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Patterns of community and species diversity in grassland vegetation of the southwestern Iberian System (Spain)/Patrones de diversidad fitocenotica y floristica en los pastizales del Sistema Iberico suroccidental.

INTRODUCTION

Plant diversity patterns in grasslands have been for a long time a question of attention for management and conservation purposes. Several studies have dealt with the soil pH, altitude, topography (aspect and slope), and grazing intensity, as the main local abiotic factors influencing the variation of the plant species composition and species richness (CRICHTLEY & al., 2002; AMEZAGA & al., 2004; Bennie & al., 2006; KLIMEK & al., 2007; MARINI & al., 2007; JIMENEZ-ALFARO & al., 2013). Topography has a strong effect on local hydrology and additionally affects patterns in local temperature and irradiation, as well as, local exposure to wind (MOESLUND & al., 2013); thus, it is considered an indirect factor for shaping local grassland vegetation patterns. Mechanisms controlling plant species richness in grasslands in Europe have been explained by the positive effect of slope angle (MAURER & al., 2006; KLIMEK & al., 2007; MARINI & al., 2007), and the negative effect of soil acidification and altitude (PIQUERAY & al., 2007; KOPEC & al., 2010). Over the worldwide, the soil pH pattern may be different across floristic regions because the direction of the relationship between richness and pH depends on whether the species pool has its evolutionary origin on soils of high or low pH (PARTEL, 2002). In Europe, it may be also considered that the higher contemporary calciphile to acidiphile richness ratio corresponds to a disproportionate decline of acidic habitats and their flora during the ice age times (EWALD, 2003), which would also explain that the majority of the central European flora is restricted to base rich, calcareous soils.

The diversity of grasslands in the southwestern Iberian Mountain System s.l. has been reported in some traditional phytosociological studies (LOPEZ GONZALEZ, 1978; BARRERA, 1985; RODRIGUEZ-ROJO & al., 2010, 2011, 2012). They pointed out important levels of community diversity and originality in this territory, as a consequence of a varied lithology, including calcareous and siliceous substrates (limestone, dolomites, slates, quartzites, red sandstones, albian sands, etc), combined with a complex relief that also appear in near mountains (RIVAS GODAY & BORJA, 1961; GAVILAN & al., 2012). Some mesic and dry grasslands are included in the Catalogue of Habitats of Special Protection in Castilla-La Mancha (Nature Conservation Law 9/1999) or in the Annex I of Habitats Directive (92/43/EC): semi-dry calcareous grasslands (Festuco-Brometea), Nardus stricta swards (species-rich Nardus grasslands), cryoturbated calcareous grasslands (alpine and subalpine calcareous grasslands). The cryoturbated calcareous grasslands colonize the ridges and slopes of the highlands, mesic grasslands fill the valley bottoms and basins, while semi-dry grasslands occupy a transitional range among both extremes but their topographic affinity has not been properly defined (RODRIGUEZ-ROJO & al., 2010).

Because of the varied lithology combined with the complexity of relief given in this territory, and under the current moderate grazing intensity and no mowing practices, we hypothesize that the main floristic gradients in the mesic and dry grasslands should be related to lithological and topographic factors, and that these can serve as good predictors of species composition. According to this, our study develops an application of numerical and indicator species analyses on dry and mesic grasslands in the southwestern Iberian System in order to give responses to the following issues: (1) to determine the main vegetation types of these grasslands, (2) to compare the resulting classification with that used traditionally by previous authors, (3) to determine the diagnostic species of each vegetation type, and (4) to assess how far topographical (slope, aspect, rock cover and altitude) and lithological factors may discriminate the community diversity and the patterns of species richness in these grasslands.

MATERIAL AND METHODS

STUDY AREA

The main rangelands prospected were Sierra de Albarracin, Sierra de Valdemeca and Serrania de Cuenca, which according to the geomorphological sectorization of GUTIERREZ & PENA (1994) belong to the central sector and southwestern area of the Iberian System (Cordillera Iberica) (Figure 1). The highest altitude is registered in Caimodorro peak (1920 m asl). According to the bioclimatic typology of RIVAS-MARTINEZ (2007), the territory is in the transition between the macrobioclimates Mediterranean, in the lowlands, and Temperate (submediterranean variant) in the uplands (Figure 2). The bioclimatic belts of the study area correspond to the upper supramediterranean, upper suprasubmediterranean and lower orosubmediterranean, with ombrotypes ranging from upper subhumid to upper humid. Biogeographically, the territory is framed into the Maestracensean and Celtiberian-Alcarrean sectors, Mediterranean Central Iberian province (RIVAS-MARTINEZ, 2011).

DATA SAMPLING

Field sampling was carried out from 2009 to 2012 during summer (mid-June to late-July). We used orthoimages, forestry maps, topographic maps and geologic maps to locate a network of grassland areas of a certain extension that were also representative of the whole territory in terms of the geological and altitudinal variation. The distribution of the sampled sites in the selected areas was designed to cover the lithological, topographical and altitudinal variation in every one of them. Most of the areas were located in the Serrania de Cuenca and Alto Tajo (Cuenca and Guadalajara provinces), while Sierra de Albarracin (Teruel) was prospected in order to complete the representation of siliceous grasslands (Figure 1). Plot size was fixed as a rule in 10x10 m2. Plant cover indices were visually estimated in percentage values (0.1, 1, 5, 10, 20, 30, 40, 50, 60, 70, 80, 90 and 100%). Altitude, slope, exposure, rock cover and bare soil cover percentages were recorded in each plot, as well as, other supplementary data (total plant cover, livestock type, grazing system, etc.). Taxonomic nomenclature follows the proposals of Flora iberica (CASTROVIEJO & al., 1986-2012) except for Compositae and Poaceae taxa that follow the Euro+Med PlantBase (Euro+Med, 2006-2013 ww2.bgbm.org/EuroPlusMed/[accessed 4-4-2013]).

NUMERICAL ANALYSES

Several types of agglomerative and divisive classification were applied to the final dataset of 160 releves and 408 species, including the beta-flexible linkage method (beta = -0.25) with Sorensen distance, Ward's method with relative Euclidean, and TWINSPAN modified according to ROLECEK & al. (2009) with three pseudo-species cut-off levels (0%, 5% and 25%) and total inertia as a heterogeneity measure. Percentage cover values were log-transformed in order to reduce the importance of dominant species (VAN DER MAAREL, 1979), excepting for the application of TWINSPAN algorithm that uses pseudo-species cut-off levels. After examining several partitions of the data set, we selected the partition with beta-flexible clustering that produced the less-chained dendrogram and more homogeneous clusters with respect to the ordination (DCA) of the same dataset. The number of clusters accepted corresponds to the alliances traditionally recognized in the literature. Nevertheless, the crispness of classification was checked using the method suggested by BOTTA-DUKAT & al. (2005) that revealed that crispness is higher at the level of four clusters and decreases gradually when the number of clusters increased. Centroids of each cluster were calculated using the Euclidean distance and 5 releves were reassigned according to their relationship to the nearest cluster centroid. The cluster membership of these 5 releves was also checked in the DCA scatter plot.

For the calculation of diagnostic species values, the size of the clusters was standardized to equal size of 15% of the data set (TICHY & CHYTRY, 2006). We only show the diagnostic species with values of the [PHI]-coefficient of association [greater than or equal to] 0.25 for at least one cluster and with a statistically significant affinity at the probability level <0.001 according to Fisher's exact test.

Using canonical correspondence analysis (CCA), the relationships between altitude, south exposure, west exposure, slope, rock cover and altitude with species composition of the vegetation data were tested. Previously, all these environmental variables were passively projected onto the DCA diagram. The power of each explanatory variable was also tested with a Monte Carlo permutation test (reduced model, 999 permutations) and a forward manual selection procedure was used to select the most important variables. Differences in altitude among particular vegetation types were tested with the Tukey post-hoc test following one-way ANOVA, while differences in slope and rock cover were tested with Kruskal-Wallis ANOVA by Ranks applied for variables with a non-gaussian distribution. An attribute plot showing changes in species richness along the first two DCA axes was constructed applying a Generalized Additive Model (GAM), using Poisson distribution and testing for nonlinearity.

Numerical analyses were performed with JUICE 7.0 software (TICHY, 2002) combined with PC-ORD v.5 (MCCUNE & MEFFORD, 1999) for Cluster Analysis, and CANOCO 4.5 package (TER BRAAK & SMILAUER, 2002) for DCA and CCA.

RESULTS

VEGETATION CLASSIFICATION AND PATTERNS

Ten clusters were considered in the dendrogram that best fit with the classification of the releves at the association level (Figure 3). However, maximum crispness occurred at the cutting level of the following four groups: A) mesic grasslands (clusters 1-4), B) semi-dry calcareous grasslands (clusters 5-7), C) dry calcareous grasslands (clusters 8-9), and D) dry siliceous grasslands (cluster 10). Frequencies of the diagnostic species in the ten clusters are summarized in Table 1.

The DCA diagram of Figure 4 shows the floristic relationships between these clusters. As gradient length of the first DCA axis was 6.24 standard deviation units, the use of an unimodal ordination is quite appropriate. The first DCA axis was positively correlated with slope and rock cover, and negatively with plant cover. Dry calcareous grasslands were positioned in the positive part of this axis, while mesic grasslands were in the negative part. The second DCA axis showed a positive correlation with altitude and dry siliceous grasslands were positioned in its positive part. South and west components of aspect had a very low correlation with first and second DCA axes. According to the Monte Carlo permutation test, only rock cover, altitude and slope were significant in CCA as explanatory variables of the variation of species composition, corresponding to the former the stronger relationship.

Next, a brief description of the vegetation types inferred from the classification, their diagnostic species and the related syntaxa is given (see also the syntaxonomical scheme included after the section of Discussion).

Group A is very heterogeneous and comprises wet meadows and mesic grasslands growing on well-developed soils, usually, at the bottom of valleys and with a 100% plant cover. Cluster 1 includes the southern Oroiberian intensively grazed grasslands of Phleo nodosi-Cynosuretum cristati (RODRIGUEZ-ROJO & al., in press.) characterized by Lolium perenne, Trifolium dubium, Bromus hordeaceus, Phleum nodosum, Trisetum flavescens, Trifolium striatum, T. campestre, etc. Cluster 2 includes grasslands growing on siliceous soils moderately wet in spring and characterized by Agrostis castellana, Jasione montana and J. crispa subsp. sessiliflora. Cluster 3 corresponds to Nardus stricta swards with Genista anglica (Genisto anglicae-Nardetum strictae) that grow in deep, siliceous and hydromorphic soils, characterized by a high number of diagnostic species such as Genista anglica, Nardus stricta, Anthoxanthum odoratum, Juncus squarrosus, Potentilla erecta, Luzula campestris, Carex leporina, etc. Finally, cluster 4 includes pastures growing on temporarily wet, base-rich and clayey soils, characterized by Deschampsia cespitosa subsp. subtriflora, Leontodon hispidus, Scorzoneroides carpetana, Carex caryophyllea, Centaurea jacea, Prunella hyssopifolia, etc, and framed into the Sanguisorbo lateriflorae-Deschampsietum refractae. Although the presence of Trifolium repens and Bellis perennis indicates floristic similarities with Cynosurus cristatus grasslands, as well as, intensive grazing, Sanguisorbo-Deschampsietum is also characterized by Plantago media and P. maritima subsp. serpentina, indicators of the calcareous nature of soils.

Group B represents the typical semi-dry calcareous grasslands of the region (Festuco-Brometea). Cluster 6 includes the Oroiberian calcareous grasslands adscribed to Cirsio microcephali-Onobrychidetum hispanicae characterized by Bromopsis erecta, Thymus pulegioides, Trifolium ochroleucon, Salvia pratensis, etc; while cluster 7 is related to the Brachypodium phoenicoides communities growing in valley bottoms at lower altitudes, which should be framed into the Maestracensean Festuco andresmolinae-Brachypodietum phoenicoidis. Cluster 5 represents a transition between the semi-dry grasslands (Cirsio-Onobrychidetum hispanicae) and the dry pastures from cryoturbated soils (Festucetum hystricis) (Figure 4). Diagnostic species of this cluster are Coronilla minima, Jasonia tuberosa, Carthamus carduncellus, Polygala monspeliaca, Onobrychis argentea subsp. hispanica, Ononis cristata, etc.; among the diagnostic species shared with Cirsio-Onobrychidetum are Cirsium acaule, Helictochloa pratensis subsp. iberica and Galium estebanii var. leioclados. Festuca hystrix and Thymus bracteatus denote the xerophytic character of this type of grasslands and its relationship with cluster 8. Plantago maritima subsp. serpentina is characteristic of clayey or loamy soils, lightly humid and basic to saline (Pedrol, 2009), and it denotes the typical substrates associated to these grasslands. The slope arrow in the DCA diagram points towards this cluster, and the ANOVA test showed significant differences in slope with respect to cluster 6 (semi-dry grasslands) but not to cluster 8 (dry grasslands) (Figure 5).

Group C contains the most typical Oroiberian short and sparse grasslands proper to upfrozen soils and framed into the Sideritido fontquerianae-Arenarion microphyllae. Cluster 8 corresponds to the Festucetum hystricis, characterized by Thymus bracteatus, Anthyllis vulneraria subsp. vulnerarioides, Poa ligulata, Carduus assoi, Dianthus pungens subsp. brachyanthus, Armeria alliacea subsp. matritensis, etc. Cluster 9 corresponds to the Paronychio capitatae-Artemisietum lanatae, mainly characterized by Artemisia pedemontana. The arrow corresponding to rock cover points towards these two clusters in the DCA diagram (Figure 4). According to the ANOVA test, rock cover is significantly higher in these two clusters, excepting for cluster 5 (transitional to semi-dry grasslands) (Figure 5). The xerophytic character of this group is reflected by the high number of annuals among its diagnostic species (Xeranthemum inapertum, Androsace maxima, Alyssum simplex, Bombycilaena erecta, etc). Richness in Iberian endemics is also correlated with rock cover (Figure 4). The endemic diagnostic species for group C are Arenaria erinacea, Arenaria obtusiflora subsp. ciliaris, Armeria alliacea subsp. matritensis, Carduus assoi, Centaurea paniculata subsp. castellana and Thymus bracteatus. Altitude is significantly higher in Festucetum hystricis than in Paronychio-Artemisietum lanatae (Figure 5).

Group D only includes the xerophile thyme-grasslands from siliceous bedrock soils (slates, sandstones and quartzites). Their floristic combination is characterized by the diagnostic species Thymus leptophyllus subsp. izcoi, Festuca rivas-martinezii, Plantago holosteum, Pilosella castellana and Koeleria crassipes, besides Rumex acetosella subsp. angiocarpus and several silicicolous therophytes (Aira caryophyllea, Evax carpetana, Scleranthus polycarpos, Logfia minima, etc). The high cover of Iberian endemics is due to the abundance of the Oroiberian Thymus leptophyllus subsp. izcoi and the Iberian Festuca rivas-martinezii. This vegetation type has been recently described (Fumano procumbentis-Thymetum izcoi G. Navarro; RODRIGUEZ-ROJO & al., 2012).

SPECIES RICHNESS

Trends in species richness along the major floristic gradients are not linear. The GAM model shows skewed unimodal trends along the two DCA axes, the decrease in species richness being more evident along the first axis (Fig. 6). Species density ranges from a minimum value of 15 species per 100 [m.sup.2], that corresponds to the clusters 5 and 9, and a maximum of 51, that corresponds also to the cluster 9. The average of the species density ranges from 25 to 35 species, and the values around 30 (28-32) are the most frequent. The higher values appear around the center of the DCA diagram and are not associated to any particular cluster. Lower species richness values are associated to both extremes of the first DCA axis, corresponding to the Nardus stricta swards in the negative extreme, and the most xerophytic grasslands in the positive extreme. On the contrary, the number of Iberian endemics increases towards the positive part of the first DCA axis where calcareous dry grasslands are positioned (cluster 8). The cumulative number of Iberian endemics for this vegetation type is equal to 19 in total (see list of Iberian endemics in Appendix 1).

DISCUSSION

Topography and lithology are the main factors influencing the vegetation diversity of grasslands in the study area. Rock cover and slope are associated to a gradient of water availability that determine the major floristic differences: the wet and mesic meadows (Molinio-Arrhenatheretea and Nardetea) are positioned in flat terrains at the valley bottoms with null rock cover, while the dry grasslands (Festuco-Ononidetea and Festucetea indigestae) show the higher values for both variables. Floristic differences linked to lithological variability (calcareous vs. siliceous substrates) were larger in dry grasslands than in mesic grasslands. Siliceous thyme-grasslands of the Fumano-Thymetum izcoi appear well differentiated against calcareous dry grasslands by a large pool of diagnostic species from Festucetea indigestae and Tuberarietalia. The lithological gradient is narrower in the mesic grasslands, opposing the associations Phleo-Cynosuretum cristati (cluster 1), from more or less decalcified soils, to Sanguisorbo-Deschampsietum hispanicae (cluster 4) from base-rich soils. However, more analysis would be needed involving other environmental variables, such as soil properties or grazing intensity, for the interpretation of the community diversity of mesic meadows.

Differences in altitude appeared among communities belonging to the same major vegetation types. For instance, although the two types of semi-dry grasslands (Festuco-Brometea) were characterized by few diagnostic species and their clusters were linked at the lowest dissimilarity level in the dendrogram, they showed altitudinal differences: Cirsio-Onobrychidetum hispanicae occurs at higher altitudes, in the upper suprasubmediterranean belt (altitudes: 1300-1600 m) whereas Festuco-Brachypodietum phoenicoidis at lower altitudes (altitudes: 1100-1500 m). However, only a community analysis of the Festuco-Brometea grasslands in all the southern Iberian System with more phytosociological data would bring out a right ecological interpretation of the environmental patterns in this grassland type. It is also interesting the relation between siliceous thyme-grasslands and altitude in the DCA diagram, although altitude is not significantly higher compared with the rest of the grassland groups. These thyme-grasslands have a wide altitudinal range, reaching the highest belts of the mountains crowned by Paleozoic quartzites (Sierra de Albarracin, Sierra Menara and Sierra de Valdemeca), above 1800 m.

The classification results were explained according to the flexible beta method selected, but the performance of the three applied clustering methods allowed us to compare differences in the linkage of some intermediate vegetation types, as it is the case of the semi-dry grasslands (group B) with moisture requirements intermediate between mesic and dry grasslands. Moreover, in combination with the ordination analysis, it is possible to reinterpret the similarity level between clusters separated in the dendrogram. For instance, cluster 5 was linked in the Ward dendrogram to the cluster node of the typical dry calcareous grasslands (Sideritido-Arenarion). But in the divisive TWINSPAN method, cluster 5 showed more affinity with the typical semidry grasslands (Cirsio-Onobrychidetum), and even their respective centroids were much closer in the DCA scatter plot where the two groups do not appear well separated. Cluster 5 has been treated here as a transitional community between Cirsio-Onobrychidetum and Festucetum hystricis and its syntaxonomical position cannot be clarified only upon the cluster dendrogram, but also upon the diagnostic species analysis and the information of the topographic variables. Onobrychis argentea subsp. hispanica, Ononis cristata, Leucanthemum pallens, Medicago sativa and Cirsium acaule may lead to interpret this type of vegetation as belonging to the Cirsio-Onobrychidetum hispanicae, but Festuca hystrix and Thymus bracteatus are also diagnostic and are even more abundant (cover values from 20 to 40%). On the other hand, the ANOVA test for the rock cover and slope variables showed that cluster 5 does not differ from cluster 8 (Festucetum hystricis) while it does from cluster 6 (Cirsio-Onobrychidetum). RIVAS-GODAY & BORJA (1961) described in Sierra de Gudar (eastern Iberian System), a submediterranean subassociation for Cirsio-Onobrychidetum characterized by Koeleria vallesiana, Carthamus carduncellus and Linum salsoloides, that would be the most frequent subassociation in our study territory. Cirsio-Onobrychidetum is limited to shady places in closed valleys, at the bottoms or hillsides, usually under the canopy of Pinus sylvestris forests (LOPEZ GONZALEZ, 1978), where the warm temperatures of summer are attenuated. In open exposed situations, in deforested valleys and steep sites, the grasslands become more xerophytic, and if the grazing intensity is high, as it usually occurs in the surroundings of rural villages, xericity is more stressed as a consequence of the effect of the erosion of the superficial soil with the bedrock coming to the surface. Only dry grasslands may colonize these exposed and topographic sites that are related to the Festucetum hystricis. However, in clay and loamy soils, a variant in transition to semidry grasslands may occur with the presence of Jasonia tuberosa, Plantago maritima subsp. serpentina, Cirsium acaule, Helictochloa pratensis subsp. iberica and Galium estebanii var. leioclados, and cluster 5 represents this vegetation type (Table 2). The presence of Jasonia tuberosa and Plantago maritima subsp. serpentina in this community also denotes a similarity with the most xerophilous grasslands of the Deschampsion mediae alliance (Plantagini serpentinae-Jasonietum tuberosae O. Bolos & Masalles in O. Bolos 1983; Prunello hyssopifoliae-Plantaginetum serpentinae Biurrun 1999; Eryngio dilatati-Jasonietum tuberosae Torres & Cano 2000) (TORRES & al., 2000). On the other hand, although Jasonia tuberosa is not present in our samples from the Sanguisorbo-Deschampsietum grasslands, LOPEZ-GONZALEZ (1978) did include it in his table of the association. The ecological factor that is associated to this floristical similarity is explained by some edaphic and lithological affinities: compact and base-rich, clayed soils.

The number of species recorded in the mountain grasslands of the southwestern Iberian System is analogous to that referred in previous studies in the Western European mountains (KOPEC & al., 2010). The DCA diagram showed that an increase in species richness is not skewed towards any of the vegetation types. According to our results, the grassland communities recorded do not differ significantly in species richness at the spatial grain of 100 [m.sup.2]. At first, it was expected that it was higher in the temperate semi-dry calcareous grasslands (Cirsio-Onobrychidetum hispanicae), reported as one of the plant communities with the world record for plant species richness (Wilson & al., 2012), but the submediterranean context of the Iberian System is far from the optimum of these temperate grasslands in central Europe. If species richness in grasslands from more hydromorphic soils is reduced by competitive exclusion of the high abundances of few species, and is also reduced in more xerophytic grasslands by hydric stress, then mesic or semi-dry grasslands would contain more species richness. This tendency may be interpreted in the species richness plot based on the fitted additive model (Fig. 6). On the other hand, the negative effect of soil acidification on plant species richness observed by previous authors (PARTEL, 2002; PIQUERAY & al., 2007; KOPEC & al., 2010) is not evident in the southern Oroiberian grasslands, maybe as a consequence of the Mediterranean character of the siliceous thyme-grasslands that are locally rich in annual species.

As a conclusion, the evaluation of the floristic value of the semi-natural grasslands in the southwestern Iberian System should take into account other criteria such as the endangered status, the level of rarity and the endemicity of the species composition. According to the latter, the Sideritido-Arenarion aggregatae grasslands, included in the Annex I of Habitats Directive, represent one of the most valuable vegetation types of this area.

SYNTAXONOMICAL SCHEME OF THE GRASSLANDS IN THE SOUTHWESTERN
IBERIAN SYSTEM

MOLINIO-ARRHENATHERETEA Tuxen 1937
  Molinietalia caeruleae Koch 1926
    Deschampsio refractae-Molinietum caeruleae (Rivas Goday & Borja
    1961) Rivas-Martinez 2002
    Cynosurion cristati Tuxen 1947
Phleo nodosi-Cynosuretum cristati Rodriguez-Rojo & Fernandez-Gonzalez
in press
  Holoschoenetalia vulgaris Br.-Bl. ex Tchou 1948
    Deschampsion mediae Br.-Bl. in Br.-Bl., Roussine & Negre 1952
      Sanguisorbo lateriflorae-Deschampsietum hispanicae
      Rivas-Martinez & G. Lopez in G. Lopez 1978 corr. Rivas-Martinez,
      Fernandez-Gonzalez, Sanchez-Mata & Pizarro 1990

FESTUCO-BROMETEA Br.-Bl. & Tuxen ex Br.-Bl. 1949
  Brometalia erecti Br.-Bl. 1936
  Teucrio pyrenaici-Bromion erecti Rivas-Martinez, Fernandez-Gonzalez
  & Loidi 1999
    Cirsio microcephalae-Onobrychidetum hispanicae Rivas Goday & Borja
    1961 corr. Rivas-Martinez, Fernandez-Gonzalez & Loidi 1999
Brachypodietalia phoenicoidis Br.-Bl. ex Molinier 1934
  Brachypodionphoenicoidis Br.-Bl. ex Molinier 1934
    Festuco andresmolinae-Brachypodietum phoenicoidis Rivas Goday &
    Borja 1961 corr. Rivas-Martinez, T.E. Diaz, Fernandez-Gonzalez,
    Izco, Loidi, Lousa & Penas 2002
FESTUCO HYSTRICIS-ONONIDETEA STRIATAE Rivas-Martinez 2002
  Festuco hystricis-Poetalia ligulatae Rivas Goday & Rivas-Martinez
  1963
    Sideritido fontquerianae-Arenarion aggregatae Rivas Goday & Borja
    1961 corr. Rivas-Martinez 2002
      Festucetum hystricis Font Quer 1954
        typicum
        variant semi-dry with Plantago maritima subsp. serpentina
      Paronychio capitatae-Artemisietum lanatae Rivas Goday & Borja
      1961
FESTUCETEA INDIGESTAE Rivas Goday & Rivas-Martinez 1971
  Jasiono sessiliflorae-Koelerietalia crassipedis Rivas-Martinez &
  Canto 1987
    Hieracio castellani-Plantaginion radicatae Rivas-Martinez & Canto
    1987
      Fumano procumbentis-Thymetum izcoi G. Navarro ex Rodriguez-Rojo
      & al. 2012


doi: 10.5209/rev_LAZA.2013.v34.n1.42664

APPENDIX 1

Iberian endemics present in the ten vegetation clusters (numbers
correspond to the synoptic table).

Cluster number                             1   2   3   4   5

Arenaria erinacea
Arenaria obtusiflora subsp. ciliaris       *           *   *
Armeria alliacea subsp. matritensis                *   *   *
Biscutella atropurpurea
Brassica repanda subsp. blancoana                          *
Carduus assoi                                              *
Centaurea cavanillesiana
Centaurea paniculata subsp. castellana                     *
Centaurea pinae
Dianthus legionensis                           *
Euphorbia minuta                                       *
Festuca marginata subsp. andres-molinae
Festuca paniculata subsp. paui
Festuca rivas-martinezii                           *
Galium lucidum subsp. fruticescens
Jasione crispa subsp. sessiliflora             *
Knautia subscaposa                             *           *
Leucanthemopsis pallida var. virescens
Linum salsoloides                                          *
Minuartia campestris
Plantago monosperma subsp. discolor                        *
Sanguisorba lateriflora                    *   *       *   *
Satureja intricata subsp. gracilis
Scorzoneroides carpetana                   *       *   *   *
Sedum nevadense                            *
Senecio minutus
Seseli cantabricum                                 *
Silene legionensis
Silene mellifera
Tanacetum vahlii
Teucrium expassum                                          *
Thymelaea pubescens
Thymus bracteatus                                          *
Thymus leptophyllus subsp. izcoi

Cluster number                             6   7   8   9   10

Arenaria erinacea                                  *   *
Arenaria obtusiflora subsp. ciliaris       *       *       *
Armeria alliacea subsp. matritensis        *       *   *   *
Biscutella atropurpurea                                    *
Brassica repanda subsp. blancoana
Carduus assoi                                      *   *
Centaurea cavanillesiana                           *
Centaurea paniculata subsp. castellana                 *
Centaurea pinae                                    *       *
Dianthus legionensis                                   *
Euphorbia minuta                               *   *
Festuca marginata subsp. andres-molinae    *
Festuca paniculata subsp. paui                     *
Festuca rivas-martinezii                                   *
Galium lucidum subsp. fruticescens             *   *
Jasione crispa subsp. sessiliflora                         *
Knautia subscaposa                         *   *   *
Leucanthemopsis pallida var. virescens             *       *
Linum salsoloides                          *       *       *
Minuartia campestris                               *
Plantago monosperma subsp. discolor                *       *
Sanguisorba lateriflora                        *
Satureja intricata subsp. gracilis                     *
Scorzoneroides carpetana                   *
Sedum nevadense
Senecio minutus                                    *
Seseli cantabricum
Silene legionensis                         *       *   *   *
Silene mellifera                           *
Tanacetum vahlii                                           *
Teucrium expassum                          *       *   *   *
Thymelaea pubescens                                *
Thymus bracteatus                          *       *       *
Thymus leptophyllus subsp. izcoi                           *


ACKNOWLEDGEMENTS

This research was carried out with the financial aid of the University of Castilla-La Mancha Research Program during 2009-2010 (TC20091114), and the Global Change, Earth Sciences and Biodiversity Program of the Spanish Ministry of Education and Science (CGL2009- 13317-C03-03, project SIVIM).

Received: 1 July 2013

Accepted: 3 December 2013

REFERENCES

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Partel, M.--2002--Local plant diversity patterns and evolutionary history at the regional scale--Ecology 83(9): 2361-2366.

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Rodriguez-Rojo, M.P. & Fernandez-Gonzalez, F.--2014--Diversity patterns and typology of Cynosurus cristatus grasslands (Cynosurion cristati Tuxen 1947) in the Iberian Peninsula. Phytocoenologia--Folia Geobot. (in press).

Rodriguez-Rojo, M.P., Madrigal, J., Crespo, G. & Fernandez-Gonzalez, F.--2010--Tipologia de los prados y pastizales calcicolas de la Serrania de Cuenca (Sistema Iberico)--In: Calleja, A., Garcia Navarro, R., Ruiz Mantecon, A. & Pelaez Suarez, R. (Eds.). Pastos: Fuente Natural de Energia. Pp. 15-21. Ed. Univ. Leon, Leon..

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M. Pilar Rodriguez-Rojo, Santiago Sardinero & Federico Fernandez-Gonzalez (*)

(*) Departamento de Ciencias Ambientales. Instituto de Ciencias Ambientales. Universidad de Castilla-La Mancha. Avda. Carlos III s/n. E-45071 Toledo, Spain. E-mail: MPilar.Rodriguez@uclm.es.

Table 1
Synoptic table of the grasslands studied in the southwestern Iberian
System. Columns correspond to the ten clusters obtained in the beta-
flexible classification, ordered according to the four main groups.
Frequency values of species in every group are shown. Dark grey shaded
values indicate [PHI] values [greater than or equal to] 50% and light
grey shading values 50% > [PHI] [greater than or equal to] 25%.

Higher level group                     A      A      A      A      B
Cluster No.                            1      2      3      4      5
N. of releves                          19     6      16     25     26

Lolium perenne                         63     .      .      16     4
Trifolium dubium                       58     .      31     12     .
Bromus hordeaceus                      74     33     6      4      4
Carex muricata subsp. lamprocarpa      37     17     .      4      .
Phleum nodosum                         95     33     56     68     38
Convolvulus arvensis                   47     17     .      4      12
Trifolium striatum                     63     67     .      8      8
Poa angustifolia                       53     .      25     20     8
Trisetum flavescens                    37     .      19     4      .
Trifolium pratense                     63     17     62     52     8
Festuca trichophylla                   89     83     75     84     50
Trifolium campestre                    84    100     19     52     46
Jasione montana                        .      67     .      .      .
Agrostis castellana                    32    100     38     12     .
Jasione crispa subsp. sessiliflora     .      50     .      .      .
Conopodium pyrenaeum                   .      50     12     4      .
Nardus stricta                         11     .      94     .      .
Genista anglica                        5      17     75     .      .
Anthoxanthum odoratum                  .      .      62     12     .
Juncus squarrosus                      .      .      44     .      .
Carex leporina                         5      .      38     4      .
Carex panicea                          5      .      38     .      .
Cerastium fontanum subsp. vulgare      .      .      38     8      .
Potentilla erecta                      .      .      31     .      .
Stachys officinalis                    .      .      44     4      .
Geum hispidum                          11     .      38     .      .
Luzula campestris                      16     17     56     4      4
Euphrasia hirtella                     11     17     56     28     .
Danthonia decumbens                    5      33     62     40     .
Helictochloa marginata                 .      .      38     .      4
Orchis coriophora                      5      17     44     4      .
Rhinanthus minor                       .      .      31     12     4
Saxifraga granulata                    5      .      38     4      .
Holcus lanatus                         32     17     56     16     .
Scorzoneroides carpetana               11     .      25     52     31
Centaurea jacea subsp. angustifolia    16     .      44     56     4
Carex caryophyllea                     .      33     25     60     4
Leontodon hispidus                     11     33     19     44     .
Lotus corniculatus                     74     33     75    100     73
Prunella hyssopifolia                  5      17     12     44     15
Coronilla minima                       .      .      .      .      81
Jasonia tuberosa                       .      .      .      .      35
Polygala monspeliaca                   5      .      .      .      38
Carthamus carduncellus                 .      17     6      12     77
Hippocrepis comosa                     .      .      .      .      31
Onobrychis argentea subsp.             11     .      .      .      50
  hispanica
Helianthemum oelandicum subsp.         .      .      .      .      58
  incanum
Ononis cristata                        5      .      .      4      46
Leucanthemum pallens                   5      .      .      4      31
Potentilla neumanniana                 .      .      .      .      50
Medicago sativa                        21     17     .      .      38
Bromopsis erecta                       5      .      .      .      42
Thymus pulegioides                     16     17     25     20     4
Trifolium ochroleucon                  11     33     19     12     12
Salvia pratensis                       .      .      .      .      4
Geum sylvaticum                        .      .      6      .      23
Filipendula vulgaris                   5      50     56     56     23
Brachypodium phoenicoides              5      .      .      8      4
Polygala calcarea                      .      .      .      8      .
Thymus bracteatus                      .      .      .      .      50
Anthyllis vulneraria subsp.           es.     .      .      .      .
  vulnerarioides
Xeranthemum inapertum                  5      .      .      .      8
Androsace maxima                       .      .      .      .      .
Poa ligulata                           .      .      .      .      .
Arenaria obtusiflora subsp.            11     .      .      .      4
  ciliaris
Bromus squarrosus                      11     .      .      12     4
Alyssum simplex                        .      .      .      .      .
Carduus assoi                          .      .      .      .      4
Teucrium chamaedrys                    .      .      .      .      .
Marrubium supinum                      .      .      .      .      4
Dianthus pungens subsp.                .      .      .      .      4
  brachyanthus
Bombycilaena erecta                    11     .      .      .      12
Helianthemum apenninum                 .      .      .      .      27
Armeria alliacea subsp. matritensis    .      .      6      16     4
Arenaria erinacea                      .      .      .      .      .
Crupina vulgaris                       .      .      .      .      4
Convolvulus lineatus                   .      .      .      .      12
Asphodelus cerasiferus                 .      .      .      .      4
Artemisia pedemontana                  .      .      .      .      .
Sideritis hirsuta                      .      .      .      .      8
Phlomis lychnitis                      .      .      .      .      4
Taeniatherum caput-medusae             .      .      .      .      .
Thymus vulgaris                        .      .      .      .      35
Thymus zygis subsp. sylvestris         .      .      .      .      .
Trigonella gladiata                    .      .      .      .      .
Centaurea paniculata subsp.            .      .      .      .      .
  castellana
Thymus leptophyllus subsp. izcoi       .      .      .      .      .
Plantago holosteum                     5      67     6      .      4
Scleranthus polycarpos                 5      33     .      .      .
Sedum amplexicaule                     16     67     .      .      .
Potentilla cinerea                     5      17     .      .      .
Evax carpetana                         .      50     .      .      .
Festuca rivas-martinezii               .      .      6      .      .
Cerastium pumilum                      21     .      6      4      8
Festuca glacilior                      .      .      .      .      12
Petrorhagia prolifera                  5      50     .      .      4
Koeleria crassipes                     .      17     .      .      .
Herniaria cinerea                      .      17     .      .      .
Crucianella angustifolia               .      17     .      .      4
Trifolium strictum                     16     17     6      .      .
Poa bulbosa                            47     17     6      12     12
Pilosella castellana                   .      50     .      .      .
Carduus carpetanus                     11     17     .      .      .
Achillea odorata                       21     33     .      16     62
Anthyllis vulneraria subsp.            .      .      .      .      8
  gandogeri
Trifolium arvense                      5      33     .      .      .
Aira caryophyllea                      21     67     38     4      .
Ranunculus paludosus                   16     33     12     4      4
Anthemis arvensis                      26     33     6      4      .
Trifolium repens                       95     33     12     72     12
Cynosurus cristatus                    68     .      56     28     .
Bellis perennis                        79     .      44     68     23
Gaudinia fragilis                      42     .      6      36     .
Hypochaeris radicata                   74    100     50     44     12
Achillea tomentosa                     .      83     .      .      .
Rumex acetosella subsp. angiocarpus    11     83     6      .      .
Logfia minima                          5      67     .      .      .
Carum verticillatum                    32     .      94     60     .
Ranunculus bulbosus subsp. aleae       42     .      88     92     27
Deschampsia caespitosa subsp.          32     .      75    100     15
  subtriflora
Plantago media                         37     .      69     96     23
Plantago maritima subsp. serpentina    5      .      .      60     .
Briza media                            5      .      56     72     4
Festuca hystrix                        .      .      .      .      81
Koeleria vallesiana                    11     .      .      4     100
Galium estebanii var. leioclados       16     .      50     20     77
Helictochloa pratensis subsp.          .      .      .      8      58
  iberica
Cirsium acaule                         16     .      25     36     62

Higher level group                     B      B      C      C      D
Cluster No.                            6      7      8      9      10
N. of releves                          11     6      26     5      20

Lolium perenne                         .      .      .      .      .
Trifolium dubium                       9      .      .      .      .
Bromus hordeaceus                      27     .      8      .      40
Carex muricata subsp. lamprocarpa      .      .      .      .      .
Phleum nodosum                         27     83     .      .      10
Convolvulus arvensis                   .      33     4      .      .
Trifolium striatum                     .      17     4      .      45
Poa angustifolia                       27     50     .      .      .
Trisetum flavescens                    9      33     .      .      .
Trifolium pratense                     55     33     .      .      .
Festuca trichophylla                   45     83     .      .      20
Trifolium campestre                    55     83     4      .      80
Jasione montana                        .      .      .      .      10
Agrostis castellana                    45     .      4      .      .
Jasione crispa subsp. sessiliflora     .      .      .      .      10
Conopodium pyrenaeum                   .      .      .      .      .
Nardus stricta                         9      .      .      .      .
Genista anglica                        .      .      .      .      5
Anthoxanthum odoratum                  .      .      .      .      .
Juncus squarrosus                      .      .      .      .      .
Carex leporina                         .      .      .      .      .
Carex panicea                          .      .      .      .      .
Cerastium fontanum subsp. vulgare      .      .      .      .      .
Potentilla erecta                      .      .      .      .      .
Stachys officinalis                    18     .      .      .      .
Geum hispidum                          .      .      .      .      .
Luzula campestris                      9      .      .      .      10
Euphrasia hirtella                     18     .      .      .      .
Danthonia decumbens                    27     .      .      .      .
Helictochloa marginata                 .      .      .      .      25
Orchis coriophora                      .      17     .      .      5
Rhinanthus minor                       .      .      .      .      .
Saxifraga granulata                    9      .      .      .      20
Holcus lanatus                         9      33     .      .      .
Scorzoneroides carpetana               9      .      .      .      .
Centaurea jacea subsp. angustifolia    27     .      .      .      .
Carex caryophyllea                     9      33     .      .      10
Leontodon hispidus                     .      .      .      .      .
Lotus corniculatus                    100    100     4      .      20
Prunella hyssopifolia                  27     33     .      .      .
Coronilla minima                       27     17     27     20     10
Jasonia tuberosa                       .      .      .      .      .
Polygala monspeliaca                   .      17     .      .      .
Carthamus carduncellus                 73     .      42     .      25
Hippocrepis comosa                     9      .      4      .      .
Onobrychis argentea subsp.             36     17     12     .      5
  hispanica
Helianthemum oelandicum subsp.         27     17     38     20     15
  incanum
Ononis cristata                        36     .      12     20     .
Leucanthemum pallens                   27     .      .      .      .
Potentilla neumanniana                 55     .      35     20     30
Medicago sativa                        27     17     .      20     .
Bromopsis erecta                       91     17     15     20     5
Thymus pulegioides                     73     17     .      .      15
Trifolium ochroleucon                  64     .      .      .      5
Salvia pratensis                       36     .      12     .      .
Geum sylvaticum                        55     17     8      .      25
Filipendula vulgaris                   82     33     4      .      20
Brachypodium phoenicoides              45    100     4      .      .
Polygala calcarea                      18     50     .      .      .
Thymus bracteatus                      18     .      96     .      15
Anthyllis vulneraria subsp.            .      .      54     .      5
  vulnerarioides
Xeranthemum inapertum                  .      .      81     60     20
Androsace maxima                       .      .      62     40     .
Poa ligulata                           .      .      58     20     15
Arenaria obtusiflora subsp.            9      .      46     .      10
  ciliaris
Bromus squarrosus                      .      .      58     40     5
Alyssum simplex                        9      .      46     20     10
Carduus assoi                          .      .      46     40     .
Teucrium chamaedrys                    .      .      31     20     .
Marrubium supinum                      .      .      35     20     10
Dianthus pungens subsp.                9      .      46     40     20
  brachyanthus
Bombycilaena erecta                    9      .      54     80     5
Helianthemum apenninum                 18     17     54     40     15
Armeria alliacea subsp. matritensis    27     .      42     20     5
Arenaria erinacea                      .      .      31     40     .
Crupina vulgaris                       .      .      31     40     .
Convolvulus lineatus                   .      17     35     20     15
Asphodelus cerasiferus                 .      .      31     60     5
Artemisia pedemontana                  .      .      19    100     .
Sideritis hirsuta                      .      .      15    100     .
Phlomis lychnitis                      .      .      4      80     5
Taeniatherum caput-medusae             .      .      4      60     .
Thymus vulgaris                        .      17     8      80     5
Thymus zygis subsp. sylvestris         .      .      .      40     .
Trigonella gladiata                    .      .      .      40     .
Centaurea paniculata subsp.            .      .      .      40     .
  castellana
Thymus leptophyllus subsp. izcoi       .      .      .      .      85
Plantago holosteum                     .      .      4      .     100
Scleranthus polycarpos                 9      .      4      .      80
Sedum amplexicaule                     9      .      12     .      95
Potentilla cinerea                     .      .      .      .      65
Evax carpetana                         .      .      .      .      65
Festuca rivas-martinezii               .      .      .      .      40
Cerastium pumilum                      9      .      31     20     70
Festuca glacilior                      .      .      12     40     55
Petrorhagia prolifera                  9      .      38     20     70
Koeleria crassipes                     .      .      .      .      35
Herniaria cinerea                      .      .      .      .      35
Crucianella angustifolia               .      .      23     20     50
Trifolium strictum                     .      .      .      .      40
Poa bulbosa                            9      17     38     .      70
Pilosella castellana                   .      .      .      .      40
Carduus carpetanus                     .      .      4      .      30
Achillea odorata                       82     33     50     20     85
Anthyllis vulneraria subsp.            .      .      8      40     35
  gandogeri
Trifolium arvense                      .      .      .      .      30
Aira caryophyllea                      18     .      .      .      55
Ranunculus paludosus                   9      .      .      .      40
Anthemis arvensis                      9      .      31     20     50
Trifolium repens                       36     17     .      .      35
Cynosurus cristatus                    9      .      .      .      .
Bellis perennis                        9      33     .      .      5
Gaudinia fragilis                      .      .      .      .      .
Hypochaeris radicata                   18     17     .      .      35
Achillea tomentosa                     .      .      8      .      45
Rumex acetosella subsp. angiocarpus    .      .      .      .      75
Logfia minima                          .      .      .      .      40
Carum verticillatum                    .      17     .      .      .
Ranunculus bulbosus subsp. aleae       18     67     .      .      .
Deschampsia caespitosa subsp.          27     33     .      .      .
  subtriflora
Plantago media                         91     67     .      .      .
Plantago maritima subsp. serpentina    18     .      .      .      .
Briza media                            73     17     .      .      .
Festuca hystrix                        27     .      96     60     5
Koeleria vallesiana                    64     50    100    100     45
Galium estebanii var. leioclados       82     83     4      .      10
Helictochloa pratensis subsp.          73     50     15     40     15
  iberica
Cirsium acaule                        100     50     8      .      5

Table 2
Festucetum hystricis
semi-dry var. with Plantago maritima subsp. serpentine
(Sideritido fontquerianae-Arenarion aggregatae, Festuco hystricis-
Poetalia ligulatae, Festuco hystricis-Ononidetea striatae)

Altitude (m asl)                 1300    1288    1272    1360    1610
Exposure                           W       W       N       W       .
Slope ([degrees])                  5      20      10      15       .
Cover (%)                         100     80      100     90      95
Releve N.                          1       2       3       4       5

Characteristics:
Koeleria vallesiana                5      10      20      10       5
Festuca hystrix                    .       1       .       .      30
Coronilla minima                   1      10      10      20       5
Carthamus carduncellus             1       .       5       1      10
Helianthemum oelandicum            1       .       .       1       5
  subsp. incanum
Achillea odorata                   5       1      0.1      1       .
Thymus bracteatus                  .       .       .      20      10
Linum salsoloides                  .      0.1      .       1       .
Plantago monosperma subsp.         .       .       .       .       5
  discolor
Carex humilis                      .       .       .       .       1
Differentials of the variant:      .       .       .       .       .
Plantago maritima subsp.          20       5       1      10       1
  serpentina
Jasonia tuberosa                  10       5       .       5       5
Cirsium acaule                     .       5       .       1       5
Galium estebanii var.              5       5       5       1       1
  leioclados
Helictochloa pratensis subsp.      .       .       1       1      0.1
  iberica
Deschampsia caespitosa subsp.      .       .       .       1       1
  subtriflora
Companions: Eryngium campestre     1       1       5      0.1      .
Pilosella officinarum             10       5       .       5       .
Lotus corniculatus                 1       1       1       .       .
Plantago lanceolata                5       1       1       .       .
Sanguisorba minor                  .       .       .       .       .
Onobrychis argentea subsp.         .       5       5       1       .
  hispanica
Potentilla neumanniana             1       .       .       .       .
Teucrium expassum                  .       1       .       1       .
Festuca trichophylla              10       5       5       5       .
Ononis cristata                    .       1       .       1       5
Ononis spinosa                     .       1       .       .       .
Trifolium campestre                1       .      0.1      .       .
Bromopsis erecta                  10       5      20       5       .
Medicago sativa                    .       .      10       .       .
Polygala monspeliaca               .      0.1      1      0.1      .
Phleum nodosum                     5       .       1       .       .
Prunella laciniata                10       5       .       .       .
Galium verum                       5       1       .       1       .
Thymus vulgaris                    .       1      0.1      .       .
Taraxacum sect. Ruderalia          .       .       .       .       .
Hippocrepis comosa                 .       .       .       .       .
Scorzoneroides carpetana           1       .       .       1       .
Leucanthemum pallens               .      0.1      .       .       .
Aegilops geniculata                1       .       1       1       .
Brachypodium distachyon            .       .       1               .
Ranunculus bulbosus subsp.         1       .       .      0.1      .
  aleae
Helianthemum apenninum             .       .       1       .       .
Cirsium echinatum                  .       .       .       .       .
Plantago media                     .       1       .       5       .
Bellis perennis                    5       1       .       1       .
Geum sylvaticum                    .       .       .       .       .
Filipendula vulgaris               .       .       .       .       .
Crepis vesicaria subsp.           0.1      .       .      0.1      .
  haenseleri
Astragalus incanus                 .       .       .       .       .
Prunella hyssopifolia              .       1       .       .       .
Arabis hirsuta                     .       .       .       .       .
Euphorbia nicaeensis               .       .       .       .       .
Klasea nudicaulis                  .       .       .       .       .
Carex flacca                       .       .       .       .       .
Bombycilaena erecta                .       .      0.1      .       .
Convolvulus arvensis               .       .       .       .       .
Helictochloa bromoides             .       5       .       .       .
Carex halleriana                   .       .       .       .       .
Plantago afra                      .       .       .       .       .
Lavandula latifolia                .       .       .       .       .
Taraxacum obovatum                 .       .       1       .       .
Globularia vulgaris                .       .       .       .       .
Cerastium pumilum                  .       .       1       .       .
Herniaria glabra                   .       .       .       .       .
Sideritis hirsuta                  .       .       .       .       .

Altitude (m asl)                 1478    1358    1392    1537    1539
Exposure                           S       N       .       .       .
Slope ([degrees])                 15      15       .       .       .
Cover (%)                         80      90      90      60      95
Releve N.                          6       7       8       9      10

Characteristics:
Koeleria vallesiana                5       1      20      20      20
Festuca hystrix                   30      20      30      20      40
Coronilla minima                   5      10       5       1       .
Carthamus carduncellus            10       5       1       5       5
Helianthemum oelandicum            1       5       5       1       1
  subsp. incanum
Achillea odorata                   .       .       .       .       1
Thymus bracteatus                 10      10      20       .       .
Linum salsoloides                  .      10       1       1       .
Plantago monosperma subsp.         .       .       .      10       .
  discolor
Carex humilis                      1       .       .       .       .
Differentials of the variant:      .       .       .       .       .
Plantago maritima subsp.           1      10       5       1       5
  serpentina
Jasonia tuberosa                   1       .       1       1       .
Cirsium acaule                    10       5       .       .       5
Galium estebanii var.              1       1       .       .       1
  leioclados
Helictochloa pratensis subsp.      1       .       .       .       .
  iberica
Deschampsia caespitosa subsp.      .       .       .       .       1
  subtriflora
Companions: Eryngium campestre    0.1     0.1      1       .      0.1
Pilosella officinarum              1       .       .       .       5
Lotus corniculatus                0.1      .       .      0.1      1
Plantago lanceolata               0.1      .       .       .       .
Sanguisorba minor                  1       .       .      0.1      .
Onobrychis argentea subsp.         .       .       .       .       .
  hispanica
Potentilla neumanniana             1       1       .       .       .
Teucrium expassum                  1       .       1       .       .
Festuca trichophylla                      0.1      .       .       .
Ononis cristata                    .       1       1       .       .
Ononis spinosa                     .       .       .       .       .
Trifolium campestre                .       .       .       .       .
Bromopsis erecta                   5       1       .       .       .
Medicago sativa                    .       .       .       .       .
Polygala monspeliaca               .       .       .       .       .
Phleum nodosum                     .       .       .       .       1
Prunella laciniata                 1       .       .       .       .
Galium verum                       1       .       1       .       5
Thymus vulgaris                    .       .       .       .       .
Taraxacum sect. Ruderalia          .       .       .       .       .
Hippocrepis comosa                 .       .       .       .       .
Scorzoneroides carpetana           .       .       .       .      0.1
Leucanthemum pallens               .       .       .       .       .
Aegilops geniculata                .       .       .       .       .
Brachypodium distachyon            .       .       .       .       .
Ranunculus bulbosus subsp.         .      0.1      .       .       .
  aleae
Helianthemum apenninum             .       .       .       .       .
Cirsium echinatum                  .       .       .       .       .
Plantago media                     .       5       .       .       .
Bellis perennis                    .       .       .       .       .
Geum sylvaticum                    .       .       .       .      0.1
Filipendula vulgaris              0.1      5       .       .       1
Crepis vesicaria subsp.            .       .       .       .       .
  haenseleri
Astragalus incanus                 .       .       .       .       .
Prunella hyssopifolia              .       .       .       .       1
Arabis hirsuta                     .       .       .       .       .
Euphorbia nicaeensis              0.1      .       .       .       .
Klasea nudicaulis                  .       .       .      0.1      1
Carex flacca                       .       1       .       .       .
Bombycilaena erecta                .       .       .       .       .
Convolvulus arvensis               .       .       .      0.1      .
Helictochloa bromoides             .       .       .       .       .
Carex halleriana                   .       5       .       .       .
Plantago afra                      .       .       .       .       .
Lavandula latifolia                .       .       .       .       .
Taraxacum obovatum                 .       .       .       .       .
Globularia vulgaris                .       5       1       .       .
Cerastium pumilum                  .       .       .       .       .
Herniaria glabra                   .       .       .      0.1     0.1
Sideritis hirsuta                  .       .       .       .       .

Altitude (m asl)                 1277    1225    1330    1267    1265
Exposure                           W       .       W      SW      NE
Slope ([degrees])                 15       .      20      10      20
Cover (%)                         70      60      80      70      70
Releve N.                         11      12      13      14      15

Characteristics:
Koeleria vallesiana               10      10       5      10      30
Festuca hystrix                   10      10      10       5      10
Coronilla minima                   5       5      10       5       .
Carthamus carduncellus             .      0.1      1       1      0.1
Helianthemum oelandicum            1       .       1       .       .
  subsp. incanum
Achillea odorata                   .       1       1       .       .
Thymus bracteatus                  .       1       1       .       .
Linum salsoloides                  .       .       .       .       .
Plantago monosperma subsp.         .       .       .       .       .
  discolor
Carex humilis                      .       .       .       .       .
Differentials of the variant:      .       .       .       .       .
Plantago maritima subsp.          10      10       5      20      0.1
  serpentina
Jasonia tuberosa                   .       1       .       .       .
Cirsium acaule                     1       .      10       .      10
Galium estebanii var.              5       .       5       5       5
  leioclados
Helictochloa pratensis subsp.     20       5       1       5       .
  iberica
Deschampsia caespitosa subsp.      .       .       .       .       .
  subtriflora
Companions: Eryngium campestre     1       1       1       1       1
Pilosella officinarum              1       5      10       5       5
Lotus corniculatus                 1       .       .       1       1
Plantago lanceolata                1       1       .       .       .
Sanguisorba minor                 0.1      1       .       1      0.1
Onobrychis argentea subsp.         .      10       5       1       .
  hispanica
Potentilla neumanniana            0.1      .       .       .       1
Teucrium expassum                  1       1      0.1      1       1
Festuca trichophylla               .       .       .       1       .
Ononis cristata                    .       .       5       .       .
Ononis spinosa                     5       .       1       1       .
Trifolium campestre               0.1      .       .       .       1
Bromopsis erecta                   .       .      10       .       .
Medicago sativa                    1       1       1       .       1
Polygala monspeliaca              0.1      .       .      0.1      .
Phleum nodosum                     .       .       .       .      0.1
Prunella laciniata                 .       .       .      0.1      1
Galium verum                       .       .       .       .       .
Thymus vulgaris                    1       1       1       1      0.1
Taraxacum sect. Ruderalia         0.1      .       .      0.1     0.1
Hippocrepis comosa                 1       .       .       1       1
Scorzoneroides carpetana           .       .       .       .       1
Leucanthemum pallens               .       1       .       .       1
Aegilops geniculata                .       1       .       .       .
Brachypodium distachyon            1       1       1       1       .
Ranunculus bulbosus subsp.         .       .       .      0.1      .
  aleae
Helianthemum apenninum             .       .       1       .       1
Cirsium echinatum                 0.1      .       .       .       .
Plantago media                     .       .       .       .       .
Bellis perennis                    .       .       .       1       .
Geum sylvaticum                    .       .       .       .       .
Filipendula vulgaris               .       .      0.1      .       .
Crepis vesicaria subsp.            .       .       1       .       .
  haenseleri
Astragalus incanus                 .      0.1      .       .       .
Prunella hyssopifolia              .       .       1       .       .
Arabis hirsuta                     .       .      0.1      .       .
Euphorbia nicaeensis               .       .       .       .       .
Klasea nudicaulis                  .       .       .       .       .
Carex flacca                       .       .       .       .       .
Bombycilaena erecta                .       .       .       .      0.1
Convolvulus arvensis              0.1      .       .       .       .
Helictochloa bromoides             .       .       .       .       .
Carex halleriana                   .       .       .       .       .
Plantago afra                      1      0.1      .       .       .
Lavandula latifolia                1       .       .       .       .
Taraxacum obovatum                 .       .       1       .       .
Globularia vulgaris                .       .       .       .       .
Cerastium pumilum                  .       .       1       .       .
Herniaria glabra                   .       .       .       .       .
Sideritis hirsuta                 0.1     0.1      .       .       .

Altitude (m asl)                 1326    1270    1342
Exposure                           N       E      SW
Slope ([degrees])                 20      15      20
Cover (%)                         70      100     70
Releve N.                         16      17      18

Characteristics:
Koeleria vallesiana               10      20      20
Festuca hystrix                   10       5      20
Coronilla minima                   .      10       5
Carthamus carduncellus            0.1      5       5
Helianthemum oelandicum            .       1       .
  subsp. incanum
Achillea odorata                   .       1       .
Thymus bracteatus                  .       5       5
Linum salsoloides                  .       .       .
Plantago monosperma subsp.         .       .       .
  discolor
Carex humilis                      .       .       .
Differentials of the variant:      .       .       .
Plantago maritima subsp.          10       1       1
  serpentina
Jasonia tuberosa                   .       .       .
Cirsium acaule                     5       5       .
Galium estebanii var.              5       5       .
  leioclados
Helictochloa pratensis subsp.      1       .       .
  iberica
Deschampsia caespitosa subsp.      .       .       .
  subtriflora
Companions: Eryngium campestre     1       1      10
Pilosella officinarum              1       1       5
Lotus corniculatus                0.1      5       1
Plantago lanceolata               0.1     0.1      1
Sanguisorba minor                  1       1       .
Onobrychis argentea subsp.        0.1     10       1
  hispanica
Potentilla neumanniana             1       1       .
Teucrium expassum                  .      0.1      .
Festuca trichophylla               .       .       .
Ononis cristata                    .       5       .
Ononis spinosa                     5       5       5
Trifolium campestre                1       .       1
Bromopsis erecta                   .       .      10
Medicago sativa                    .       .      0.1
Polygala monspeliaca               .       .      0.1
Phleum nodosum                     .       .       .
Prunella laciniata                 5      0.1      .
Galium verum                       .       .       .
Thymus vulgaris                    1       .       .
Taraxacum sect. Ruderalia          .       .       1
Hippocrepis comosa                 5       5       .
Scorzoneroides carpetana           1       .       1
Leucanthemum pallens               1      0.1      .
Aegilops geniculata                .       .       .
Brachypodium distachyon            .       .       .
Ranunculus bulbosus subsp.        0.1      .       .
  aleae
Helianthemum apenninum             .       .      0.1
Cirsium echinatum                  .       .       .
Plantago media                     .       1       .
Bellis perennis                    .       .       .
Geum sylvaticum                    .      0.1      .
Filipendula vulgaris               .       .       .
Crepis vesicaria subsp.            .      0.1      .
  haenseleri
Astragalus incanus                 .       .       1
Prunella hyssopifolia              .       .       .
Arabis hirsuta                     .       .       .
Euphorbia nicaeensis              0.1      .       .
Klasea nudicaulis                  .       .       .
Carex flacca                       .      0.1      .
Bombycilaena erecta                .       .       .
Convolvulus arvensis               .       .       .
Helictochloa bromoides             .       .       .
Carex halleriana                   .       1       .
Plantago afra                      .       .       .
Lavandula latifolia               0.1      .       .
Taraxacum obovatum                 .       .       .
Globularia vulgaris                .       .       .
Cerastium pumilum                  .       .       .
Herniaria glabra                   .       .       .
Sideritis hirsuta                  .       .       .

Other species: Carex caryophyllea 5 in1; Bromus squarrosus, Centaurea
jacea 1 in 1; Scabiosa columbaria subsp. affinis 1 in 2; Festuca
gracilior 10 in 3; Bupleurum baldense, Convolvulus lineatus, Medicago
rigidula, Melilotus sulcatus, Poa bulbosa, Myosotis ramosissima,
Rhinanthus minor 1 in 3; Alyssum alyssoides, Crucianella angustifolia,
Daucus carota, Leontodon longirostris, Veronica arvensis 0.1 in 3;
Fritillaria lusitanica 0.1 in 5; Brachypodium phoenicoides 5 in 6;
Aphyllanthes monspeliensis, Cuscuta epithymum subsp. kotschyi 1;
Knautia subscaposa in 6; Asphodelos cerasiferus, Festuca fenas,
Marrubium supinum, Salvia pratensis, Thesium humifusum 0.1 in 6; Briza
media 0.1 in 7; Arenaria obtusiflora subsp. ciliaris, Brassica repanda
subsp. blancoana 0.1 in 8; Klasea nudicaulis, Ornithogalum narbonense
0.1 in 9; Ranunculus paludosis 1 in 10; Armeria alliacea subsp.
matritensis 0.1 in 10; Poa angustifolia 1 in 11; Arenaria modesta,
Phlomis lychnitis 0.1 in 11; Dorycnium pentaphyllum 0.1 in 12; Logfia
gallica, Xeranthemum inapertum 0.1 in 13; Helictochloa marginata 1 in
15; Hypochaeris radicata 0.1 in 15; Cichorium intybus 0.1 in 15.

Localities: 1, 2: Cuenca, Sierra de zafrilla, Loma de la Herradura,
30TXK0557, 20.06.2010; 3: Guadalajara, Pinilla de Molina, La Hoya del
Perdigon, 30TwL9403, 17.06.2010; 4: Cuenca, Sierra de Valdemeca, Cerro
del Bu, 30TXK0453, 20.06.2010; 5: Guadalajara, Checa, P.N. Alto Tajo,
Los Asperones, 30TXK0275, 2.07.2010; 6: Cuenca, Sierra de zafrilla,
Cerro del zarzal, 30TXK1853, 2.07.2009; 7: Guadalajara, Cueva del
Hierro, 30TwK8093, 9.06.2009;8: Cuenca, Sierra de Valdemeca, El
Bujedal, 30TXK0652, 20.06.2010; 9.10: Cuenca, Serrania de Cuenca,
quinto de los Prados, 30TwK8989, 7.07.2009; 11: Cuenca, Serrania de
Cuenca, Loma del Ocejon, 30TwK9966, 10.06.2009; 12: Cuenca, Serrania
de Cuenca, El Monton de Tierra, 30TXK0161, 11.06.2009; 13: Cuenca,
Serrania de Cuenca, Puntal de la Salceda, 30TXK0851, 20.06.2010; 14:
Cuenca, Serrania de Cuenca, Riscos de la Carrascosa, 30TwK8879,
10.06.2009; 15: Cuenca, Serrania de Cuenca, Monte Sierra de las
Canales, 30TXK0064, 11.06.2009; 16: Cuenca, Serrania de Cuenca, Prados
de la Cierva, 30TwK9843, 11.06.2009; 17: Cuenca, Sierra de Tragacete,
Cerro de San Miguel, 30TXK0064, 11.06.2009; 18: Cuenca, Sierra de
Valdemiguete, Herreria de los Chorros, 30TXK0164, 11.06.2009.
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Title Annotation:articulo en ingles
Author:Rodriguez-Rojo, M. Pilar; Sardinero, Santiago; Fernandez-Gonzalez, Federico
Publication:Lazaroa
Date:Jan 1, 2013
Words:8029
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