Research Paper |
Corresponding author: Idoia Biurrun ( idoia.biurrun@ehu.es ) Academic editor: Arkadiusz Nowak
© 2021 Itziar García-Mijangos, Asun Berastegi, Idoia Biurrun, Iwona Dembicz, Monika Janišová, Anna Kuzemko, Denys Vynokurov, Didem Ambarlı, Javier Etayo, Goffredo Filibeck, Ute Jandt, Rayna Natcheva, Oktay Yildiz, Jürgen Dengler.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
García-Mijangos I, Berastegi A, Biurrun I, Dembicz I, Janišová M, Kuzemko A, Vynokurov D, Ambarlı D, Etayo J, Filibeck G, Jandt U, Natcheva R, Yildiz O, Dengler J (2021) Grasslands of Navarre (Spain), focusing on the Festuco-Brometea: classification, hierarchical expert system and characterisation. Vegetation Classification and Survey 2: 195-231. https://doi.org/10.3897/VCS/2021/69614
|
Aims: To clarify the syntaxonomic position of the grasslands in Navarre, with special focus on the dry grasslands, and to characterise the resulting syntaxonomic units in terms of diagnostic species and ecological conditions. Study area: Navarre (northern Spain). Methods: We sampled 119 plots of 10 m2 following the standardised EDGG methodology and analysed them together with 839 plots of similar size recorded in the 1990. For the classification, we used the modified TWINSPAN algorithm, complemented by the determination of diagnostic species with phi coefficients of association, which led to the creation of an expert system. We conducted these steps in a hierarchical manner for each syntaxonomic rank. We visualised the position of the syntaxa along environmental gradients by means of NMDS. Species richness, and structural and ecological characteristics of the syntaxa were compared by ANOVAs. Results: We could clearly identify five phytosociological classes: Lygeo-Stipetea, Festuco-Brometea, Molinio-Arrhenatheretea, Nardetea strictae, and Elyno-Seslerietea. Within the Festuco-Brometea a xeric and a meso-xeric order could be distinguished, with two alliances each, and eight associations in total: Thymelaeo-Aphyllanthetum, Jurineo-Festucetum, Helianthemo-Koelerietum, Prunello-Plantaginetum, Carduncello-Brachypodietum, Helictotricho-Seslerietum, Calamintho-Seselietum and Carici-Teucrietum. Conclusions: The combination of numerical methods allowed a consistent and more objective classification of grassland types in Navarre than previous approaches. At the association level, we could largely reproduce the units previously described with traditional phytosociological methods. By contrast, at higher syntaxonomic level, our analyses suggest significant modifications. Most importantly, a major part of the units traditionally included in the Festuco-Ononidetea seem to fall within the Festuco-Brometea. We could show that bryophytes and lichens are core elements of these grasslands and particularly the Mediterranean ones of Lygeo-Stipetea, both in terms of biodiversity and of diagnostic species. We conclude that the combination of our different numerical methods is promising for deriving more objective and reproducible delimitations of syntaxa in a hierarchical manner.
Taxonomic references:
Syntaxonomic reference:
Abbreviations: ANOVA = analysis of variance; EDGG = Eurasian Dry Grassland Group; NMDS: non-metric multidimensional scaling; TWINSPAN = Two-Way Indicator Species Analysis.
diagnostic species, electronic expert system, Elyno-Seslerietea, Festuco-Brometea, Festuco-Ononidetea, grassland, Lygeo-Stipetea, modified TWINSPAN, Molinio-Arrhenatheretea, Nardetea strictae, Navarre, vegetation classification
Grasslands represent one of the most extensive and diverse formations of the world, yet undervalued and under-researched. Grasslands are spontaneously occurring herbaceous vegetation types that are mostly dominated by grasses (Poaceae) or other graminoids (Cyperaceae, Juncaceae) and have a relatively high herb-layer cover (usually > 10%), while woody species (dwarf shrubs, shrubs and trees), if present at all, have a significantly lower cover than the herbs (
Since the second half of the 20th century, European grasslands have experienced two extremes of the land-use gradient, and both resulted in the loss of grassland biodiversity (
During the last decades, a great effort on grassland classification has been made, based on large vegetation-plot databases and numerical analysis in several countries or regions across Europe to delimit and define the different syntaxonomic units. Several studies have been developed at a regional, up to continental scale on dry-grasslands (
Phytosociological studies in the Iberian Peninsula have been broadly developed in the last century and were synthesized in the syntaxonomic checklist of Spain (
Navarre region, located in northern Iberian Peninsula, is a bioclimatically diverse region where Alpine, Atlantic and Mediterranean biogeographical areas converge. The long history of grazing and management throughout the area has resulted in the broad spread of grasslands. The region has an important elevational and precipitation gradient that allows the coexistence of dry and mesic grasslands as well as alpine and Mediterranean semi-arid communities (
According to
The high grassland diversity in Navarre reflects the richness of grassland habitats of interest for European Community (
The large amount of data available related to grassland in the region of Navarre and its strategic geographical position where different climatic conditions converge provide a unique opportunity to clarify grassland syntaxonomy, especially those from submediterranean areas. More specifically, we aim to 1) Identify the main grassland types in Navarre using numerical and reproducible methods, 2) Compare our results with existing traditional classifications at the level of alliance or association 3) Define the diagnostic species of syntaxa including bryophytes and lichens. 4) Characterise and differentiate associations with regard to topographic, edaphic and climatic variables.
Navarre is a territory of 10,391 km2 located in the north-central part of the Iberian Peninsula. There is a wide elevational range in the region, from 25 m a.s.l. in Endarlatsa, 15 km from the Cantabrian Sea in the north, to 2,466 m a.s.l. in the Mesa de los Tres Reyes in the western Pyrenees. The bioclimate is temperate in the northern part of the region, and Mediterranean in the south, with large submediterranean areas in the central part (
Several types of deciduous forests prevail in the temperate zone, where secondary grasslands, mainly mesic and meso-xeric grasslands, are an important component of the landscape. Sclerophyllous woodlands dominate in the Mediterranean areas of southern Navarre, with Mediterranean grasslands and garrigues as secondary vegetation. In the Pyrenees, alpine grasslands and scrubs occur above 1,700 m a.s.l., in the subalpine belt mostly as secondary vegetation replacing Pinus uncinata woodlands, and as potential natural vegetation in the alpine belt, above ca. 2,100 m a.s.l. (
Geological diversity also has a great influence on the vegetation. Shales, quartzites or granites from the Palaeozoic are common in the northern area of Navarre, mostly in the Atlantic region. Red sandstones and conglomerates from the Triassic surround these Palaeozoic rocks. Limestones, marls and dolomites from the Jurassic and Cretaceous period, and also limestones, marls, flysch substrates, but also calcarenites from the continental Tertiary are dominant in all of the central area of Navarre. From the continental Tertiary, sandstones, clays, slimes, but also limestones and gypsum are dominant in the south of Navarre, mostly in the Mediterranean region (
We took 119 10-m² plots sampled following the standard EDGG methodology (
Furthermore, we included those 839 vegetation plots from
Although these plots from the additional dataset were evenly distributed across the region and all grassland types, we wish to highlight that four of the 11 classes represented in
The combination of both datasets resulted in a total of 958 vegetation plots. The data from EDGG expedition are stored in and available from the GrassPlot database (
Soil samples were collected in each EDGG plot. Samples were taken with a hand shovel from the uppermost 5–10 cm at five random points within the plot, merged in a mixed sample and air-dried. The coarse fragment of the samples was determined by dry screening (Ø > 2 mm) and soil texture was determined by the Bouyoucos hydrometer method (
We retrieved climatic data from CHELSA dataset version 1.2 (
Before numerical analysis, we unified species taxonomy and nomenclature. Vascular plants were named according to Euro+Med PlantBase (
For the initial unsupervised classifications, we used the modified version of TWINSPAN (
In the case of very large datasets, classification is highly dependent on the selection of attributes (species) used. The more attributes used, the data become more scattered (
We created the confusion matrix comparing the original (expert-based) and new numerical (unsupervised) classifications (see Suppl. material
In the case of the classes, we found that three of the traditional classes shared a significant number of frequent species and therefore, we decided to merge them and re-run the previous steps to achieve the final expert system and the final set of diagnostic species of classes. We continued then, with the same approach, with our main target class Festuco-Brometea to search for the most plausible division into orders. Criteria were based on how well the resulting units were floristically and ecologically characterised and how closely they matched the general syntaxonomic system of Europe. Next, we continued in each of the resulting orders to find an appropriate division into alliances and finally for each of the alliances we analysed the appropriate subdivision into associations separately. For each syntaxonomic level and cluster we therefore followed the procedure of: (1) running modified TWINSPAN, (2) identifying a reasonable number of syntaxa of the next lower level and (3) determining their diagnostic species. In the case of order and alliance we selected the relevés that matched both the expert and TWINSPAN classification, but for associations we used only the TWINSPAN results, (4) translating these into an expert system, (5) appling this expert system to the data including the type relevés of all associations included in Festuco-Brometea (details provided in Suppl. material
We followed the fourth edition of the International Code of Phytosociological Nomenclature (ICPN;
We determined diagnostic species using the phi coefficient of association (
To visualize the gradient of vascular plant species composition across the vegetation types, we used non-metric multidimensional scaling (NMDS;
Differences among classes regarding structural, topographic, bioclimatic and soil variables, as well as regarding richness values, were analysed by means of analyses of variance (ANOVAs) in the R programming language (
At the level of ten groups, the TWINSPAN analysis resulted in a division of the data where the classification into seven classes proposed by
The synoptic table with the diagnostic species for each cluster of the modified TWINSPAN analysis is presented in Suppl. material
The relationship between the previous expert-based classification (
Relationship between the original classification and the expert system classification. In each column the number of relevés and the proportion related to the total of relevés belonging to the original classification (in brackets) that match the expert system are shown.
Syntaxonomic classes (original classification) | Expert System classification | Nº rel. per class | |||||
LYG (%) | FES (%) | MOL (%) | NAR (%) | SES (%) | Non-classified | ||
Lygeo–Stipetea | 25 (96) | 1 (4) | 26 | ||||
Stipo–Trachynietea | 10 (100) | 10 | |||||
Ononido–Rosmarinetea | 5 (13) | 33 (87) | 38 | ||||
Festuco–Brometea | 8 (4) | 131 (61) | 34 (16) | 40 (19) | 2 | 215 | |
Festuco–Ononidetea | 158 (75) | 1 (< 1) | 2 (1) | 48 (23) | 1 | 210 | |
Molinio–Arrhenatheretea | 13 (6) | 185 (86) | 17 (8) | 1 | 216 | ||
Nardetea strictae | 149 (96) | 5 (3) | 1 | 155 | |||
Sedo–Scleranthetea | 11 (85) | 1 (8) | 1 | 13 | |||
Elyno–Seslerietea | 1 (2) | 3 (5) | 58 (93) | 2 | 64 | ||
Carici–Kobresietea | 2 | 2 | |||||
Caricetea curvulae | 1 | 1 | |||||
Poetea bulbosae | 6 | 2 | 8 | ||||
Nº relevés per group | 54 | 339 | 220 | 223 | 114 | 8 | 958 |
The expert system analysis included in this group LYG most relevés that were originally classified in the class Lygeo-Stipetea. Communities dominated by therophytes of Stipo-Trachynietea and those from Poetea bulbosae were also classified in this group, as they shared many annual species: Bombycilaena erecta, Catapodium rigidum, Linum strictum, Trachynia distachya, etc. LYG also includes some relevés from the subassociation Thymelaeo-Aphyllanthetum brachypodietosum retusi of the class Ononido-Rosmarinetea and from the association Elytrigio campestris-Brachypodietum phoenicoidis of Festuco-Brometea.
Photo plate showing typical stands of four of the five distinguished vegetation classes (for Festuco-Brometea, see Figures
These communities are characterised by the presence of hard-leaved grasses such as Brachypodium retusum, Helictochloa bromoides, Lygeum spartum and Stipa parviflora and dwarf chamaephytes as well as many therophytes (Table
Abridged constancy table of the five grassland classes considered in this study. Values are percentage constancies, and species are ordered by decreasing phi-values in the respective syntaxon, respectively by decreasing overall constancy for non-diagnostic species. In the upper part vascular plants are given, in the lower part bryophytes and lichens, whose constancies and fidelities have been calculated based only on the plots of the EDGG Field Workshop. In the table, the 15 vascular plant taxa and the eight non-vascular plant taxa with the highest fidelity in a class are shown, plus all taxa that are diagnostic for multiple classes and all taxa with at least 10% overall constancy. Diagnostic species (phi ≥ 0.25) are highlighted in grey, highly diagnostic species (phi ≥ 0.5) in dark grey. The complete constancy table combined with the table of the underlying 958 vegetation plots is given in Suppl. material
Class | All | LYG | FES | MOL | NAR | SES |
---|---|---|---|---|---|---|
# plots | 958 | 54 | 339 | 220 | 223 | 114 |
# plots with bryophyte/lichen treatment | 119 | 19 | 64 | 8 | 11 | 17 |
Class LYG (47 taxa) | ||||||
Linum strictum | 3.9 | 52 | 2 | 1 | . | . |
Brachypodium retusum | 8.4 | 52 | 14 | 1 | . | . |
Catapodium rigidum | 4.3 | 43 | 5 | 1 | . | . |
Lygeum spartum | 2.0 | 33 | <1 | . | . | . |
Asterolinon linum-stellatum | 2.2 | 33 | 1 | <1 | . | . |
Artemisia herba-alba | 1.8 | 31 | . | . | . | . |
Thymus vulgaris subsp. vulgaris | 9.9 | 50 | 20 | . | . | . |
Polygala monspeliaca | 2.4 | 33 | 1 | . | . | . |
Trachynia distachya | 3.0 | 35 | 2 | <1 | . | 1 |
Teucrium capitatum subsp. capitatum | 4.3 | 35 | 6 | . | . | . |
Bombycilaena erecta | 2.8 | 31 | 3 | . | . | . |
Euphorbia exigua | 3.9 | 33 | 5 | <1 | . | . |
Plantago lagopus subsp. lagopus | 1.5 | 26 | . | . | . | . |
Plantago albicans | 1.5 | 26 | . | . | . | . |
Atractylis humilis | 1.8 | 24 | 1 | . | . | . |
[…] | ||||||
Class FES (21 taxa) | ||||||
Bromopsis erecta subsp. erecta | 27.1 | 2 | 65 | 6 | 5 | 11 |
Carthamus mitissimus | 19.4 | 7 | 51 | 1 | 3 | 2 |
Carex humilis | 14.5 | 4 | 40 | . | . | 1 |
Potentilla tabernaemontani | 19.5 | 6 | 48 | 1 | 4 | 10 |
Coronilla minima | 14.2 | 7 | 38 | <1 | . | 1 |
Festuca rectifolia | 17.7 | 4 | 42 | . | 2 | 15 |
Seseli montanum subsp. montanum | 14.1 | 2 | 33 | 1 | 6 | 4 |
Helictochloa pratensis subsp. iberica | 20.9 | 2 | 46 | <1 | 4 | 30 |
Geum sylvaticum | 6.8 | . | 18 | . | 1 | 1 |
Scabiosa columbaria subsp. columbaria | 11.1 | . | 25 | 4 | 3 | 5 |
Medicago lupulina | 20.1 | 6 | 39 | 18 | 3 | 8 |
Onobrychis conferta subsp. hispanica | 6.3 | 2 | 17 | 1 | . | . |
Sanguisorba minor aggr. | 16.0 | 17 | 34 | 6 | 6 | 1 |
Teucrium chamaedrys | 6.8 | 4 | 18 | . | . | 2 |
Trifolium montanum subsp. montanum | 6.2 | . | 15 | 2 | 1 | . |
[…] | ||||||
Class MOL (33 taxa) | ||||||
Holcus lanatus | 11.6 | . | 3 | 44 | 1 | . |
Ranunculus acris subsp. friesianus | 7.8 | . | 1 | 32 | <1 | . |
Agrostis stolonifera subsp. stolonifera | 9.1 | 2 | 3 | 34 | <1 | . |
Trifolium fragiferum | 6.6 | . | . | 28 | 1 | . |
Ranunculus repens | 6.5 | . | . | 26 | 2 | . |
Poa trivialis subsp. trivialis | 8.9 | . | 6 | 30 | . | . |
Lolium perenne | 11.1 | 2 | 4 | 35 | 7 | . |
Schedonorus arundinaceus subsp. arundinaceus | 6.4 | . | 2 | 25 | . | . |
Juncus articulatus | 4.9 | . | . | 21 | <1 | . |
Juncus inflexus | 4.3 | . | . | 19 | . | . |
Centaurea debeauxii | 6.8 | . | 4 | 23 | 1 | . |
Anthoxanthum odoratum | 10.5 | . | 4 | 30 | 9 | 2 |
Rumex acetosa subsp. acetosa | 4.8 | . | <1 | 19 | . | 3 |
Potentilla reptans | 6.7 | 6 | 2 | 24 | 1 | . |
Veronica chamaedrys subsp. chamaedrys | 5.0 | . | <1 | 18 | 3 | . |
[…] | ||||||
Class NAR (17 taxa) | ||||||
Potentilla erecta | 16.7 | . | 1 | 5 | 63 | 4 |
Galium saxatile | 11.5 | . | <1 | <1 | 48 | 1 |
Agrostis capillaris | 35.4 | . | 18 | 27 | 86 | 23 |
Festuca microphylla | 40.2 | . | 22 | 19 | 94 | 48 |
Polygala serpyllifolia | 8.6 | . | 2 | <1 | 33 | . |
Nardus stricta | 8.5 | . | . | . | 34 | 5 |
Danthonia decumbens | 17.1 | . | 10 | 9 | 47 | 4 |
Agrostis curtisii | 5.7 | . | . | . | 25 | . |
Jasione laevis subsp. laevis | 5.8 | . | . | . | 25 | 1 |
Carex pilulifera subsp. pilulifera | 5.5 | . | . | <1 | 23 | . |
Calluna vulgaris | 7.4 | . | . | . | 27 | 8 |
Veronica officinalis | 5.0 | . | . | <1 | 21 | 1 |
Helictochloa marginata subsp. marginata | 6.7 | . | 3 | . | 24 | 2 |
Trifolium alpinum | 3.7 | . | . | . | 16 | . |
Vaccinium myrtillus | 2.7 | . | . | . | 11 | 1 |
[…] | ||||||
Class SES (46 taxa) | ||||||
Helictotrichon sedenense subsp. sedenense | 5.6 | . | . | . | <1 | 46 |
Carex sempervirens subsp. sempervirens | 5.4 | . | 1 | . | <1 | 43 |
Alchemilla plicatula aggr. | 13.5 | . | 4 | . | 19 | 61 |
Festuca gautieri subsp. scoparia | 4.7 | . | <1 | . | <1 | 38 |
Poa alpina | 10.4 | . | 5 | . | 12 | 50 |
Androsace villosa subsp. villosa | 4.4 | . | 1 | . | . | 32 |
Paronychia kapela subsp. serpyllifolia | 4.6 | . | 1 | . | 2 | 32 |
Agrostis schleicheri | 3.9 | . | 1 | . | . | 29 |
Carex ornithopoda | 4.8 | . | 2 | . | 2 | 32 |
Ranunculus carinthiacus | 4.1 | . | 1 | . | 2 | 29 |
Sesleria caerulea subsp. caerulea | 4.3 | . | 2 | . | <1 | 29 |
Trifolium thalii | 7.9 | . | <1 | . | 13 | 38 |
Silene acaulis | 3.4 | . | . | . | 1 | 26 |
Aster alpinus | 3.5 | . | 1 | . | <1 | 25 |
Saxifraga paniculata | 2.9 | . | <1 | . | . | 24 |
[…] | ||||||
Anthyllis vulneraria | 13.6 | 4 | 22 | 1 | 1 | 43 |
Diagnostic for multiple classes (13 taxa) | ||||||
Eryngium campestre | 16.1 | 37 | 32 | 9 | 1 | . |
Genista scorpius | 7.0 | 30 | 15 | . | . | . |
Koeleria vallesiana | 25.1 | 37 | 53 | . | . | 34 |
Dactylis glomerata | 21.0 | 52 | 20 | 43 | 3 | 2 |
Carex flacca subsp. flacca | 25.3 | . | 45 | 35 | 4 | 3 |
Pilosella officinarum | 29.6 | 13 | 41 | 5 | 48 | 13 |
Thymus praecox | 42.5 | . | 63 | 1 | 43 | 82 |
Carex caryophyllea | 27.9 | 4 | 31 | 4 | 51 | 30 |
Helianthemum canum subsp. canum | 14.6 | 2 | 27 | . | 1 | 39 |
Teucrium pyrenaicum | 11.0 | . | 24 | <1 | <1 | 18 |
Trifolium repens | 30.0 | . | 17 | 47 | 52 | 8 |
Campanula scheuchzeri | 7.0 | . | 1 | <1 | 15 | 25 |
Plantago alpina | 6.5 | . | 1 | . | 20 | 13 |
Companion species | ||||||
Lotus corniculatus | 44.3 | 4 | 53 | 32 | 54 | 43 |
Plantago lanceolata | 37.5 | 7 | 50 | 47 | 29 | 11 |
Trifolium pratense | 32.0 | 4 | 29 | 46 | 37 | 20 |
Bellis perennis | 31.1 | 7 | 26 | 43 | 41 | 15 |
Achillea millefolium | 25.6 | 2 | 24 | 24 | 41 | 12 |
Hypochaeris radicata | 24.8 | 15 | 19 | 30 | 40 | 9 |
Plantago media | 24.7 | . | 31 | 24 | 30 | 9 |
Galium pumilum | 23.7 | 2 | 40 | 8 | 15 | 34 |
Briza media subsp. media | 22.2 | . | 38 | 25 | 9 | 9 |
Brachypodium rupestre | 21.5 | . | 32 | 19 | 18 | 10 |
Ranunculus bulbosus subsp. bulbosus | 21.5 | 2 | 28 | 20 | 24 | 11 |
Cerastium fontanum subsp. vulgare | 17.7 | . | 11 | 22 | 35 | 6 |
Daucus carota | 16.1 | 19 | 22 | 32 | . | . |
Linum catharticum subsp. catharticum | 15.7 | . | 28 | 3 | 13 | 18 |
Galium verum subsp. verum | 14.3 | 13 | 20 | 12 | 15 | 2 |
Potentilla montana | 13.6 | . | 17 | 1 | 26 | 10 |
Cynosurus cristatus | 13.0 | . | 13 | 27 | 10 | . |
Prunella vulgaris | 11.5 | . | 7 | 26 | 12 | 3 |
Helianthemum nummularium | 11.3 | . | 21 | <1 | 8 | 15 |
Blackstonia perfoliata | 11.0 | 17 | 25 | 5 | . | . |
Hippocrepis comosa | 10.9 | 2 | 20 | <1 | 5 | 18 |
Leontodon saxatilis subsp. saxatilis | 10.9 | 19 | 12 | 15 | 9 | . |
Colchicum montanum | 10.8 | . | 20 | 1 | 11 | 5 |
Trifolium campestre | 10.8 | 13 | 19 | 12 | 3 | . |
Phleum pratense | 10.5 | 11 | 17 | 15 | 1 | . |
Erica vagans | 10.4 | . | 17 | . | 17 | 4 |
[…] | ||||||
Bryophytes and lichens (based on plots from the Field Workshop) | ||||||
Class LYG (12 taxa) | ||||||
Seirophora lacunosa | 5.9 | 37 | . | . | . | . |
Gyalolechia fulgens | 5.9 | 37 | . | . | . | . |
Didymodon acutus | 23.5 | 58 | 25 | . | . | 6 |
Squamarina cartilaginea | 6.7 | 37 | . | . | . | 6 |
Weissia condensa | 7.6 | 32 | 3 | . | . | 6 |
Fulgensia poeltii | 4.2 | 21 | 2 | . | . | . |
Lathagrium cristatum | 4.2 | 21 | 2 | . | . | . |
Enchylium tenax | 10.1 | 32 | 8 | . | . | 6 |
[…] | ||||||
Class FES (5 taxa) | ||||||
Cladonia rangiformis | 14.3 | . | 27 | . | . | . |
Cladonia convoluta | 8.4 | . | 16 | . | . | . |
Eurhynchiastrum pulchellum | 5.9 | . | 11 | . | . | . |
Campyliadelphus chrysophyllus | 12.6 | . | 20 | . | . | 12 |
Cladonia foliacea | 4.2 | . | 8 | . | . | . |
Class MOL (2 taxa) | ||||||
Brachythecium laetum | 4.2 | . | . | 50 | 9 | . |
Rhytidiadelphus squarrosus | 2.5 | . | 3 | 13 | . | . |
Class NAR (3 taxa) | ||||||
Tortula acaulon | 1.7 | . | . | . | 18 | . |
Lophocolea minor | 0.8 | . | . | . | 9 | . |
Tortula inermis | 0.8 | . | . | . | 9 | . |
Class SES (20 taxa) | ||||||
Cladonia pocillum | 5.9 | . | 3 | . | . | 29 |
Tortella tortuosa | 20.2 | . | 23 | . | 9 | 47 |
Fissidens dubius | 15.1 | . | 19 | . | . | 35 |
Mnium marginatum | 2.5 | . | . | . | . | 18 |
Polytrichum juniperinum | 2.5 | . | . | . | . | 18 |
Scapania calcicola | 2.5 | . | . | . | . | 18 |
Tortella inclinata | 10.9 | 5 | 8 | . | 9 | 35 |
Ptychostomum capillare aggr. | 11.8 | 5 | 13 | . | . | 29 |
[…] | ||||||
Ditrichum pusillum | 10.1 | 11 | 6 | . | 9 | 29 |
Diagnostic for multiple classes | ||||||
Tortella squarrosa | 31.1 | 32 | 41 | . | . | 29 |
Abietinella abietina | 10.9 | . | 19 | 13 | . | . |
Ctenidium molluscum | 31.1 | . | 44 | 13 | . | 47 |
Flexitrichum gracile | 21.0 | . | 33 | . | . | 24 |
Companion species | ||||||
Homalothecium lutescens | 34.5 | 16 | 42 | 50 | 9 | 35 |
Weissia controversa | 23.5 | 42 | 23 | 13 | 9 | 18 |
Hypnum cupressiforme | 22.7 | 5 | 28 | . | 36 | 24 |
Pseudoscleropodium purum | 13.4 | . | 20 | 13 | 9 | 6 |
Calliergonella cuspidata | 12.6 | . | 20 | 13 | 9 | . |
After applying the expert system most relevés of Festuca-Brometea, Festuco-Ononidetea and Ononido-Rosmarinetea were classified in the FES group (Table
This group (FES) occupies the transition areas between the Pyrenees and Cantabrian mountains and the Mediterranean region (Figure
Comparison of climatic, structural, ecological and diversity characteristics among the five classes. The p-values and significance levels refer to ANOVAs.
Parameter | LYG | FES | MOL | NAR | SES | p-value | Sig. |
Total number of relevés | 54 | 339 | 220 | 223 | 114 | ||
Number of relevés from EDGG FW | 19 | 64 | 8 | 11 | 17 | ||
Mean ± SD | Mean ± SD | Mean ± SD | Mean ± SD | Mean ± SD | |||
Parameters calculated for all relevés | |||||||
Geographical and climatic parameters | |||||||
Elevation [m a.s.l.] | 439 ± 157 | 853 ± 286 | 577 ± 272 | 1265 ± 378 | 1752 ± 386 | <0.001 | *** |
Mediterranity index | 1.36 ± 0.46 | 0.66 ± 0.19 | 0.77 ± 0.31 | 0.41 ± 0.1 | 0.36 ± 0.08 | <0.001 | *** |
Annual mean temperature [°C] | 13.2 ± 1.3 | 10.5 ± 1.5 | 11.8 ± 1.6 | 8.0 ± 2.4 | 5.3 ± 2.7 | <0.001 | *** |
Mean annual precepitation [mm] | 686 ± 260 | 1232 ± 283 | 1134 ± 331 | 1751 ± 271 | 1865 ± 232 | <0.001 | *** |
Parameters calculated for relevés from EDGG Field Workshop | |||||||
Vegetation structure | |||||||
Cover vegetation total [%] | 67 ± 22 | 81 ± 19 | 98 ± 2 | 86 ± 9 | 55 ± 22 | <0.001 | *** |
Cover shrub layer [%] | 1 ± 1 | 1 ± 3 | 0 ± 0 | 0 ± 0 | 0 ± 0 | 0.138 | |
Cover herb layer [%] | 55 ± 25 | 76 ± 20 | 98 ± 2 | 77 ± 25 | 51 ± 22 | <0.001 | *** |
Cover cryptogam layer [%] | 19 ± 21 | 16 ± 18 | 31 ± 32 | 1 ± 2 | 10 ± 10 | 0.005 | ** |
Cover litter [%] | 16 ± 17 | 9 ± 14 | 8 ± 12 | 6 ± 6 | 14 ± 25 | 0.365 | n.s. |
Herb layer maximum height [cm] | 66 ± 26 | 65 ± 31 | 108 ± 32 | 31 ± 17 | 24 ± 19 | <0.001 | *** |
Species richness | |||||||
Species richness (total) | 35.6 ± 6.8 | 55.3 ± 14.5 | 45.3 ± 14.7 | 40.5 ± 6.9 | 44.0 ± 11.7 | <0.001 | *** |
Species richness (vascular plants) | 29.2 ± 7.5 | 48.0 ± 11.9 | 43.5 ± 14.0 | 37.5 ± 6.4 | 34.4 ± 7.7 | <0.001 | *** |
Species richness (cryptogams) | 6.4 ± 4.2 | 7.3 ± 4.9 | 2.0 ± 1.7 | 2.9 ± 2.0 | 9.6 ± 6.2 | <0.001 | *** |
Species richness (bryophytes) | 3.2 ± 2.0 | 6.3 ± 4.2 | 2.0 ± 1.7 | 2.5 ± 1.6 | 7.2 ± 5.6 | <0.001 | *** |
Species richness (lichens) | 3.2 ± 3.2 | 1.0 ± 1.3 | 0.0 ± 0.0 | 0.4 ± 0.7 | 2.4 ± 2.3 | <0.001 | *** |
Topography | |||||||
Southing (cosine of aspect) | 0.1 ± 0.6 | -0.3 ± 0.68 | -0.46 ± 0.65 | 0.24 ± 0.69 | 0.08 ± 0.89 | 0.019 | * |
Inclination [°] | 8 ± 9 | 16 ± 13 | 6 ± 6 | 26 ± 9 | 32 ± 11 | <0.001 | *** |
Maximum microrelief [cm] | 7 ± 7 | 9 ± 8 | 4 ± 3 | 9 ± 4 | 29 ± 26 | <0.001 | *** |
Soil parameters | |||||||
Soil depth mean [cm] | 12 ± 6 | 16 ± 8 | 17 ± 5 | 36 ± 16 | 6 ± 5 | <0.001 | *** |
Soil depth CV | 54 ± 32 | 50 ± 40 | 49 ± 34 | 30 ± 16 | 97 ± 51 | 0.001 | *** |
Cover rocks and stones [%] | 6 ± 13 | 7 ± 14 | 0 ± 0 | 2 ± 3 | 35 ± 23 | <0.001 | *** |
Cover gravel [%] | 19 ± 29 | 6 ± 15 | 0 ± 0 | 1 ± 1 | 13 ± 16 | 0.011 | * |
Cover fine soil [%] | 75 ± 35 | 88 ± 22 | 100 ± 0 | 97 ± 3 | 52 ± 32 | <0.001 | *** |
Coarse fragments [%] | 16 ± 13 | 22 ± 17 | 15 ± 14 | 12 ± 8 | 24 ± 16 | 0.139 | n.s. |
Fine fragments < 2mm [%] | 84 ± 13 | 78 ± 17 | 85 ± 14 | 88 ± 8 | 76 ± 16 | 0.139 | n.s. |
pH | 7.69 ± 0.24 | 7.52 ± 0.42 | 7.66 ± 0.99 | 6.8 ± 0.29 | 7.46 ± 0.38 | <0.001 | *** |
Electrical conductivity [µS/cm] | 283 ± 184 | 232 ± 86 | 168 ± 78 | 146 ± 80 | 310 ± 158 | 0.002 | ** |
CaCO3 [%] | 40.7 ± 10.5 | 26.7 ± 19.1 | 8.5 ± 8.5 | 4 ± 1.1 | 4.6 ± 1.8 | <0.001 | *** |
Organic matter [%] | 0.6 ± 0.6 | 1.4 ± 0.8 | 1.2 ± 0.3 | 1.3 ± 0.2 | 2.2 ± 0.7 | <0.001 | *** |
86% of the relevés previously assigned to the Molinio-Arrhenetheretea were included in the group MOL, together with 16% of the relevés of Festuco-Brometea. This group is characterized by several diagnostic species of the class Molinio-Arrhenatheretea, such as Agrostis stolonifera subsp. stolonifera, Anthoxanthum odoratum, Holcus lanatus, Juncus articulatus, J. inflexus, Lolium perenne, Poa trivialis subsp. trivialis, Ranunculus acris subsp. friesianus, R. repens and Trifolium fragiferum subsp. fragiferum, among other species (Table
The relevés from Festuco-Brometea class classified in the group MOL had been originally assigned to the associations Seseli-Brachypodietum and Elytrigio-Brachypodietum phoenicoidis from Festuco-Brometea. The presence of Agrimonia eupatoria, Agrostis stolonifera subsp. stolonifera, Bromus hordeaceus subsp. hordeaceus, Poa trivialis subsp. trivialis, Potentilla reptans, Ranunculus acris subsp. friesianus and Schedonorus arundinaceus subsp. arundinaceus relates these relevés to this group (MOL).
This group is widely distributed throughout the study area (Figure
Table
Relevés from Festuco-Brometea included in this group correspond to communities of the association Calamintho-Seselietum montani that grow in places with a very humid ombroclimate, which causes acidification of the soil leading to the presence of acidophilous species diagnostic of Nardetea. As regards Molinio-Arrhenatheretea, relevés originally assigned to the association Merendero-Cynosuretum were classified in this group. In both cases, the species shared with Nardetea were Agrostis capillaris, Carex pilulifera subsp. pilulifera, Danthonia decumbens, Festuca microphylla, Galium saxatile, Helictochloa marginata subsp. marginata, Luzula campestris, Jasione laevis subsp. laevis, Polygala serpyllifolia, Potentilla erecta, among others.
The relevés of this group are widely distributed in the montane and subalpine belts of the Pyrenees and Basque-Cantabrian mountains under temperate climate (Figure
The expert system classification within the group SES included most of the relevés of the class Elyno-Seslerietea and 23% of relevés from Festuco-Ononidetea. Agrostis schleicheri, Alchemilla plicatula aggr., Androsace villosa subsp. villosa, Carex ornithopoda subsp. ornithopoda, C. sempervirens subsp. sempervirens, Festuca gautieri subsp. scoparia, Helictotrichon sedenense subsp. sedenense, Paronychia kapela subsp. serpyllifolia, Poa alpina, Ranunculus carinthiacus, Sesleria caerulea subsp. caerulea, Silene acaulis and Trifolium thalii are diagnostic species of this group (Table
Relevés of Festuco-Ononidetea included in this group correspond to communities of the Pyrenean subalpine alliance Festucion scopariae, which share most of the diagnostic species of the group, such as Aster alpinus, Minuartia verna subsp. verna, and Saxifraga paniculata, in addition to those aforementioned.
This group SES includes the plots at highest elevations in calcareous mountains, in the upper montane and subalpine belts. In these cases, they share territories with the previous group NAR, but in rocky calcareous places (Figure
The NMDS ordination diagram clearly differentiated between the five groups defined by our class expert system (Figure
The differences between classes regarding elevation and climatic conditions can be seen in Table
Comparison of nine ecological variables among the five classes. For elevation and Mediterraneity Index, all relevés were analysed, whereas for the rest of variables only relevés from EDGG Field Workshop were used. Letters represent homogeneous groups (at α = 0.05) according to Tukey’s post-hoc test following a significant ANOVA.
Regarding soil, topographic and structural variables (Table
The total species richness is highest in Festuco-Brometea, although differences with the second richer class Molinio-Arrhenatheretea are not significant (Figure
Comparison of species richness divided into four groups (total species, vascular plants, bryophytes and lichens) among the five classes using the relevés from EDGG Field Workshop. Letters represent homogeneous groups (at α = 0.05) according to Tukey’s post-hoc test following a significant ANOVA.
The TWISPAN analysis for the group FES related to the class Festuco-Brometea resulted in four main divisions that can be interpreted at order and alliance levels (Figure
Diagnostic species for order 1 were Carex humilis, Galium lucidum subsp. fruticescens, Helianthemum apenninum subsp. apenninum and Koeleria vallesiana (Table
Abridged constancy table of the class Festuco-Brometea and its subordinate syntaxa. Values are percentage constancies, and species are ordered by decreasing phi-values in the respective syntaxon, respectively by decreasing overall constancy for non-diagnostic species. In the upper part vascular plants are given, in the lower part bryophytes and lichens, whose constancies and fidelities have been calculated based on the plots of the EDGG Field Workshop only where they have been recorded (in italics if based on data from a single plot, with ? if no such data were available for any plot). In the table, the 15 vascular plant taxa and the eight non-vascular plant taxa with the highest fidelity in a syntaxon are shown, plus all taxa that are diagnostic for multiple syntaxa and all taxa with at least 10% overall constancy. Diagnostic species (phi ≥ 0.25) for higher syntaxa highlighted in light grey, diagnostic species for associations in dark grey, while differential species of associations within the respective alliance are given with a frame. The complete constancy table combined with the table of the underlying 339 vegetation plots is given in Suppl. material
Class | Class | Ord. | Ord. | All. | All. | All. | All. | Assoc. | Assoc. | Assoc. | Assoc. | Assoc. | Assoc. | Assoc. | Assoc. |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Order | 1 | 2 | 1 | 1 | 2 | 2 | 1 | 1 | 1 | 2 | 2 | 2 | 2 | 2 | |
Alliance | 1.1 | 1.2 | 2.1 | 2.2 | 1.1 | 1.2 | 1.2 | 2.1 | 2.1 | 2.2 | 2.2 | 2.2 | |||
Association | 1.1.1 | 1.2.1 | 1.2.2 | 2.1.1 | 2.1.2 | 2.2.1 | 2.2.2 | 2.2.3 | |||||||
# plots | 339 | 139 | 200 | 52 | 87 | 40 | 160 | 52 | 25 | 61 | 14 | 26 | 12 | 78 | 69 |
Field Workshop (with bryophytes + lichens) | 64 | 23 | 41 | 18 | 5 | 8 | 33 | 18 | <1 | 5 | 1 | 7 | 1 | 20 | 12 |
Ord. 1 (4 taxa) | |||||||||||||||
Koeleria vallesiana | 53.1 | 92 | 26 | 85 | 97 | 15 | 29 | 85 | 100 | 95 | 36 | 4 | <1 | 27 | 36 |
Carex humilis | 40.1 | 66 | 22 | 65 | 67 | 8 | 26 | 65 | 96 | 56 | 14 | 4 | 42 | 28 | 19 |
Helianthemum apenninum subsp. apenninum | 13.6 | 29 | 3 | 19 | 34 | 3 | 3 | 19 | 32 | 36 | <1 | 4 | <1 | 3 | 4 |
Galium lucidum subsp. fruticescens | 9.7 | 21 | 2 | 15 | 24 | 3 | 2 | 15 | 24 | 25 | <1 | 4 | 17 | <1 | 1 |
All. 1.1 (38 taxa + 1 multiple diagnostic taxon) | |||||||||||||||
Brachypodium retusum | 14.5 | 29 | 5 | 75 | 1 | 18 | 1 | 75 | <1 | 2 | 43 | 4 | 8 | 1 | <1 |
Thymus vulgaris subsp. vulgaris | 20.1 | 45 | 3 | 81 | 24 | 8 | 1 | 81 | 44 | 16 | 7 | 8 | <1 | 1 | 1 |
Lavandula latifolia | 7.1 | 17 | 1 | 42 | 1 | 3 | <1 | 42 | 4 | <1 | <1 | 4 | <1 | <1 | <1 |
Aphyllanthes monspeliensis | 14.7 | 24 | 9 | 58 | 3 | 10 | 8 | 58 | 4 | 3 | 14 | 8 | <1 | 13 | 4 |
Coris monspeliensis | 6.8 | 14 | 2 | 38 | <1 | 8 | <1 | 38 | <1 | <1 | 14 | 4 | <1 | <1 | <1 |
Teucrium chamaedrys | 18.0 | 35 | 7 | 60 | 20 | 10 | 6 | 60 | 16 | 21 | 14 | 8 | 42 | 3 | 3 |
Helictochloa bromoides | 6.2 | 13 | 2 | 35 | <1 | 8 | <1 | 35 | <1 | <1 | 14 | 4 | <1 | <1 | <1 |
Bupleurum rigidum subsp. rigidum | 4.7 | 11 | 1 | 29 | <1 | 3 | <1 | 29 | <1 | <1 | 7 | <1 | <1 | <1 | <1 |
Linum appressum | 9.7 | 17 | 5 | 40 | 2 | 8 | 4 | 40 | 4 | 2 | 21 | <1 | 8 | 4 | 4 |
Genista scorpius | 15.0 | 23 | 10 | 58 | 2 | 35 | 3 | 58 | 4 | 2 | 50 | 27 | 8 | 4 | 1 |
Dorycnium pentaphyllum subsp. pentaphyllum | 16.5 | 24 | 12 | 58 | 3 | 33 | 6 | 58 | 4 | 3 | 64 | 15 | 33 | 5 | 3 |
Santolina villosa | 3.8 | 9 | 1 | 23 | <1 | <1 | 1 | 23 | <1 | <1 | <1 | <1 | 8 | <1 | <1 |
Coronilla minima | 38.3 | 60 | 23 | 79 | 49 | 25 | 23 | 79 | 72 | 41 | 36 | 19 | 8 | 29 | 17 |
Fumana ericifolia | 3.5 | 8 | 1 | 21 | <1 | <1 | 1 | 21 | <1 | <1 | <1 | <1 | <1 | <1 | 1 |
Linum narbonense | 6.2 | 12 | 3 | 29 | 1 | 5 | 2 | 29 | <1 | 2 | 14 | <1 | 8 | 3 | <1 |
[…] | |||||||||||||||
Asperula cynanchica | 17.7 | 22 | 15 | 44 | 9 | 8 | 16 | 44 | 4 | 11 | 14 | 4 | 17 | 18 | 14 |
Catananche caerulea | 13.6 | 17 | 12 | 42 | 1 | 23 | 9 | 42 | <1 | 2 | 21 | 23 | <1 | 17 | 1 |
Festuca marginata subsp. andres-molinae | 10.3 | 14 | 8 | 35 | 2 | 20 | 4 | 35 | 8 | <1 | 29 | 15 | <1 | 9 | <1 |
Assoc. 1.1.1 (7 taxa) | |||||||||||||||
Rhamnus alaternus subsp. alaternus | 1.2 | 3 | <1 | 8 | <1 | <1 | <1 | 8 | <1 | <1 | <1 | <1 | <1 | <1 | <1 |
Odontites kaliformis | 1.2 | 3 | <1 | 8 | <1 | <1 | <1 | 8 | <1 | <1 | <1 | <1 | <1 | <1 | <1 |
Lithodora fruticosa | 1.2 | 3 | <1 | 8 | <1 | <1 | <1 | 8 | <1 | <1 | <1 | <1 | <1 | <1 | <1 |
Gladiolus illyricus | 1.2 | 3 | <1 | 8 | <1 | <1 | <1 | 8 | <1 | <1 | <1 | <1 | <1 | <1 | <1 |
Atractylis humilis | 1.2 | 3 | <1 | 8 | <1 | <1 | <1 | 8 | <1 | <1 | <1 | <1 | <1 | <1 | <1 |
Aster willkommii | 1.2 | 3 | <1 | 8 | <1 | <1 | <1 | 8 | <1 | <1 | <1 | <1 | <1 | <1 | <1 |
Helichrysum stoechas subsp. stoechas | 1.8 | 4 | <1 | 10 | 1 | <1 | <1 | 10 | <1 | 2 | <1 | <1 | <1 | <1 | <1 |
All. 1.2 (8 taxa + 1 multiple diagnostic) | |||||||||||||||
Sedum album | 14.2 | 27 | 6 | 2 | 41 | <1 | 7 | 2 | 48 | 39 | <1 | <1 | 8 | <1 | 14 |
Ononis striata | 4.7 | 12 | <1 | 2 | 17 | <1 | <1 | 2 | 20 | 16 | <1 | <1 | <1 | <1 | <1 |
Asperula pyrenaica | 12.7 | 19 | 8 | 4 | 29 | 3 | 9 | 4 | 28 | 30 | <1 | 4 | <1 | 4 | 17 |
Plantago atrata subsp. discolor | 5.6 | 12 | 2 | 2 | 17 | <1 | 2 | 2 | 12 | 20 | <1 | <1 | <1 | 3 | 1 |
Anthyllis vulneraria | 22.1 | 29 | 18 | 8 | 41 | 15 | 18 | 8 | 40 | 43 | 14 | 15 | <1 | 15 | 23 |
Thymus praecox | 62.8 | 60 | 65 | 33 | 76 | 23 | 76 | 33 | 80 | 74 | 21 | 23 | 42 | 67 | 91 |
Oreochloa confusa | 3.8 | 7 | 2 | <1 | 11 | <1 | 2 | <1 | 16 | 8 | <1 | <1 | <1 | <1 | 4 |
Dianthus pungens subsp. brachyanthus | 3.2 | 6 | 1 | <1 | 10 | <1 | 1 | <1 | 16 | 8 | <1 | <1 | <1 | <1 | 3 |
Brimeura amethystina | 3.2 | 6 | 1 | <1 | 10 | <1 | 1 | <1 | 4 | 13 | <1 | <1 | <1 | 1 | 1 |
Assoc. 1.2.1 (13 taxa) | |||||||||||||||
Arenaria grandiflora subsp. grandiflora | 13.0 | 29 | 2 | 4 | 44 | <1 | 3 | 4 | 88 | 25 | <1 | <1 | <1 | <1 | 6 |
Festuca hystrix | 5.9 | 14 | <1 | 4 | 21 | <1 | <1 | 4 | 56 | 7 | <1 | <1 | <1 | <1 | <1 |
Helianthemum canum subsp. canum | 26.8 | 49 | 12 | 12 | 71 | <1 | 14 | 12 | 100 | 59 | <1 | <1 | <1 | 12 | 20 |
Klasea nudicaulis | 6.5 | 15 | 1 | 4 | 22 | <1 | 1 | 4 | 48 | 11 | <1 | <1 | <1 | <1 | 1 |
Anthyllis montana | 2.9 | 7 | <1 | <1 | 11 | <1 | <1 | <1 | 32 | 3 | <1 | <1 | <1 | <1 | <1 |
Jurinea humilis | 3.5 | 8 | 1 | <1 | 13 | <1 | 1 | <1 | 28 | 7 | <1 | <1 | <1 | <1 | 1 |
Erucastrum nasturtiifolium subsp. sudrei | 1.8 | 4 | <1 | <1 | 7 | <1 | <1 | <1 | 20 | 2 | <1 | <1 | <1 | <1 | <1 |
Trinia glauca | 7.7 | 17 | 1 | 10 | 22 | <1 | 1 | 10 | 36 | 16 | <1 | <1 | <1 | 3 | <1 |
Festuca ovina aggr. | 11.2 | 18 | 7 | 8 | 24 | <1 | 8 | 8 | 44 | 16 | <1 | <1 | 8 | 8 | 9 |
Aster alpinus | 1.2 | 3 | <1 | <1 | 5 | <1 | <1 | <1 | 16 | <1 | <1 | <1 | <1 | <1 | <1 |
Paronychia kapela subsp. kapela | 0.9 | 2 | <1 | <1 | 3 | <1 | <1 | <1 | 12 | <1 | <1 | <1 | <1 | <1 | <1 |
Arenaria erinacea | 0.9 | 2 | <1 | <1 | 3 | <1 | <1 | <1 | 12 | <1 | <1 | <1 | <1 | <1 | <1 |
Scilla verna | 6.8 | 10 | 5 | 4 | 14 | 3 | 5 | 4 | 28 | 8 | <1 | 4 | <1 | 5 | 4 |
Assoc. 1.2.2 (5 taxa + 1 multiple diagnostic taxon + 1 differential taxon) | |||||||||||||||
Conopodium arvense | 4.4 | 9 | 2 | <1 | 14 | <1 | 2 | <1 | <1 | 20 | <1 | <1 | <1 | 3 | 1 |
Sedum acre | 2.9 | 6 | 1 | <1 | 10 | <1 | 1 | <1 | <1 | 15 | <1 | <1 | <1 | <1 | 1 |
Deschampsia media subsp. hispanica | 3.2 | 7 | 1 | <1 | 11 | 3 | <1 | <1 | <1 | 16 | <1 | 4 | <1 | <1 | <1 |
Helianthemum salicifolium | 1.5 | 4 | <1 | <1 | 6 | <1 | <1 | <1 | <1 | 8 | <1 | <1 | <1 | <1 | <1 |
Erodium glandulosum | 1.5 | 4 | <1 | <1 | 6 | <1 | <1 | <1 | <1 | 8 | <1 | <1 | <1 | <1 | <1 |
Medicago lupulina | 38.9 | 33 | 43 | 12 | 46 | 33 | 46 | 12 | 4 | 64 | 21 | 38 | 8 | 54 | 43 |
Ord. 2 (12 taxa) | |||||||||||||||
Lotus corniculatus | 52.8 | 24 | 73 | 13 | 30 | 58 | 77 | 13 | 20 | 34 | 57 | 58 | 17 | 85 | 78 |
Trifolium pratense | 28.6 | 4 | 46 | 2 | 6 | 45 | 46 | 2 | <1 | 8 | 29 | 54 | 8 | 62 | 35 |
Briza media subsp. media | 38.3 | 17 | 54 | 25 | 11 | 45 | 56 | 25 | 4 | 15 | 43 | 46 | 33 | 83 | 29 |
Trifolium ochroleucon | 15.6 | 1 | 26 | <1 | 2 | 20 | 27 | <1 | <1 | 3 | 7 | 27 | <1 | 47 | 9 |
Trifolium repens | 16.5 | 2 | 27 | 2 | 2 | 20 | 28 | 2 | <1 | 3 | <1 | 31 | <1 | 27 | 35 |
Cynosurus cristatus | 12.7 | <1 | 22 | <1 | <1 | 28 | 20 | <1 | <1 | <1 | 7 | 38 | <1 | 35 | 7 |
Ranunculus bulbosus subsp. bulbosus | 28.0 | 12 | 40 | 17 | 8 | 45 | 38 | 17 | 4 | 10 | 21 | 58 | 8 | 44 | 38 |
Plantago lanceolata | 50.4 | 32 | 64 | 23 | 37 | 50 | 67 | 23 | 12 | 48 | 43 | 54 | <1 | 73 | 72 |
Endressia castellana | 9.4 | <1 | 16 | <1 | <1 | 10 | 18 | <1 | <1 | <1 | <1 | 15 | 25 | 23 | 10 |
Ononis spinosa | 13.0 | 2 | 21 | 2 | 2 | 28 | 19 | 2 | <1 | 3 | 7 | 38 | <1 | 35 | 4 |
Trisetum flavescens subsp. flavescens | 11.8 | 2 | 19 | 4 | 1 | 20 | 18 | 4 | <1 | 2 | <1 | 31 | <1 | 31 | 7 |
Festuca nigrescens | 9.1 | 1 | 15 | <1 | 1 | 13 | 16 | <1 | <1 | 2 | <1 | 19 | 17 | 24 | 4 |
All. 2.1 (11 taxa + 1 multiple diagnostic taxon) | |||||||||||||||
Schedonorus arundinaceus subsp. fenas | 5.9 | <1 | 10 | <1 | <1 | 38 | 3 | <1 | <1 | <1 | 36 | 38 | <1 | 5 | 1 |
Centaurea jacea | 17.1 | 6 | 25 | 13 | 2 | 55 | 17 | 13 | <1 | 3 | 57 | 54 | 17 | 29 | 3 |
Arrhenatherum elatius | 6.2 | 1 | 10 | 4 | <1 | 33 | 4 | 4 | <1 | <1 | 29 | 35 | 33 | 3 | <1 |
Convolvulus arvensis | 4.4 | 1 | 7 | 2 | <1 | 28 | 2 | 2 | <1 | <1 | 21 | 31 | <1 | 3 | 1 |
Brachypodium phoenicoides | 10.0 | 8 | 12 | 19 | 1 | 43 | 4 | 19 | <1 | 2 | 43 | 42 | <1 | 4 | 4 |
Blackstonia perfoliata | 24.8 | 23 | 26 | 58 | 2 | 68 | 16 | 58 | <1 | 3 | 79 | 62 | <1 | 27 | 6 |
Daucus carota | 21.8 | 10 | 30 | 17 | 6 | 53 | 24 | 17 | <1 | 8 | 36 | 62 | 17 | 44 | 4 |
Carex flacca subsp. flacca | 45.1 | 26 | 59 | 58 | 7 | 80 | 53 | 58 | 4 | 8 | 86 | 77 | 58 | 81 | 22 |
Agrimonia eupatoria | 4.1 | 1 | 7 | 2 | <1 | 20 | 3 | 2 | <1 | <1 | 14 | 23 | <1 | 6 | <1 |
Poa compressa | 3.2 | 1 | 5 | <1 | 1 | 18 | 2 | <1 | <1 | 2 | 7 | 23 | <1 | 4 | <1 |
Dactylis glomerata | 19.8 | 15 | 23 | 27 | 8 | 48 | 17 | 27 | 4 | 10 | 29 | 58 | 58 | 21 | 6 |
Medicago sativa subsp. sativa | 1.2 | <1 | 2 | <1 | <1 | 10 | <1 | <1 | <1 | <1 | 7 | 12 | <1 | <1 | <1 |
Assoc. 2.1.1 (7 taxa + 2 differential taxa) | |||||||||||||||
Plantago maritima subsp. serpentina | 6.8 | 6 | 7 | 15 | 1 | 25 | 3 | 15 | <1 | 2 | 57 | 8 | <1 | 3 | 3 |
Festuca capillifolia | 5.3 | 4 | 7 | 6 | 2 | 28 | 1 | 6 | 4 | 2 | 50 | 15 | <1 | 3 | <1 |
Jasonia tuberosa | 6.2 | 9 | 4 | 21 | 2 | 15 | 1 | 21 | <1 | 3 | 43 | <1 | <1 | 1 | 1 |
Prunella hyssopifolia | 11.5 | 8 | 14 | 19 | 1 | 35 | 9 | 19 | <1 | 2 | 57 | 23 | <1 | 15 | 3 |
Agrostis stolonifera subsp. stolonifera | 2.7 | <1 | 5 | <1 | <1 | 18 | 1 | <1 | <1 | <1 | 29 | 12 | <1 | 3 | <1 |
Lotus tenuis | 0.6 | <1 | 1 | <1 | <1 | 5 | <1 | <1 | <1 | <1 | 14 | <1 | <1 | <1 | <1 |
Lathyrus latifolius | 0.6 | <1 | 1 | <1 | <1 | 5 | <1 | <1 | <1 | <1 | 14 | <1 | <1 | <1 | <1 |
Assoc. 2.1.2 (28 taxa + 3 differential taxa) | |||||||||||||||
Phleum pratense | 17.1 | 4 | 26 | 10 | 1 | 53 | 19 | 10 | <1 | 2 | 14 | 73 | <1 | 28 | 13 |
Poa trivialis subsp. trivialis | 5.6 | <1 | 10 | <1 | <1 | 28 | 5 | <1 | <1 | <1 | <1 | 42 | <1 | 8 | 3 |
Vicia parviflora | 3.2 | <1 | 6 | <1 | <1 | 23 | 1 | <1 | <1 | <1 | <1 | 35 | <1 | 1 | 1 |
Brachypodium phoenicoides x rupestre | 9.4 | 2 | 15 | 6 | <1 | 40 | 8 | 6 | <1 | <1 | 7 | 58 | 8 | 12 | 4 |
Xeranthemum cylindraceum | 2.7 | 1 | 4 | 2 | <1 | 18 | 1 | 2 | <1 | <1 | <1 | 27 | <1 | <1 | 1 |
Trifolium campestre | 18.6 | 7 | 27 | 6 | 8 | 40 | 23 | 6 | 8 | 8 | <1 | 62 | <1 | 24 | 26 |
Vicia sativa subsp. nigra | 5.9 | 2 | 9 | 2 | 2 | 25 | 4 | 2 | <1 | 3 | <1 | 38 | 8 | 6 | 1 |
Iris spuria subsp. maritima | 1.8 | <1 | 3 | <1 | <1 | 13 | 1 | <1 | <1 | <1 | <1 | 19 | <1 | 1 | <1 |
Gaudinia fragilis | 2.1 | <1 | 4 | <1 | <1 | 13 | 1 | <1 | <1 | <1 | <1 | 19 | <1 | 3 | <1 |
Trifolium angustifolium | 2.4 | 1 | 4 | 2 | <1 | 13 | 1 | 2 | <1 | <1 | <1 | 19 | <1 | <1 | 3 |
Lathyrus pratensis subsp. pratensis | 1.5 | <1 | 3 | <1 | <1 | 10 | 1 | <1 | <1 | <1 | <1 | 15 | <1 | 1 | <1 |
Jacobaea vulgaris | 3.2 | <1 | 6 | <1 | <1 | 18 | 3 | <1 | <1 | <1 | 7 | 23 | <1 | 5 | <1 |
Allium oleraceum | 1.8 | <1 | 3 | <1 | <1 | 10 | 1 | <1 | <1 | <1 | <1 | 15 | <1 | 1 | 1 |
Cerastium fontanum subsp. vulgare | 10.6 | 4 | 15 | <1 | 7 | 23 | 13 | <1 | 4 | 7 | <1 | 35 | <1 | 14 | 14 |
Picris hieracioides | 7.1 | 4 | 9 | 12 | <1 | 23 | 6 | 12 | <1 | <1 | 7 | 31 | <1 | 10 | 1 |
[…] | |||||||||||||||
Eryngium campestre | 32.4 | 29 | 35 | 58 | 13 | 58 | 29 | 58 | 8 | 15 | 29 | 73 | 25 | 31 | 28 |
Galium verum subsp. verum | 19.8 | 9 | 27 | 6 | 11 | 38 | 24 | 6 | <1 | 16 | 14 | 50 | 17 | 22 | 29 |
Anacamptis pyramidalis | 10.6 | 7 | 13 | 19 | <1 | 28 | 9 | 19 | <1 | <1 | 14 | 35 | <1 | 19 | <1 |
All. 2.2 (10 taxa + 1 multiple diagnostic taxon) | |||||||||||||||
Brachypodium rupestre | 32.4 | 12 | 47 | 12 | 11 | 3 | 58 | 12 | 4 | 15 | 7 | <1 | 92 | 73 | 36 |
Festuca microphylla | 22.4 | 4 | 35 | 2 | 6 | 3 | 43 | 2 | <1 | 8 | 7 | <1 | 25 | 46 | 43 |
Achillea millefolium | 24.5 | 7 | 37 | 6 | 8 | 13 | 43 | 6 | 12 | 7 | 7 | 15 | 8 | 47 | 42 |
Agrostis capillaris | 17.7 | 3 | 28 | <1 | 5 | 5 | 34 | <1 | 4 | 5 | <1 | 8 | 8 | 37 | 35 |
Erica vagans | 17.1 | 3 | 27 | 6 | 1 | 13 | 31 | 6 | 4 | <1 | <1 | 19 | 58 | 41 | 14 |
Potentilla montana | 16.8 | 3 | 27 | <1 | 5 | 8 | 31 | <1 | 4 | 5 | <1 | 12 | 25 | 24 | 39 |
Helianthemum nummularium | 21.2 | 7 | 31 | <1 | 11 | <1 | 39 | <1 | <1 | 16 | <1 | <1 | 17 | 36 | 46 |
Danthonia decumbens | 10.3 | 1 | 17 | <1 | 1 | 3 | 21 | <1 | <1 | 2 | <1 | 4 | <1 | 29 | 14 |
Scabiosa columbaria subsp. columbaria | 25.4 | 12 | 35 | 10 | 14 | 10 | 41 | 10 | <1 | 20 | 7 | 12 | 42 | 56 | 23 |
Gentiana verna subsp. verna | 5.6 | 1 | 9 | <1 | 1 | <1 | 11 | <1 | <1 | 2 | <1 | <1 | <1 | 9 | 16 |
Assoc. 2.2.1 (21 taxa + 1 multiple diagnostic taxon + 2 differential taxa) | |||||||||||||||
Vincetoxicum hirundinaria subsp. intermedium | 3.8 | 4 | 4 | 4 | 3 | <1 | 5 | 4 | 4 | 3 | <1 | <1 | 58 | 1 | <1 |
Sesleria autumnalis | 2.7 | 1 | 4 | <1 | 2 | <1 | 4 | <1 | <1 | 3 | <1 | <1 | 50 | 1 | <1 |
Tanacetum corymbosum subsp. corymbosum | 2.9 | 1 | 4 | 2 | 1 | <1 | 5 | 2 | <1 | 2 | <1 | <1 | 33 | 5 | <1 |
Genista hispanica subsp. occidentalis | 15.9 | 7 | 22 | 10 | 6 | 10 | 25 | 10 | 8 | 5 | 7 | 12 | 67 | 32 | 9 |
Euphorbia characias | 2.1 | 2 | 2 | 4 | 1 | <1 | 3 | 4 | <1 | 2 | <1 | <1 | 25 | <1 | 1 |
Euphorbia amygdaloides | 0.6 | <1 | 1 | <1 | <1 | <1 | 1 | <1 | <1 | <1 | <1 | <1 | 17 | <1 | <1 |
Cruciata laevipes | 0.6 | <1 | 1 | <1 | <1 | <1 | 1 | <1 | <1 | <1 | <1 | <1 | 17 | <1 | <1 |
Teucrium pyrenaicum | 23.9 | 22 | 25 | 10 | 30 | 3 | 31 | 10 | 28 | 30 | 7 | <1 | 67 | 29 | 26 |
Pimpinella major subsp. major | 1.2 | <1 | 2 | <1 | <1 | <1 | 3 | <1 | <1 | <1 | <1 | <1 | 17 | 1 | 1 |
Helictotrichon cantabricum | 6.8 | 6 | 8 | 12 | 2 | 13 | 6 | 12 | 4 | 2 | 21 | 8 | 42 | 5 | 1 |
Viola alba aggr. | 3.5 | 4 | 3 | 12 | <1 | <1 | 4 | 12 | <1 | <1 | <1 | <1 | 25 | 4 | <1 |
Dianthus hyssopifolius subsp. hyssopifolius | 4.1 | 3 | 5 | <1 | 5 | <1 | 6 | <1 | <1 | 7 | <1 | <1 | 25 | 3 | 7 |
Helleborus foetidus | 0.9 | <1 | 2 | <1 | <1 | 3 | 1 | <1 | <1 | <1 | <1 | 4 | 17 | <1 | <1 |
Senecio lagascanus | 1.8 | 3 | 1 | 4 | 2 | <1 | 1 | 4 | <1 | 3 | <1 | <1 | 17 | <1 | <1 |
Echium vulgare subsp. vulgare | 2.1 | 1 | 3 | 2 | 1 | <1 | 3 | 2 | <1 | 2 | <1 | <1 | 17 | 3 | 1 |
Assoc. 2.2.2 (4 taxa + 1 multiple diagnostic taxon + 2 differential taxa) | |||||||||||||||
Leontodon hispidus | 9.1 | 1 | 15 | 2 | 1 | <1 | 18 | 2 | <1 | 2 | <1 | <1 | <1 | 32 | 6 |
Plantago media | 31.3 | 9 | 47 | 8 | 9 | 25 | 53 | 8 | 8 | 10 | 14 | 31 | 8 | 72 | 39 |
Linum catharticum subsp. catharticum | 27.7 | 12 | 39 | 19 | 7 | 25 | 43 | 19 | <1 | 10 | 29 | 23 | <1 | 63 | 28 |
Polygala vulgaris subsp. vulgaris | 7.1 | 1 | 12 | 2 | <1 | 5 | 13 | 2 | <1 | <1 | <1 | 8 | <1 | 24 | 1 |
Prunella vulgaris | 6.8 | 1 | 11 | 2 | 1 | 5 | 12 | 2 | <1 | 2 | <1 | 8 | <1 | 21 | 4 |
Holcus lanatus | 2.9 | <1 | 5 | <1 | <1 | 3 | 6 | <1 | <1 | <1 | <1 | 4 | <1 | 12 | <1 |
Assoc. 2.2.3 (10 taxa + 1 multiple diagnostic taxon) | |||||||||||||||
Carex caryophyllea | 31.3 | 14 | 43 | 6 | 20 | 13 | 51 | 6 | 8 | 25 | 7 | 15 | 17 | 36 | 72 |
Festuca rectifolia | 42.5 | 45 | 41 | 6 | 68 | 8 | 49 | 6 | 36 | 80 | <1 | 12 | 17 | 26 | 81 |
Bellis perennis | 25.7 | 12 | 35 | 6 | 16 | 18 | 39 | 6 | <1 | 23 | 7 | 23 | <1 | 31 | 57 |
Colchicum montanum | 20.4 | 17 | 23 | 2 | 25 | 3 | 28 | 2 | 8 | 33 | <1 | 4 | <1 | 19 | 43 |
Alchemilla plicatula aggr. | 3.5 | 1 | 6 | <1 | 1 | <1 | 7 | <1 | <1 | 2 | <1 | <1 | <1 | 1 | 14 |
Aira caryophyllea subsp. caryophyllea | 9.1 | 5 | 12 | 2 | 7 | 8 | 13 | 2 | <1 | 10 | <1 | 12 | <1 | 4 | 26 |
Vicia pyrenaica | 2.9 | 1 | 4 | <1 | 2 | <1 | 5 | <1 | <1 | 2 | <1 | <1 | <1 | <1 | 12 |
Poa alpina | 4.7 | 3 | 6 | <1 | 5 | <1 | 8 | <1 | <1 | 7 | <1 | <1 | <1 | 1 | 16 |
Cerastium arvense | 8.3 | 9 | 8 | <1 | 14 | <1 | 10 | <1 | 8 | 15 | <1 | <1 | <1 | <1 | 23 |
Erinus alpinus | 3.8 | 3 | 5 | <1 | 5 | <1 | 6 | <1 | <1 | 5 | <1 | <1 | <1 | <1 | 13 |
Euphrasia salisburgensis | 2.1 | <1 | 4 | <1 | <1 | <1 | 4 | <1 | <1 | <1 | <1 | <1 | <1 | 1 | 9 |
Other species (class character species and companion species) | |||||||||||||||
Bromopsis erecta subsp. erecta | 65.2 | 60 | 69 | 69 | 55 | 53 | 73 | 69 | 32 | 66 | 29 | 65 | 75 | 82 | 61 |
Carthamus mitissimus | 50.7 | 56 | 47 | 58 | 55 | 23 | 53 | 58 | 44 | 61 | 14 | 27 | 33 | 63 | 45 |
Potentilla tabernaemontani | 48.1 | 56 | 43 | 52 | 59 | 20 | 48 | 52 | 36 | 69 | 21 | 19 | 8 | 36 | 68 |
Helictochloa pratensis subsp. iberica | 45.7 | 48 | 44 | 52 | 46 | 18 | 51 | 52 | 36 | 51 | 7 | 23 | 42 | 51 | 51 |
Pilosella officinarum | 41.3 | 31 | 49 | 40 | 25 | 23 | 55 | 40 | 8 | 33 | 29 | 19 | <1 | 59 | 61 |
Galium pumilum | 39.8 | 27 | 49 | 44 | 17 | 48 | 49 | 44 | 4 | 23 | 36 | 54 | 58 | 60 | 33 |
Sanguisorba minor aggr. | 33.9 | 29 | 37 | 44 | 21 | 20 | 41 | 44 | 8 | 26 | 14 | 23 | 25 | 54 | 30 |
Seseli montanum subsp. montanum | 32.7 | 32 | 34 | 23 | 37 | 35 | 33 | 23 | 32 | 39 | 21 | 42 | 8 | 32 | 38 |
Hippocrepis comosa | 20.4 | 12 | 27 | 13 | 10 | 10 | 31 | 13 | 8 | 11 | 14 | 8 | 8 | 41 | 22 |
Hypochaeris radicata | 18.9 | 10 | 25 | 15 | 7 | 25 | 25 | 15 | <1 | 10 | 43 | 15 | 17 | 32 | 19 |
Geum sylvaticum | 18.0 | 12 | 23 | 12 | 11 | 10 | 26 | 12 | 8 | 13 | 7 | 12 | 17 | 32 | 20 |
Onobrychis conferta subsp. hispanica | 16.8 | 20 | 15 | 31 | 14 | 8 | 16 | 31 | 16 | 13 | 14 | 4 | <1 | 28 | 6 |
Trifolium montanum subsp. montanum | 15.3 | 6 | 22 | 2 | 9 | 15 | 23 | 2 | 8 | 10 | 7 | 19 | <1 | 33 | 14 |
Clinopodium alpinum subsp. pyrenaeum | 14.2 | 19 | 11 | 13 | 23 | <1 | 13 | 13 | 8 | 30 | <1 | <1 | <1 | 10 | 19 |
Leucanthemum pallens | 13.6 | 9 | 17 | 19 | 3 | 23 | 15 | 19 | <1 | 5 | 14 | 27 | 17 | 26 | 3 |
Filipendula vulgaris | 12.7 | 6 | 17 | <1 | 10 | 13 | 18 | <1 | 12 | 10 | 7 | 15 | 33 | 13 | 20 |
Astragalus monspessulanus subsp. monspessulanus | 12.4 | 19 | 8 | 15 | 21 | 5 | 9 | 15 | 28 | 18 | 7 | 4 | <1 | 10 | 9 |
Leontodon saxatilis subsp. saxatilis | 12.1 | 8 | 15 | 8 | 8 | 25 | 13 | 8 | <1 | 11 | 21 | 27 | 8 | 12 | 14 |
Prunella laciniata | 12.1 | 4 | 18 | 8 | 1 | 8 | 21 | 8 | <1 | 2 | <1 | 12 | 8 | 26 | 17 |
Thymelaea ruizii | 11.5 | 12 | 12 | 21 | 6 | 10 | 12 | 21 | 4 | 7 | 14 | 8 | 8 | 17 | 7 |
Ononis pusilla | 10.3 | 19 | 4 | 25 | 16 | 5 | 4 | 25 | 8 | 20 | 14 | <1 | <1 | 3 | 6 |
Bellis sylvestris | 10.0 | 14 | 8 | 23 | 8 | 10 | 7 | 23 | 8 | 8 | 7 | 12 | <1 | 10 | 4 |
[…] | |||||||||||||||
Bryophytes and lichens (based on plots from the Field Workshop) | |||||||||||||||
Ord. 1 (1 taxon) | |||||||||||||||
Tortella squarrosa | 40.6 | 70 | 24 | 72 | 60 | 13 | 27 | 72 | ? | 60 | <1 | 14 | <1 | 20 | 42 |
All. 1.1 (1 taxon) | |||||||||||||||
Flexitrichum gracile | 32.8 | 48 | 24 | 61 | <1 | <1 | 30 | 61 | ? | <1 | <1 | <1 | <1 | 30 | 33 |
All. 1.2 (8 taxa) | |||||||||||||||
Cladonia foliacea | 7.8 | 13 | 5 | <1 | 60 | <1 | 6 | <1 | ? | 60 | <1 | <1 | <1 | <1 | 17 |
Didymodon acutus | 25.0 | 30 | 22 | 22 | 60 | <1 | 27 | 22 | ? | 60 | <1 | <1 | <1 | 40 | 8 |
Lathagrium cristatum | 1.6 | 4 | <1 | <1 | 20 | <1 | <1 | <1 | ? | 20 | <1 | <1 | <1 | <1 | <1 |
Pseudocrossidium hornschuchianum | 1.6 | 4 | <1 | <1 | 20 | <1 | <1 | <1 | ? | 20 | <1 | <1 | <1 | <1 | <1 |
Scytinium schraderi | 1.6 | 4 | <1 | <1 | 20 | <1 | <1 | <1 | ? | 20 | <1 | <1 | <1 | <1 | <1 |
Encalypta vulgaris | 3.1 | 4 | 2 | <1 | 20 | <1 | 3 | <1 | ? | 20 | <1 | <1 | <1 | <1 | 8 |
Didymodon vinealis | 4.7 | 9 | 2 | 6 | 20 | <1 | 3 | 6 | ? | 20 | <1 | <1 | <1 | 5 | <1 |
Ditrichum pusillum | 6.3 | 9 | 5 | 6 | 20 | <1 | 6 | 6 | ? | 20 | <1 | <1 | <1 | 5 | 8 |
Ord. 2 (4 taxa) | |||||||||||||||
Pseudoscleropodium purum | 20.3 | <1 | 32 | <1 | <1 | 25 | 33 | <1 | ? | <1 | <1 | 29 | <1 | 40 | 25 |
Cladonia rangiformis | 26.6 | 9 | 37 | 6 | 20 | 13 | 42 | 6 | ? | 20 | 100 | <1 | <1 | 20 | 83 |
Eurhynchiastrum pulchellum | 10.9 | <1 | 17 | <1 | <1 | 25 | 15 | <1 | ? | <1 | <1 | 29 | <1 | 20 | 8 |
Fissidens dubius | 18.8 | 4 | 27 | 6 | <1 | 13 | 30 | 6 | ? | <1 | 100 | <1 | 100 | 25 | 33 |
All. 2.1 (2 taxa) | |||||||||||||||
Calliergonella cuspidata | 20.3 | <1 | 32 | <1 | <1 | 50 | 27 | <1 | ? | <1 | 100 | 43 | <1 | 35 | 17 |
Weissia controversa | 23.4 | 17 | 27 | 17 | 20 | 50 | 21 | 17 | ? | 20 | 100 | 43 | 100 | 25 | 8 |
Assoc. 2.1.2 (6 taxa) | |||||||||||||||
Oxyrrhynchium hians | 6.3 | 4 | 7 | 6 | <1 | 38 | <1 | 6 | ? | <1 | <1 | 43 | <1 | <1 | <1 |
Fissidens taxifolius | 14.1 | 4 | 20 | 6 | <1 | 38 | 15 | 6 | ? | <1 | <1 | 43 | <1 | 25 | <1 |
Brachytheciastrum velutinum | 1.6 | <1 | 2 | <1 | <1 | 13 | <1 | <1 | ? | <1 | <1 | 14 | <1 | <1 | <1 |
Brachythecium rutabulum | 1.6 | <1 | 2 | <1 | <1 | 13 | <1 | <1 | ? | <1 | <1 | 14 | <1 | <1 | <1 |
Plagiomnium undulatum | 1.6 | <1 | 2 | <1 | <1 | 13 | <1 | <1 | ? | <1 | <1 | 14 | <1 | <1 | <1 |
Weissia condensa | 3.1 | 4 | 2 | 6 | <1 | 13 | <1 | 6 | ? | <1 | <1 | 14 | <1 | <1 | <1 |
All. 2.2 (6 taxa) | |||||||||||||||
Entodon concinnus | 12.5 | <1 | 20 | <1 | <1 | <1 | 24 | <1 | ? | <1 | <1 | <1 | <1 | 20 | 33 |
Abietinella abietina | 18.8 | 9 | 24 | 11 | <1 | <1 | 30 | 11 | ? | <1 | <1 | <1 | <1 | 30 | 33 |
Enchylium tenax | 7.8 | <1 | 12 | <1 | <1 | <1 | 15 | <1 | ? | <1 | <1 | <1 | <1 | 20 | 8 |
Cladonia convoluta | 15.6 | 9 | 20 | 11 | <1 | <1 | 24 | 11 | ? | <1 | <1 | <1 | <1 | 20 | 33 |
Thuidium assimile | 6.3 | <1 | 10 | <1 | <1 | <1 | 12 | <1 | ? | <1 | <1 | <1 | <1 | 10 | 17 |
Cladonia cariosa | 4.7 | <1 | 7 | <1 | <1 | <1 | 9 | <1 | ? | <1 | <1 | <1 | <1 | 10 | 8 |
Assoc. 2.2.2 (4 taxa) | |||||||||||||||
Barbula unguiculata | 6.3 | <1 | 10 | <1 | <1 | <1 | 12 | <1 | ? | <1 | <1 | <1 | <1 | 20 | <1 |
Campyliadelphus chrysophyllus | 20.3 | 13 | 24 | 17 | <1 | 13 | 27 | 17 | ? | <1 | <1 | 14 | <1 | 40 | 8 |
Rhytidiadelphus squarrosus | 3.1 | <1 | 5 | <1 | <1 | <1 | 6 | <1 | ? | <1 | <1 | <1 | <1 | 10 | <1 |
Thuidium delicatulum | 3.1 | <1 | 5 | <1 | <1 | <1 | 6 | <1 | ? | <1 | <1 | <1 | <1 | 10 | <1 |
Assoc. 2.2.3 (23 taxa) | |||||||||||||||
Exsertotheca crispa | 14.1 | <1 | 22 | <1 | <1 | <1 | 27 | <1 | ? | <1 | <1 | <1 | <1 | 15 | 50 |
Ptychostomum capillare aggr. | 12.5 | 4 | 17 | <1 | 20 | <1 | 21 | <1 | ? | 20 | <1 | <1 | <1 | 5 | 50 |
- Ptychostomum capillare | 7.8 | 4 | 10 | <1 | 20 | <1 | 12 | <1 | ? | 20 | <1 | <1 | <1 | 5 | 25 |
- Ptychostomum elegans | 6.3 | <1 | 10 | <1 | <1 | <1 | 12 | <1 | ? | <1 | <1 | <1 | <1 | <1 | 33 |
Tortella tortuosa | 23.4 | 17 | 27 | 17 | 20 | <1 | 33 | 17 | ? | 20 | <1 | <1 | <1 | 20 | 58 |
Hypnum cupressiforme | 28.1 | 17 | 34 | 17 | 20 | 13 | 39 | 17 | ? | 20 | <1 | 14 | <1 | 30 | 58 |
Cetraria islandica | 3.1 | <1 | 5 | <1 | <1 | <1 | 6 | <1 | ? | <1 | <1 | <1 | <1 | <1 | 17 |
Lophocolea heterophylla | 3.1 | <1 | 5 | <1 | <1 | <1 | 6 | <1 | ? | <1 | <1 | <1 | <1 | <1 | 17 |
Tortella inclinata | 7.8 | 9 | 7 | 11 | <1 | <1 | 9 | 11 | ? | <1 | <1 | <1 | <1 | <1 | 25 |
Bryum argenteum | 4.7 | <1 | 7 | <1 | <1 | <1 | 9 | <1 | ? | <1 | <1 | <1 | <1 | 5 | 17 |
[…] | |||||||||||||||
Other species (class character species and companion species) | |||||||||||||||
Ctenidium molluscum | 43.8 | 35 | 49 | 44 | <1 | 38 | 52 | 44 | ? | <1 | 100 | 29 | <1 | 55 | 50 |
Homalothecium lutescens | 42.2 | 35 | 46 | 44 | <1 | 63 | 42 | 44 | ? | <1 | 100 | 57 | <1 | 40 | 50 |
Syntrichia ruralis aggr. | 12.5 | 13 | 12 | 11 | 20 | <1 | 15 | 11 | ? | 20 | <1 | <1 | <1 | 5 | 33 |
[…] |
Climatic structural, ecological and diversity characteristics of the orders and alliances within the Festuco-Brometea. The p-values and significance levels refer to ANOVAs.
Parameter | Alliances | p-value | Sig. | |||
1.1 | 1.2 | 2.1 | 2.2 | |||
Total number of relevés | 52 | 87 | 40 | 160 | ||
Number of relevés from EDGG FW | 18 | 5 | 8 | 33 | ||
Mean ± SD | Mean ± SD | Mean ± SD | Mean ± SD | |||
Parameters calculated for all relevés | ||||||
Geographical and climatic parameters | ||||||
Elevation [m a.s.l.] | 602±151 | 1030±215 | 561±139 | 912±273 | <0.001 | *** |
Mediterranity index | 0.83±0.19 | 0.6±0.12 | 0.91±0.15 | 0.58±0.15 | <0.001 | *** |
Annual mean temperature [°C] | 11.9±1.0 | 9.9±1.0 | 12.1±0.7 | 10.0±1.5 | <0.001 | *** |
Mean annual precipitation [mm] | 1025±205 | 1287±210 | 920±167 | 1346±27.0 | <0.001 | *** |
Parameters calculated for relevés from EDGG Field Workshop | ||||||
Vegetation structure | ||||||
Cover vegetation total [%] | 74±27 | 68±16 | 91±7 | 85±14 | 0.033 | * |
Cover shrub layer [%] | 4±4 | 0±0 | 1±2 | 0±1 | <0.001 | *** |
Cover herb layer [%] | 69±22 | 62±15 | 79±26 | 81±16 | 0.068 | . |
Cover cryptogam layer [%] | 15±17 | 9±7 | 34±32 | 14±12 | 0.021 | * |
Cover litter [%] | 11±13 | 3±4 | 26±27 | 5±7 | 0.001 | ** |
Herb layer maximum height [cm] | 77±32 | 38±15 | 86±33 | 58±27 | 0.005 | ** |
Species richness | ||||||
Species richness (total) | 48.2±10.1 | 50.6±1.5 | 55.3±17.9 | 59.8±15.4 | 0.041 | * |
Species richness (vascular plants) | 42.9±8.9 | 46.0±3.8 | 50.6±16.8 | 50.5±12.2 | 0.151 | n.s |
Species richness (cryptogams) | 5.3±2.7 | 4.6±2.6 | 4.6±3.0 | 9.4±5.7 | 0.004 | ** |
Species richness (bryophytes) | 4.7±2.4 | 3.4±2.1 | 4.5±2.9 | 8.0±4.8 | 0.007 | ** |
Species richness (lichens) | 0.6±0.8 | 1.2±0.8 | 0.1±0.4 | 1.4±1.5 | 0.029 | * |
Topography | ||||||
Southing (cosine of aspect) | -0.1±0.7 | 0.8±0.2 | -0.5±0.6 | -0.5±0.5 | <0.001 | *** |
Inclination [°] | 19±11 | 14±5 | 11±7 | 16±15 | 0.563 | n.s |
Maximum microrelief [cm] | 7±4 | 9±7 | 7±7 | 11±9 | 0.240 | n.s |
Soil parameters | ||||||
Soil depth mean [cm] | 14±6 | 8±5 | 21±9 | 17±9 | 0.034 | * |
Soil depth CV | 48±26 | 100±82 | 35±29 | 46±35 | 0.020 | * |
Cover rocks and stones [%] | 6±11 | 9±13 | 1±4 | 8±16 | 0.600 | n.s |
Cover gravel [%] | 10±21 | 15±17 | 0±0 | 3±10 | 0.112 | n.s |
Cover fine soil [%] | 84±26 | 76±18 | 99±4 | 89±22 | 0.261 | n.s |
Coarse fragments [%] | 17±15 | 30±25 | 30±19 | 22±16 | 0.278 | n.s |
Fine fragments < 2mm [%] | 83±15 | 70±25 | 70±19 | 78±16 | 0.278 | n.s |
pH | 7.65±0.37 | 7.31±0.86 | 7.32±0.36 | 7.52±0.38 | 0.229 | n.s |
Electrical conductivity [µS/cm] | 188±60 | 213±141 | 225±103 | 261±80 | 0.031 | * |
CaCO3 [%] | 42.9±9 | 27.3±23.2 | 17.9±11.6 | 19.5±19 | <0.001 | *** |
Organic matter [%] | 0.8±0.3 | 1.8±1.5 | 1.3±0.4 | 1.6±0.8 | 0.001 | ** |
Ecological characteristics of the associations within the Festuco-Brometea. The p-values and significance levels refer to ANOVAs.
Parameter | Association | p-values | Sig. | |||||||
1.1.1 | 1.2.1 | 1.2.2 | 2.1.1 | 2.1.2 | 2.2.1 | 2.2.2 | 2.2.3 | |||
Total number of relevés | 52 | 25 | 61 | 14 | 26 | 12 | 78 | 69 | ||
Elevation [m a.s.l.] | 602 | 1113 | 989 | 578 | 552 | 800 | 797 | 1060 | <0.001 | *** |
Mediterraneity index | 0.83 | 0.61 | 0.59 | 0.89 | 0.91 | 0.57 | 0.64 | 0.51 | <0.001 | *** |
Annual mean temperature [ºC] | 11.8 | 9.8 | 9.9 | 12.1 | 12.1 | 10.5 | 10.5 | 9.4 | <0.001 | *** |
Mean annual precipitation [mm] | 1025 | 1242 | 1302 | 905 | 928 | 1369 | 1258 | 1445 | <0.001 | *** |
The NMDS analysis in Figure
The order 2 was defined by Briza media subsp. media, Cynosurus cristatus, Lotus corniculatus, Trifolium ochroleucon and T. pratense subsp. pratense, as diagnostic species (Table
Inside the alliance 2.1 two groups were distinguished. Each one was related to one association previously described according to the analysis of their types: group 2.1.1 to the association Prunello-Plantaginetum serpentinae and group 2.1.2 to the association Carduncello-Brachypodietum phoenicoidis.
Finally, alliance 2.2 was split into three groups corresponding to the associations Helictotricho-Seslerietum hispanicae, Calamintho-Seselietum montani and Carici-Teucrietum pyrenaici according to the position of their type relevés. The latter is mainly distributed in the calcareous Cantabrian and Pyrenean mountains (Figure
The alliance 1.2 is distributed in the highest elevations but also shows by far the highest values of southing; alliance 2.2 is also found in high elevations, and both share lower mediterraneity values compared to alliances 1.1 and 2.1; the two latter alliances show similar values of high temperature and low precipitation but 2.1 occurs in the most thermic and less rainy areas (Table
Comparison of four ecological variables among the four alliances. For elevation and Mediterraneity Index, all relevés were analysed, whereas for the rest of variables only relevés from EDGG Field Workshop were used. Letters represent homogeneous groups (at α = 0.05) according to Tukey’s post-hoc test following a significant ANOVA.
The total species richness is similar among the different alliances, as well as richness of vascular plants and lichens (Figure
Comparison of species richness divided into four groups (total species, vascular plants, bryophytes and lichens) among the four alliances in Festuco-Brometea using the relevés from EDGG Field Workshop. Letters represent homogeneous groups (at α = 0.05) according to Tukey’s post-hoc test following a significant ANOVA.
Association 1.1.1 – Thymelaeo ruizii-Aphyllanthetum monspeliensis
(relevès in Suppl. material
Characterisation: Grasslands usually growing on the middle part of slopes, characterised by the dominance of Brachypodium retusum and Bromopsis erecta subsp. erecta. Typical Mediterranean grasses such as Brachypodium retusum, B. phoenicoides, Dactylis glomerata subsp. hispanica, Festuca marginata subsp. andres-molinae and Helictochloa bromoides, and chamaephytes as Helianthemum apenninum subsp. apenninum, Lavandula latifolia and Thymus vulgaris subsp. vulgaris are frequent in this association. In addition, typical species of submediterranean and temperate grasslands are also common: Carex humilis, Koeleria vallesiana, Plantago lanceolata, Potentilla tabernaemontani, Sanguisorba minor, and Teucrium pyrenaicum.
Photo plate showing typical stands of the associations included in order 1 of the Festuco-Brometea. A Thymelaeo ruizii-Aphyllanthetum monspeliensis, A1 Overview, A2 Orchis papilionacea, endangered in Navarre; B Jurineo humilis-Festucetum hystricis, B1 Festuca hystrix, B2 Anthyllis montana; C Helianthemo incani-Koelerietum vallesianae, C1 Festuca rectifolia, C2 overview. Photos: A. Berastegi (A2, B1, B2, C1); J. Dengler (A1, C2).
Ecology and distribution: These grasslands are typical of temperate submediterranean transitional areas, at elevations between 400 to 1,100 m a.s.l. The sampled stands are grazed or recently abandoned. They are distributed in the middle part of Navarre region, as serial stages of Quercus faginea, Q. pubescens and Q. rotundifolia forests, and main land use are the cereal crops. They are usually found in carbonate soils developed on marls, limestones, flysch, conglomerates and sandstones, in the meso-supramediterranean and mesotemperate-supratemperate sub-humid to humid belts (
Syntaxonomy: This unit matches quite well with the association Thymelaeo ruizii-Aphyllanthetum monspeliensis, described from the submediterranean central areas in Navarre by Braun-Blanquet (1966) as a dwarf-shrub community. However,
Association 1.2.1 – Jurineo humilis-Festucetum hystricis
(relevès in Suppl. material
Characterisation: These grasslands grow on ridges and flat summit areas that are very windy, in mountain ranges usually above 900 m a.s.l. located in the transition between temperate and Mediterranean climates, in areas where cryoturbation phenomenon usually occurs. Carex humilis, Helianthemum canum subsp. canum and Koeleria vallesiana show a very high constancy in these open grasslands, but they are characterised by species like Anthyllis montana, Arenaria grandiflora subsp. grandiflora, Festuca hystrix, Jurinea humilis and Klasea nudicaulis, most of them typical of the high Mediterranean mountains.
Ecology and distribution: These communities can be found at elevations between 650 and 1,350 m a.s.l., although more commonly above 900 m, in the supramediterranean and supratemperate subhumid-humid belts (
Syntaxonomy: This unit fits quite well with the association Jurineo humilis-Festucetum hystricis.
Association 1.2.2 – Helianthemo incani-Koelerietum vallesianae
(relevès in Suppl. material
Characterisation: These communities are dominated by dry grassland species such as Carex humilis, Coronilla minima, Festuca rectifolia, Helianthemum canum subsp. canum, Helictochloa pratensis subsp. iberica, Koeleria vallesiana, Potentilla tabernaemontani, or Thymus praecox. Typical species of meso-xeric grasslands such as Bromopsis erecta subsp. erecta or Carthamus mitissimus are also common. From a physiognomic point of view, they are characterised by being short grasslands, with a cover of around 70-90%, in which some creeping chamaephytes can be important.
Ecology and distribution: The association represents pastures which are subject to intense livestock use, mainly by sheep, especially in the summer period. It occurs on different types of carbonate substrates (limestones, calcarenites, conglomerates, flysch), although mainly on limestone. They develop in the mountain ranges of the transition between the Atlantic and Mediterranean regions, also reaching the westernmost Pyrenean mountains, mostly in the montane belt.
Syntaxonomy: This unit matches well with the association Helianthemo incani-Koelerietum vallesianae, which was originally included in the class Festuco-Ononidetea, order Ononidetalia striatae, alliance Genistion occidentalis (
Association 2.1.1 – Prunello hyssopifoliae-Plantaginetum serpentinae
(relevès in Suppl. material
Characterisation: These communities are characterised by species like Festuca capillifolia, Jasonia tuberosa, Plantago maritima subsp. serpentina or Prunella hyssopifolia. Other species with high frequency are Blackstonia perfoliata, Carex flacca subsp. flacca, Centaurea jacea or Dorycnium pentaphyllum subsp. pentaphyllum.
Ecology and distribution: They are typical of the submediterranean climate and can be found at elevations from 410 to 1,000 m a.s.l., in the colline and montane belts. These communities develop in micro-depressions in loamy or clayey soils, which, due to their impermeable nature, are subject to temporary waterlogging. During the rainy season, these areas can become flooded, while in periods of strong sunshine they dry out completely. They are relatively frequent in the areas of blue-grey loams in the central part of Navarre, as serial stages of Quercus pubescens and Q. faginea forests, and main land use are the cereal crops.
Syntaxonomy: This unit matches quite well with the Prunello hyssopifoliae-Plantaginetum serpentinae association, originally placed in the class Molinio-Arrhenatheretea, although as a quite deviant community from the alliance Deschampsion mediae that often occurs in mosaic with meso-xeric grasslands; thus, typical dry grassland species are common (
Association 2.1.2 – Carduncello mitissimi-Brachypodietum phoenicoidis
(relevès in Suppl. material
Characterisation: Grasslands growing usually on the middle or bottom part of slopes, characterised by Blackstonia perfoliata, Brachypodium phoenicoides (including its hybrid with B. rupestre), Bromopsis erecta subsp. erecta, Carex flacca subsp. flacca, Eryngium campestre or Phleum pratense. Some other typical Festuco-Brometea species also occur: Carthamus mitissimus, Centaurea jacea, Ranunculus bulbosus subsp. bulbosus or Trifolium ochroleucon. Species of the class Molinio-Arrhenatheretea are also common, including Lotus corniculatus, Plantago lanceolata, Trifolium campestre and T. pratense.
Photo plate showing typical stands of the associations included in order 2 of the Festuco-Brometea. A Carduncello mitissimi-Brachypodietum phoenicoidis; B Helictotricho cantabrici-Seslerietum hispanicae; C Calamintho acini-Seselietum montani, D Carici ornithopodae-Teucrietum pyrenaici. Photos: J. Dengler (A, C, D); A. Berastegi (B).
Ecology and distribution: These dry grasslands are typical for the submediterranean climate type and can be found at elevations between 400 and 1,040 m a.s.l., in the supramediterranean and mesotemperate belts. They appear on clayey soils developed from calcareous materials (marl and limestone). They are distributed in the middle area of Navarre region, as serial stages of Quercus pubescens and Q. faginea forests, and main land use are the cereal crops. The sampled stands are grazed with low intensity or have been recently abandoned.
Syntaxonomy: This unit matches well with the association Carduncello mitissimi-Brachypodietum phoenicoidis, originally included in the order Brachypodietalia phoenicoidis (
Association 2.2.1 – Helictotricho cantabrici-Seslerietum hispanicae
(relevès in Suppl. material
Characterisation: These communities, dominated by the grasses Brachypodium rupestre, Helictotrichon cantabricum or Sesleria autumnalis, develop on rocky, steep slopes on limestone, usually with large crevices. In addition to the abovementioned species, it is common to find species such as Bromopsis erecta subsp. erecta, Carex flacca subsp. flacca, Dactylis glomerata, Galium pumilum, Teucrium pyrenaicum or Vincetoxicum hirundinaria subsp. intermedium. Some scrub species such as Dorycnium pentaphyllum subsp. pentaphyllum, Erica vagans or Genista hispanica subsp. occidentalis are also present, sometimes with relevant cover.
Ecology and distribution: These rocky grasslands are typical for the temperate climate and can be found at elevations between 460 and 1,050 m a.s.l., in the colline and montane belts These communities develop mainly in the context of the series of Quercus ilex, Fagus sylvatica and Quercus pubescens. However, their main role is as a permanent natural community on steep calcareous slopes.
Syntaxonomy: This unit roughly matches with the association Helictotricho cantabrici-Seslerietum hispanicae described by
Association 2.2.2 – Calamintho acini-Seselietum montani
(relevès in Suppl. material
Characterisation: Basophilous grasslands characterised by Brachypodium rupestre, Briza media subsp. media, Bromopsis erecta subsp. erecta, Carex flacca subsp. flacca, Lotus corniculatus or Plantago media. Some other taxa typical in these communities are Carthamus mitissimus, Helictochloa pratensis subsp. iberica, Linum catharticum subsp. catharticum, Potentilla tabernaemontani, Ranunculus bulbosus subsp. bulbosus, Thymus praecox and Trifolium ochroleucon. Species such as Achillea millefolium or Trifolium pratense are also common within the most mesic stands.
Ecology and distribution: These meso-xeric grasslands are typical for the temperate climate with submediterranean features and can be found at elevations between 230 and 1,400 m a.s.l., in the colline and montane belts. They develop on more or less deep soils, as serial stages of Fagus sylvatica and Quercus pubescens forests.
Syntaxonomy: This unit matches quite well with the association Calamintho acini-Seselietum montani described by
Association 2.2.3 – Carici ornithopodae-Teucrietum pyrenaici
(relevès in Suppl. material
Characterisation: These grasslands are characterised by species such as Clinopodium alpinum subsp. pyrenaeum, Festuca rectifolia, Helictochloa pratensis subsp. iberica, Seseli montanum subsp. montanum or Teucrium pyrenaicum, and some orophilous plants such as Poa alpina and Vicia pyrenaica. Typical elements of Festuco-Brometea such as Bromopsis erecta subsp. erecta, Carex caryophyllea, Helianthemum nummularium, etc. also occur, as well as others of Nardetea and Molinio-Arrhenatheretea such as Festuca microphylla, Lotus corniculatus or Plantago lanceolata.
Ecology and distribution: They are typical for the temperate climate and can be found at elevations between 560 and 1,720 m a.s.l., mostly in the montane belt. They usually grow on shallow soils (rendzina) developed on limestones, in the beech forest belt, and main land use is summer grazing (transterminant herds).
Syntaxonomy: This unit roughly matches with the association Carici ornithopodae-Teucrietum pyrenaici described by
Although our results largely concur with the previous classification of grasslands in Navarre (
The class Festuco-Ononidetea was proposed by
The class Elyno-Seslerietea gathers alpine and subalpine calcicolous swards of the nemoral mountain ranges of Europe. In Navarre, they belong to the Alpine-Pyrenean order Seslerietalia caeruleae and the alliance Primulion intricatae (
As regards the class Carici-Kobresietea bellardii, although our analysis included these communities in Elyno-Seslerietea, we kept it as a separate class, as it was only represented by two relevés in our dataset. Actually, these cryophytic alpine grasslands are very scarce in Navarre, so our geographic scope is not suitable to decide on the separation or grouping of both classes.
The class Nardetea strictae was defined as secondary oligotrophic grasslands and groups mesophilous or acidophilous, fairly grazed, tussock grasslands dominated by Nardus stricta from the montane to alpine belts with humid and hyper-humid ombroclimate (
According to our results, the association Merendero-Cynosuretum should also be included in the class Nardetea strictae. This association was originally included in the alliance Cynsurion cristati of the Molinio-Arrhenatheretea class (
Our analysis included relevés previously classified in the alliance Sedion pyrenaici from the class Sedo-Scleranthetea in Nardetea strictae. However, we have to consider the reduced context of our study, so we kept this class as a separate unit. In Navarre, these communities shaped by succulent species and dwarf chamaephytes growing on siliceous lithosols and rock surfaces (
Molinio-Arrhenetheretea is the most diverse class in Navarre regarding the number of associations.
The class Lygeo-Stipetea gathers Mediterranean pseudo-steppes on calcareous substrates and relict Mediterranean steppes on deep clayey soils (
Our analyses placed relevés of the classes Poetea bulbosae and Stipo-Trachynietea distachyae in Lygeo-Stipetea. However, our dataset contained only a small number of relevés from these classes and thus we cannot make any decision about the grouping of these classes within Lygeo-Stipetea. Therefore, we kept both classes as independent units.
In Navarre, the class Festuco-Brometea is composed of dry grasslands dominated by hemicryptophytes that develop on non-hygromorphic soils in temperate and submediterranean climates (
The numerical analysis clearly separates two groups that can be interpreted as two orders. Order 1 groups the more xerophytic relevès with Mediterranean influence which occupy an intermediate position between the orders Brachypodietalia pinnati and the more Mediterranean communities of Festuco-Ononidetea and Ononido-Rosmarinetea. This order would be a vicariant of Astragalo-Potentilletalia and Stipo-Festucetalia pallentis from central-southern Europe (
Communities in this order 1 are included in two alliances. Alliance 1 includes the association Thymelaeo-Aphyllanthetum monspeliensis, originally included in the alliance Helianthemo italici-Aphyllanthion monspeliensis (class Ononido-Rosmarinetea) by Braun-Blanquet (1966). Subsequently most Spanish phytosociologists have also placed it there, including the Spanish checklist (
Alliance 2 in this order 1 includes two associations that were previously classified in two different orders of the class Festuco-Ononidetea: Jurineo humilis-Festucetum hystricis in the order Festuco-Poetalia ligulatae and Helianthemo incani-Koelerietum vallesianae in Ononidetalia striatae (
Order 2 is related to Brachypodietalia pinnati and includes grasslands that usually develop in areas with a temperate climate, in well-constituted soils with relatively good water retention capacity and normally high total vegetation cover. Calamintho-Seselietum represents one of the typical associations of this order. This order also includes grasslands growing in rocky steep slopes from areas of high rainfall (Helictotricho-Seslerietum hispanicae and Carici-Teucrietum pyrenaici), as well as dry grasslands from submediterranean areas, but the latter ones are restricted to soils or topographic situations that allow relatively good water retention (Prunello-Plantaginetum serpentinae and Carduncello-Brachypodietum phoenicoidis).
Rocky grasslands from this order 2 (Helictotricho-Seslerietum hispanicae and Carici-Teucrietum pyrenaici) are included in Ononidetalia striatae in the Spanish checklist, but our analysis has shown that they have a strong floristic relationship with grasslands of Brachypodietalia pinnati. In fact, both associations were originally included in this order (
Alliance 2.1, which includes the associations Prunello-Plantaginetum serpentinae and Carduncello-Brachypodietum phoenicoidis in Navarre, could be assigned to the alliance Brachypodion phoenicoidis. More comprehensive analyses would be needed to confirm, as this alliance is distributed along the Western Mediterranean region, and its type association Brachypodietum phoenicoidis Br.-Bl. 1924 was described in Mediterranean France (
Class: Festuco-Brometea Br.-Bl. et Tx. ex Klika et Hadač 1944
Order 1: ???
Alliance 1.1: ???
1.1.1: Thymelaeo ruizii-Aphyllanthetum monspeliensis Br.-Bl. et P. Montserrat in Br.-Bl. 1966
Alliance 1.2: ???
1.2.1: Jurineo humilis-Festucetum hystricis Peralta et al. in
1.2.2: Helianthemo incani-Koelerietum vallesianae Berastegi et al. in
Order 2: Brachypodietalia pinnati Korneck 1974 nom. cons. propos. (= Brometalia erecti Koch 1926)
Nomenclatural remark:
Alliance 2.1: ???
2.1.1: Prunello hyssopifoliae-Plantaginetum serpentinae F. Prieto et al. ex
2.1.2: Carduncello mitissimi-Brachypodietum phoenicoidis García-Mijangos et al. in
Alliance 2.2: Potentillo montanae-Brachypodion pinnati Br.-Bl. 1967
2.2.1: Helictotricho cantabrici-Seslerietum hispanicae Br.-Bl. 1967
2.2.2: Calamintho acini-Seselietum montani Br.-Bl. 1967
2.2.3: Carici ornithopodae-Teucrietum pyrenaici
Grasslands of Festuco-Brometea showed the highest total species richness, and specifically meso-xeric grasslands of the association Calamintho-Seselietum montani, which have previously been highlighted as species rich grasslands (
Up to now vegetation ecologists in the Southern European countries, and particularly in the Mediterranean region, rarely considered bryophytes and lichens as part of the vegetation - unlike many of their colleagues in temperate and boreal Europe. This is reflected by the fact that for example,
The combination of numerical methods allows a standardisation of the classification of grassland types. In fact, with our expert system we could largely reproduce the associations previously recognised in the region. Moreover, some often “diagnostic” species mentioned in the literature could be confirmed by our numerical analyses of a large dataset, while others were not supported by the data. However, at the class level, we found significant deviations from the Iberian syntaxonomic tradition (
Our study provides, for the first time, an electronic expert system for the grasslands of Navarre, which allows a standardised assignment of any new relevé, thus is of enormous value, particularly for practitioners. We provide, also for the first time, a detailed databased characterisation and comparison of the syntaxa in terms of their environmental conditions and biodiversity. We were also able to show that bryophytes and lichens, contrary to past assumptions, are core elements of these grasslands and in particular, the Mediterranean ones of Lygeo-Stipetea, both in terms of biodiversity and of diagnostic species. Therefore, they should also be taken into account in Mediterranean phytosociology.
Once the main five phytosociological classes were differentiated, our study focused on the analysis of the Festuco-Brometea. Therefore, an in-depth analysis based on expert systems of the rest of the classes would be desirable. Moreover, classes whose status could not be resolved due to a small/marginal dataset or due to plot sizes being too small, should be specifically addressed in future studies with better/more data from a larger area.
Finally, it can be emphasised that we have provided important insights from the western part of Europe that complement the extensive studies of
The vegetation-plot data underlying this study are stored and available in the GrassPlot database (https://edgg.org/databases/GrassPlot; dataset code ES_A;
I.G.M, I.B. and A.B. organized the 7th EDGG Field Workshop in Navarre (Spain); as EDGG Field Workshop Coordinator during the Field Workshop, J.D. ensured consistent application of the EDGG methodology, I.G.M. identified the vascular plant species collected during the Field Workshop, J.E. identified the lichens, R.N. identified the bryophytes and added ecological aspects, and O.Y. analysed the soil samples and described methodological aspects; A.B. compiled 839 relevés used in the paper from 1996 to 1999; I.G.M. together with J.D. developed the numerical classification, implemented the expert system and identified the diagnostic species with the collaboration of I.B., A.B., A.K., M.J., and D.V.; M.J. developed the NMDS ordination, I.D. and D.V. analysed differences between syntaxa by means of ANOVAs and J.D. calculated and analysed biodiversity patterns; I.G.M. led the writing of the manuscript with substantial inputs from A.B., I.B. and J.D.; all authors critically revised the manuscript.
We are grateful to EDGG and IAVS for financial support for some of the participants. I.G. and I.B. were supported by the Basque Government (IT936‐16). We would like to give special thanks to F. Urra and I. Ibarrola for helping with the organization of the field workshop in Navarre. Thanks to all Field Workshop participants and other helpers who did not opt-in as authors.
Itziar Garcia-Mijangos (itziar.garcia@ehu.eus), ORCID: https://orcid.org/0000-0002-6642-7782
Asun Berastegi (aberastg@gan-nik.es), ORCID: https://orcid.org/0000-0003-0456-3305
Idoia Biurrun (Corresponding author, idoia.biurrun@ehu.eus), ORCID: https://orcid.org/0000-0002-1454-0433
Iwona Dembicz (i.dembicz@gmail.com), ORCID: https://orcid.org/0000-0002-6162-1519
Monika JaniŠová (monika.janisova@gmail.com), ORCID: https://orcid.org/0000-0002-6445-0823
Anna Kuzemko (anyameadow.ak@gmail.com), ORCID: https://orcid.org/0000-0002-9425-2756
Denys Vynokurov (denys.vynokurov@gmail.com), ORCID: https://orcid.org/0000-0001-7003-6680
Didem Ambarlı (didem.ambarli@gmail.com), ORCID: https://orcid.org/0000-0001-5589-9373
Javier Etayo (jetayosa@educacion.navarra.es), ORCID: https://orcid.org/0000-0003-0392-0710
Goffredo Filibeck (filibeck@unitus.it), ORCID: https://orcid.org/0000-0002-4187-9467
Ute Jandt (ute.jandt@botanik.uni-halle.de), ORCID: https://orcid.org/0000-0002-3177-3669
Rayna Natcheva (renimoss@bio.bas.bg)
Oktay Yildiz (oktayyildiz@duzce.edu.tr), ORCID: https://orcid.org/0000-0002-8058-4506
Jürgen Dengler (dr.juergen.dengler@gmail.com), ORCID: https://orcid.org/0000-0003-3221-660X
Geographic, environmental and structural data of the relevés
Ordered relevé table and complete constancy table of the five classes
Confusion matrix comparing TWINSPAN and original classification
Expert system for the Festuco-Brometea order 1 alliances
Expert system for the Festuco-Brometea order 2 alliances
Expert system for the Festuco-Brometea alliance 1.2 associations
Expert system for the Festuco-Brometea alliance 2.1 associations
Expert system for the Festuco-Brometea alliance 2.2 associations
Ordered relevé table and complete constancy table of the Festuco-Brometea