a plant community of Phlomis purpurea and Juniperus oxycedrus subsp. lagunae (Table V). Finally, we ascribe this group of phytocoenoses to an upper syntaxonomical level, as suggested in the International Code of Phytosociological Nomenclature (Weber et al., 2000). Thus, we propose the alliance Juniperion oxycedro-lagunae all. nov. whose typus is the association Pistacio terebinthi-Juniperetum lagunae (Art. 17). This alliance comprises all acidophilous and neutro-basophilous plant formations dominated by the taxon Juniperus oxycedrus subsp. lagunae, with an outstanding distribution in the LusoExtremaduran subprovince under thermotypes from upper thermomediterranean to supramediterranean, and with an ombrotype which is at least subhumid. This alliance would then be included in the Pistacio-Rhamnetalia alaterni order, as part of the Quercetea ilicis class. As characteristic species of the alliance let us mention: Juniperus oxycedrus subsp. lagunae, J. oxycedrus subsp. oxycedrus, Pistacia terebinthus and Echinospartum ibericum. IV. CONCLUSIONS The permanent character of the edaphoxerophilous communities of Juniperus oxycedrus subsp. lagunae, which are restricted to areas where the dominant ecological factor is the xericity of substrate, is a clear feature of all the territories in the centre and south of the Iberian Peninsula (Spain and Portugal). This feature makes them clearly distinguishable from the rest of the juniper groves dominated by Juniperus oxycedrus subsp. oxycedrus and other climactic plant formations in the territory. This fact justifies the creation of a new alliance, namely, Juniperion oxycedro-lagunae, with a wide distribution in the siliceous and neutro-basic substrates of the Luso-Extremaduran biogeographical subprovince, to which four new associations and one community are described. Syntaxonomical scheme QUERCETEA ILICIS Br.-Bl. ex A. & O. Bolòs 1950 O. Pistacio lentisci-Rhamnetalia alaterni Rivas-Martínez 1975 All. Juniperion oxycedro-lagunae all. nov. Ass. Pistacio terebinthi-Juniperetum lagunae Rodríguez Torres & Cano ass. nova Ass. Stipo tenacissimae-Juniperetum lagunae ass. nova Ass. Echinosparto iberici-Juniperetum lagunae ass. nova Ass. Cytiso eriocarpi-Juniperetum lagunae Pinto & Cano ass. nova Community of Phlomis purpurea - Juniperus oxycedrus subsp. lagunae

96

Table IV, Tableau IV.- Stipo tenacissimae-Juniperetum lagunae ass. nova (group A4, Fig. 6). Relevé reference Altitude (m) Area (m2) Coverage (%) Aspect Slope (%) Number of order

R28 940 500 50 S 30 1

R29 950 500 80 NW 20 2

R30 940 500 90 N 30 3

R31 600 500 70 W 40 4

R32 860 500 70 N 20 5

Character taxa of association and upper units 2 Juniperus oxycedrus subsp. lagunae Stipa tenacissima . 1 Quercus rotundifolia Scilla autumnalis 1 Quercus coccifera . . Pistacia terebinthus . Rhamnus lycioides Phillyrea angustifolia . Phillyrea latifolia . Olea europaea var. sylvestris .

3 1 3 . . . . . . .

4 . 3 . . . . . . .

3 . 1 . + . . . . .

3 1 1 . + . . . . .

3 1 + . 1 . . . . .

Accompanying taxa Retama sphaerocarpa Dactylis hispanica Dianthus lusitanus Avenula sulcata Cistus albidus Cistus salvifolius Helichrysum stoechas Stipa gigantea Phagnalon saxatile Umbilicus rupestris Sedum brevifolium Thymus zygis Cynosurus echinatus Crocus nudiflorus Asphodelus aestivus Halimium umbellatum subsp. viscosum Daphne gnidium Thymus mastichina Agrostis castellana Staehelina dubia Teucrium gnaphalodes Lavandula stoechas subsp. sampaiana Teucrium pseudochamaepytis Phlomis lychnitis Thapsia villosa Gastridium ventricosum Asparagus acutifolius Rosa canina Ballota nigra Ruta chalepensis Osyris alba Urginea maritima Dorycnium pentaphyllum Santolina rosmarinifolia Ruta montana Carex hallerana Ceterach officinarum Hyparrhenia hirta Cheilanthes tinaei Cistus ladanifer Adenocarpus telonensis Cytisus multiflorus Cytisus striatus subsp. eriocarpus

+ + + . 1 2 + 1 + . . 1 + + + 1 . . . . . . . . . . . . . . . . . . . . . . . . . . .

. 1 + + 2 2 . + + + . + . + + . + . . . . . . . . . . . . . . . . . . . . . . . . . .

2 . . . . . 1 + . . . 1 . . + . + + . . . . . . . . . . . . . . . . . . . . . . . . .

. 1 . + . . + . . . . 1 . . 1 . + 1 + + + 1 + + . . . . . . . . . . . . . . . . . . .

. + . . .

+ 1 1 + + + 1 + 1 + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

R33 R34 860 820 500 500 70 100 SE NE 30 20 6 7

1 . . . . + . . + . . . + . + 1 + + + + + + + + . . . . . . . . . . . . .

R73 780 500 95 SW 30 8

R74 820 500 95 SW 25 9

4 + + . 1 . . . . .

4 . + . 4 . 1 . . +

4 . 2 . 2 + + 1 . .

4 1 1 . + . . . . .

5 + . . + . . 1 . .

. + . . 3 . + . . . . . . . + + + . . . . 1 1 + . . + . . . . . . . . . . . . . . . .

+ . . . . + 1 . 1 + . + . . + . + + + 1 . + 1 + . . + . . + + + + + + 1 + + + 1 1 1 .

. + . .

+ + 1 . . . . . . . . . . . . . . + . . . + + . . . . . . . + 1 . . . . . . . 2 . . .

. . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . + . .

1 . . . + . . . . . . . . . . + . . . . . 1 . . . + + . . . . + . + . 1 . 1

R102 R103 800 875 500 500 85 100 SE SW 25 20 10 11

97 Genista hirsuta Cheilanthes maderensis Rosmarinus officinalis Samolus valerandi Helichrysum italicum subsp. serotinum Sedum sediforme Arrhenatherum bulbosum Asphodelus albus Erica scoparia Lavandula stoechas subsp. luisieri

. . . . . . . . . .

. . . . . . . . . .

. . . . . . . . . .

. . . . . . . . . .

. . . . . . . . . .

. . . . . . . . . .

. . . . . . . . . .

. . . . . . . . . .

1 + . . . . . . . .

. . 2 + + + + + . .

. . + . . . . + + +

Localities: Sierra de los Yébenes (Toledo, Spain) (R28, R29, R30). Subida a Las Nieves (Toledo, Spain) (R31). Mora de Toledo (Toledo, Spain) (R32, R33, R34). Puerto Lápice (Ciudad Real, Spain) (R73, R74). Marjaliza (Toledo, Spain) (R102, R103).

Appendix 1. Authors and sites of the bibliographical relevés used. Appendix 1. Auteurs et sources des relevés utilisés. Phytocoenoses

Relevé references

Bibliographic reference

Echinosparto iberici-Juniperetum lagunae R141-154 Made by the authors, table I Made by the authors, table II Cytiso eriocarpi-Juniperetum lagunae R13-18, R21-27 Made by the authors, table III Pistacio terebinthi-Juniperetum lagunae R1-3, R5-8, R10-11, R101, R104-108 Made by the authors, table IV Stipo tenacissimae-Juniperetum lagunae R28-34, R73-74, R102-103 Community of Phlomis purpurea Made by the authors, table V Juniperus oxycedrus subsp. lagunae R19-20 Community of Phlomis purpurea Delgado (2001), p. 70, table 14, 2 relevés Juniperus oxycedrus subsp. lagunae R122-123 Pérez Latorre et al. (1999), p. 153, table 5, 3 relevés Cytiso tribracteolati-Juniperetum oxycedri R35-37 Sánchez Mata (1989), p. 308, table 111, 4 relevés Festuco elegantis-Juniperetum oxycedri R38-41 Cano (1988), table 49, relevés 1-6 Pyro bourgaeanae-Quercetum rotundifoliae R43-48 Pyro bourgaeanae-Quercetum rotundifoliae Cano (1988), table 49, relevés 7-10 quercetosum fagineae R49-52 Pyro bourgaeanae-Quercetum rotundifoliae Cano (1988), table 49, relevés 11-13 myrtetosum communis R53-55 Cano (1988), table 48, 9 relevés Poterio agrimonioidis-Quercetum suberis R56-64 Penas (1980), 8 relevés Junipero oxycedri-Quercetum rotundifoliae R65-72 Laorga (1986), table 100, 7 relevés Hyacinthoido hispanicae-Quercetum cocciferae R75-81 Melendo (1998), p. 436, table XXIV. 2, relevés 1-2, 5-9 Pyro bourgaeanae-Quercetum rotundifoliae R82-83, R84-88 Pyro bourgaeanae-Quercetum rotundifoliae Melendo (1998), p. 436, table XXIV. 2, relevés 20-31 myrtetosum communis R89-100 Pyro bourgaeanae -Quercetum rotundifoliae García Cruz (1999), p.315, table 23.1.3, relevés 1-5 myrtetosum communis R109-113 Pyro bourgaeanae -Quercetum rotundifoliae García Cruz (1999), p.315, table 23.1.3, relevés 6-8 quercetosum fagineae R114-116 Belmonte (1986), table 74, relevés 1-4 Pyro bourgaeanae -Quercetum rotundifoliae R117-120 Ruiz Téllez (1986), table 75, 1 relevé Junipero oxycedri-Quercetum rotundifoliae R121 Pyro bourgaeanae -Quercetum rotundifoliae Delgado (2001), p. 52, table 8, relevé 1 myrtetosum communis R124 Delgado (2001), p. 52, table 8, relevés 2-3 Pyro bourgaeanae -Quercetum rotundifoliae R125-126 Pyro bourgaeanae -Quercetum rotundifoliae Delgado (2001), p. 52, table 8, relevés 4-6 quercetosum suberis R127-129 Sánchez Pascual (1994), table 81-A, 6 relevés Pyro bourgaeanae -Quercetum rotundifoliae R130-135 Amor et al. (1993), p. 120, table 14, 5 relevés Pyro bourgaeanae -Quercetum rotundifoliae R136-140 Without syntaxonomic position (Made by the authors)

R4, R9 R12 R42

Ladera norte del Oso (Real de S. Vicente, Toledo, Spain) Barranco de Castrejón (La Puebla de Montalbán, Toledo, Spain) Navalasno del Conde (Sierra Morena, Jaén, Spain)

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