Research Article |
Corresponding author: Fotios Xystrakis ( fotios.xystrakis@fri.gr ) Academic editor: Federico Fernández-González
© 2022 Fotios Xystrakis, Minas Chasapis, Eleni Eleftheriadou, Dimitrios Samaras, Konstantinos Theodoropoulos.
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:
Xystrakis F, Chasapis M, Eleftheriadou E, Samaras D, Theodoropoulos K (2022) The optimization of typical species inventory of habitat types of a NATURA 2000 site using a phytosociological approach. Plant Sociology 59(2): 1-16. https://doi.org/10.3897/pls2022592/01
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The definition of typical species inventories of the 92/43/EEC Directive habitat types is a valuable information for the optimization of the conservation status assessment. Habitat-specific assessment protocols and predefined local inventories of typical species provide a method for a relatively fast and accurate assessment of the criterion “structures and functions”. Habitat types are often defined and described on the basis of a phytosociological description of vegetation units, mainly at the syntaxonomical level of alliance. Therefore, the definition of typical species inventories can be based on phytosociological approaches. Within this concept we surveyed the vegetation of a NATURA 2000 Special Area of Conservation in northern Greece in order to optimize and downscale the existing region-wide inventories of typical species. In total, we sampled 164 relevés in beech and in thermophilous deciduous broadleaved forests. The relevés were assigned to vegetation units and habitat types using numerical approaches and their differential and constant taxa were defined. We used these taxa to draw up the optimized, site-specific inventories of typical species for seven habitat types of community interest and one habitat type of national interest.
Constant taxa, differential taxa, GR1240002, Greece Habitats Directive, Mt. Tzena
Among the main research questions in vegetation science is the description and typification of plant communities, in a way that recognizable and reproducible units are identified wherever similar ecological conditions are met (
Phytosociological analysis and classification of plant assemblages has also strong relations with nature conservation (
Similarly in Greece, a first attempt for the development of a crosswalk between habitat types and phytosociological syntaxa is presented by
The assessment of typical species is included as a sub-criterion for the assessment of ‘structures and functions’ (
In Greece, phytosociological methods and concepts were applied to elaborate habitat-specific protocols for the assessment of the habitat types of the 92/43/EEC Directive (
In this paper we describe and classify the forest vegetation of a Natura 2000 site, applying phytosociological concepts and methods that ensure objectivity. The identified vegetation units are then assigned to habitat types, and site-specific inventories of typical species are formed. These adapted floristic inventories are to replace the existing, large-scale typical species lists of evaluation protocols, and used as the basis for the assessment of the criterion “structures and functions”. This substantially supports the optimization of the implementation of the 92/43/EEC Directive.
The study area consists of Mt. Tzena, which is a mountain situated in central Macedonia, Greece, at the borders with the Republic of North Macedonia (Fig.
The vegetation of the mountain has been recently described and mapped at an area of 5,326.75 ha by
Regarding the protection regime of the study area, Mt. Tzena is part of the Special Area of Conservation (SAC) of the 92/43/EEC Directive of the European Natura 2000 network under the code GR1240002 – Ori Tzena. It largely coincides with the Special Protection Area (SPA) of the bird’s Directive GR1240007 – Ori Tzena kai Pinovo. SAC GR1240002 consists of two distinct mountains, Mt. Tzena and Mt. Pinovo, the former covering the eastern half of the SAC.
Thirteen habitat types of the 92/43/EEC Directive have been described in the SAC, with one of them, 6230*- Species-rich Nardus grasslands, on siliceous substrates in mountain areas (and submountain areas in Continental Europe), being a priority habitat type. Among these thirteen habitat types, seven are forest habitat types and five of them, 9110 (Luzulo-Fagetum beech forests), 9130 (Asperulo-Fagetum beech forests), 9150 (Medio-European limestone beech forests of the Cephalanthero-Fagion), 91M0 (Pannonian-Balkanic turkey oak –sessile oak forests) and 9280 (Quercus frainetto woods) are under consideration in this paper. The other two, 9270-Hellenic beech forests with Abies borisii-regis and 92C0-Platanus orientalis and Liquidambar orientalis woods (Platanion orientalis), cover a relatively small surface area and were not considered in the analysis. In addition to these aforementioned habitat types, a forest habitat type of national interest that includes the extended Carpinus orientalis and Ostrya carpinifolia forests has been considered. It is coded under the 4-digit code 9254 and described as “Forests of Ostrya, Carpinus orientalis and mixed thermophilous forests” (
To study the vegetation of Mt. Tzena, 105 relevés were recorded in beech-dominated stands and 59 relevés were recorded in thermophilous deciduous forests, i.e. pure or mixed stands of Quercus pubescens, Q. petraea subsp. polycarpa, Q. frainetto, Carpinus orientalis and Ostrya carpinifolia. The location of the relevés is shown in Figure
To distinguish vegetation units, we applied agglomerative hierarchical clustering. Firstly, relevés were separated into two distinct vegetation data sets on the basis of species dominance: beech and thermophilous broad-leaved forests. These two main datasets reflect the two major groups of forest habitat types that occur in the study area: the temperate beech-dominated forests and the thermophilous Quercus spp. and Carpinus/Ostrya dominated forests. In each data set, we removed taxa that occur in less than 3% of the total number of plots in order to reduce the effect of rare species (
Having selected the final number of clusters, we identified the diagnostic/differential taxa of vegetation units applying: (a) the multilevel analysis for the identification of indicator taxa presented in
Each relevé was assigned to an already described syntaxon at the association level. We used the works of
In order to refine the typical species inventory, vegetation relevés have to be first assigned to a habitat type, or eventually to one of the vicariants of habitat types that were identified by
To refine the typical species inventory and form site-specific evaluation protocols, we (a) narrowed down the typical taxa in the assessment protocol proposed by
Vegetation survey resulted in 509 plant taxa in 164 relevés (see Suppl. material
The outputs of the cluster analysis allowed for the identification of four vegetation groups in the beech dominated forests of the study area (Table
Indicator (differential) and constant species (>25% constancy) of beech-dominated forests (applying
taxon | group 1 | group 2 | group 3 | group 4 | group 1 | group 2 | group 3 | group 4 |
---|---|---|---|---|---|---|---|---|
indicator | constancy | |||||||
Constant taxa | ||||||||
Fagus sylvatica subsp. sylvatica | - | - | - | - | 100 | 100 | 100 | 100 |
Euphorbia amygdaloides | - | - | - | - | 79 | 65 | 38 | 57 |
Lactuca muralis | - | - | - | - | 57 | 97 | 59 | 65 |
Neottia nidus-avis | - | - | - | - | 71 | 35 | 54 | 70 |
Poa nemoralis | - | - | - | - | 100 | 94 | 97 | 83 |
Luzula sylvatica | - | - | - | - | 29 | 19 | 54 | 39 |
Cystopteris fragilis | - | - | - | - | 7 | 29 | 14 | 17 |
Saxifraga rotundifolia | - | - | - | - | 21 | 35 | 8 | 26 |
Veronica urticifolia | - | - | - | - | 7 | 29 | 5 | 26 |
Prenanthes purpurea | - | - | - | - | 14 | 29 | 30 | 13 |
Abies borisii-regis | - | - | - | - | 14 | 13 | 14 | 30 |
Differential taxa | ||||||||
Campanula trachelium | p | n | n | n | 71 | 16 | 14 | 0 |
Doronicum orientale | p | n | n | n | 64 | 0 | 11 | 0 |
Galium mollugo aggr. | p | n | n | n | 36 | 3 | 5 | 4 |
Galium pseudaristatum | p | n | n | n | 57 | 0 | 16 | 0 |
Juniperus oxycedrus subsp. deltoides | p | n | n | n | 36 | 3 | 3 | 4 |
Lathyrus laxiflorus | p | n | n | n | 64 | 13 | 11 | 0 |
Lathyrus niger | p | n | n | n | 21 | 0 | 0 | 0 |
Lathyrus venetus | p | n | n | n | 50 | 0 | 3 | 0 |
Platanthera chlorantha | p | n | n | n | 36 | 0 | 0 | 0 |
Primula veris | p | n | n | n | 57 | 10 | 8 | 13 |
Quercus frainetto | p | n | n | n | 36 | 3 | 0 | 0 |
Quercus pubescens | p | n | n | n | 21 | 0 | 0 | 0 |
Rosa arvensis | p | n | n | n | 64 | 13 | 3 | 0 |
Silene atropurpurea | p | n | n | n | 21 | 0 | 0 | 0 |
Silene italica subsp. italica | p | n | n | n | 50 | 3 | 5 | 4 |
Silene viridiflora | p | n | n | n | 29 | 0 | 3 | 0 |
Sorbus torminalis | p | n | n | n | 21 | 0 | 0 | 0 |
Trifolium pignantii | p | n | n | n | 57 | 3 | 16 | 0 |
Festuca koritnicensis | p | n | n | - | 21 | 0 | 0 | 4 |
Hippocrepis emerus subsp. emeroides | p | n | n | - | 36 | 0 | 3 | 13 |
Campanula sparsa | p | n | - | n | 21 | 0 | 3 | 0 |
Carex flacca subsp. serrulata | p | - | n | n | 21 | 3 | 0 | 0 |
Cyclamen hederifolium | p | - | n | n | 29 | 10 | 0 | 4 |
Dactylis glomerata subsp. glomerata | p | - | n | n | 64 | 29 | 0 | 13 |
Potentilla micrantha | p | - | n | n | 79 | 45 | 22 | 13 |
Fraxinus ornus | p | - | n | - | 43 | 16 | 8 | 22 |
Silene vulgaris | p | - | n | - | 29 | 10 | 3 | 22 |
Viola alba subsp. alba | p | - | n | - | 50 | 23 | 11 | 17 |
Primula acaulis | p | - | n | n | 29 | 19 | 3 | 0 |
Dioscorea communis | p | - | - | n | 21 | 6 | 3 | 0 |
Poa bulbosa subsp. bulbosa | p | - | - | n | 21 | 3 | 3 | 0 |
Sedum cepaea | p | - | - | n | 21 | 3 | 5 | 0 |
Campanula persicifolia | p | n | p | n | 43 | 3 | 27 | 0 |
Campanula spatulata | p | n | p | n | 50 | 3 | 41 | 4 |
Festuca heterophylla | p | - | p | n | 79 | 45 | 89 | 26 |
Hieracium racemosum | p | n | p | n | 71 | 16 | 54 | 13 |
Luzula forsteri | p | n | p | n | 93 | 13 | 54 | 0 |
Polypodium vulgare | p | - | p | n | 29 | 19 | 41 | 0 |
Veronica vindobonensis | p | - | p | n | 93 | 45 | 78 | 17 |
Hieracium murorum | p | n | p | p | 93 | 16 | 62 | 52 |
Aremonia agrimonoides subsp. agrimonoides | p | p | n | p | 43 | 48 | 8 | 48 |
Cephalanthera rubra | p | n | n | p | 71 | 13 | 8 | 52 |
Acer hyrcanum | p | - | n | p | 36 | 13 | 8 | 43 |
Sanicula europaea | p | p | n | n | 36 | 35 | 0 | 4 |
Hedera helix subsp. helix | p | p | n | n | 64 | 35 | 5 | 9 |
Brachypodium sylvaticum subsp. sylvaticum | p | p | n | n | 21 | 23 | 0 | 0 |
Aegopodium podagraria | n | p | n | n | 14 | 48 | 8 | 4 |
Anemone ranunculoides | n | p | - | n | 0 | 42 | 14 | 0 |
Calamintha grandiflora | n | p | - | - | 0 | 55 | 32 | 13 |
Dactylorhiza saccifera | - | p | - | n | 7 | 29 | 16 | 0 |
Dryopteris filix-mas | n | p | n | n | 0 | 52 | 3 | 13 |
Epilobium montanum | n | p | n | n | 0 | 65 | 5 | 17 |
Galeobdolon montanum | n | p | n | n | 14 | 77 | 16 | 13 |
Geranium robertianum | n | p | n | n | 0 | 61 | 8 | 13 |
Geum urbanum | n | p | n | - | 0 | 23 | 0 | 4 |
Hordelymus europaeus | n | p | n | n | 0 | 45 | 3 | 0 |
Moehringia trinervia | - | p | n | n | 29 | 39 | 8 | 9 |
Polystichum aculeatum | n | p | n | - | 0 | 26 | 0 | 4 |
Pteridium aquilinum | - | p | - | n | 36 | 45 | 35 | 9 |
Pulmonaria rubra | n | p | n | n | 7 | 42 | 0 | 0 |
Rubus hirtus | n | p | n | n | 7 | 77 | 14 | 0 |
Salvia glutinosa | n | p | - | - | 0 | 23 | 3 | 4 |
Scilla subnivalis | - | p | n | n | 21 | 42 | 11 | 4 |
Symphytum tuberosum subsp. angustifolium | n | p | - | n | 7 | 48 | 27 | 9 |
Urtica dioica | n | p | n | n | 0 | 23 | 0 | 0 |
Viola reichenbachiana/riviniana | - | p | - | n | 36 | 61 | 32 | 13 |
Cardamine bulbifera | n | p | p | n | 7 | 71 | 38 | 0 |
Galium odoratum | n | p | p | n | 0 | 65 | 30 | 0 |
Melica uniflora | - | p | p | n | 14 | 42 | 38 | 9 |
Lilium martagon | - | p | n | p | 14 | 32 | 5 | 39 |
Avenella flexuosa | n | - | p | - | 0 | 13 | 22 | 9 |
Hypopitys monotropa | - | - | p | n | 14 | 3 | 22 | 0 |
Luzula luzuloides subsp. luzuloides | n | - | p | n | 14 | 29 | 57 | 9 |
Arabis turrita | - | n | n | p | 21 | 13 | 3 | 48 |
Carex digitata | n | - | - | p | 7 | 13 | 11 | 39 |
Cotoneaster nebrodensis | n | n | n | p | 0 | 0 | 0 | 26 |
Doronicum columnae | n | - | n | p | 0 | 19 | 3 | 35 |
Orthilia secunda | n | n | - | p | 0 | 0 | 11 | 30 |
Polygonatum odoratum | - | - | n | p | 14 | 6 | 3 | 26 |
Rosa villosa | n | n | n | p | 0 | 0 | 0 | 30 |
Rubus idaeus | n | - | n | p | 0 | 19 | 0 | 26 |
Sesleria robusta | - | n | - | p | 29 | 0 | 11 | 52 |
Solidago virgaurea | - | n | - | p | 21 | 6 | 16 | 39 |
Sorbus aria | - | n | n | p | 7 | 3 | 0 | 26 |
Sorbus X thuringiaca | n | n | n | p | 0 | 0 | 0 | 22 |
The assignment of the relevés to the syntaxa (units) that are described by
The site characteristics of the relevés forming each group are shown in Figure
Number of relevés of beech dominated forests assigned to each syntaxonomic unit described by
Number of relevés | ||||
Unit ( |
group 1 | group 2 | group 3 | group 4 |
u2: mesophytic beech forests of NE Greece | . | . | 2 | 1 |
u3: acidophytic forests (on gneiss) of NE Greece | . | . | 1 | . |
u7: mesophytic beech forests of Mt. Voras on humid nutrient-rich soils (Lamiastro montani-Fagetum sylvaticae) | . | 18 | 5 | 2 |
u8: forests of Mt. Voras occurring in warmer and drier sites (Galium odoratum-Fagus sylvatica) | . | . | 4 | 2 |
u10: forests on calcareous substrate at high elevations (Cardamine graeca-Fagus sylvatica) | . | 2 | 4 | 6 |
u11: Forests on calcareous substrate at lower altitudes (subcom. with Fraxinus ornus of the Lathyro alpestris-Fagetum sylvaticae) | 5 | 9 | 6 | 11 |
u12: Lathyro alpestris-Fagetum sylvaticae | 2 | 1 | 13 | 1 |
u13: thermophytic beech forests of Mt. Chortiatis (southwestern part of NE Greece) (Rubus canescens-Fagus sylvatica) | . | . | 1 | . |
u14: thermophytic beech forests of Mt. Cholomon (southwestern part of NE Greece) | 7 | 1 | 1 | . |
sum | 14 | 31 | 37 | 23 |
Unit (Bergmeir and Dimopoulos 2001) | group 1 | group 2 | group 3 | group 4 |
u2: Galium odoratum-Fagus sylvatica comm. | . | . | 1 | . |
u3: Lamiastro montani-Fagetum sylvaticae | . | 20 | 4 | 3 |
u5: Orthilio secundae-Fagetum (subcom. with Luzula luzuloides) | 2 | 3 | 23 | 16 |
u6: Orthilio secundae-Fagetum (subcom. with Abies borisii-regis) | . | . | . | 1 |
u11: Lathyro alpestris-Fagetum sylvaticae (subcom. with Galium odoratum) | 2 | 5 | 2 | 3 |
u13: Rubus canescens-Fagus sylvatica comm. | 10 | 3 | 7 | . |
sum | 14 | 31 | 37 | 23 |
Among the indicator species of this group, taxa of the alliance Quercion frainetto such as Primula veris, Quercus frainetto, Trifolium pignantii are in abundance. Furthermore, it can be observed that relevés of this group occur in the lowest elevational zone of the examined Fagus forests (Figure
Group 2 Group 2 is related to unit 7 of
Relevés of this group should be included in the habitat type 9130 (Asperulo fagetum beech forests) (
Group 3 is a large, rather heterogeneous group. It consists of 37 relevés, the majority of which (36) occurs on siliceous bedrock. It reflects an intermediate character between cool and warm habitats. Relevés of this group are associated with various units described by
Likewise in group 1 and group 2, a considerable number of relevés are similar to habitat type 9280, which includes beech forests with a significant presence of floristic elements of the alliance Quercion frainetto (
Group 4 consists of 23 relevés, the majority of them being located on calcareous substrate. It is strongly affiliated with units 10-forests on calcareous substrate at high elevations (Cardamine graeca-Fagus sylvatica) and 11-forests on calcareous substrate at lower elevations (subcom. with Fraxinus ornus of the Lathyro alpestris-Fagetum sylvaticae)) of
The majority of the relevés of this group are associated with the habitat types 9150 (Medio-European limestone beech forests of the Cephalanthero-Fagion) that include a large part of beech forests developing on calcareous substrates in Greece (
Comparing the updated typical species inventories with the respective inventories presented in the original protocols, it is made obvious that substantial changes took place. Out of 29 typical species in the existing evaluation protocol of the habitat type 9110, only 11 are also included in the new inventory, which counts 26 taxa. Regarding habitat type 9130, out of 36 species, 19 are also present in the updated protocol (in these taxa Urtica dioica is also included). Thirty-five taxa are considered as new addition to the updated inventory. Regarding habitat type 9150, 16 out of 44 taxa of the existing evaluation protocol are considered in the updated inventory while 20 more taxa have been included additionally. Finally, in habitat type 9280, 31 out of 53 taxa are included in the updated inventory while 28 new taxa are added. These changes are mostly related to the extended sampling that took place in this NATURA 2000 area. The existing typical species inventories were compiled using a limited number of relevés from each NATURA 2000 site.
The outputs of the cluster analysis justified the acceptance of three groups in the data set that includes the relevés dominated by thermophilous deciduous broadleaved species (Table
These three groups correspond to the i) Ostrya carpinifolia-Carpinus orientalis-Quercus pubescens, ii) Quercus pubescens-Carpinus orientalis, and iii) Quercus frainetto and Q. petraea forest formations respectively. The assignment of the relevés to the syntaxa (units) identified in Greece by
In terms of site characteristics (Figure
Indicator (differential) and constant species of thermophilous deciduous broadleaved forests (applying
group 1 | group 2 | group 3 | group 1 | group 2 | group 3 | |
---|---|---|---|---|---|---|
indicator | constancy | |||||
Constant taxa | ||||||
Asplenium ceterach | - | - | - | 41 | 16 | 17 |
Hypericum perforatum | - | - | - | 36 | 16 | 17 |
Pilosella piloselloides/bauhini | - | - | - | 27 | 11 | 11 |
Thymus longicaulis/sibthorpii | - | - | - | 45 | 16 | 17 |
Acer hyrcanum | - | - | - | 73 | 95 | 67 |
Arabis turrita | - | - | - | 77 | 89 | 56 |
Brachypodium sylvaticum subsp. sylvaticum | - | - | - | 82 | 42 | 44 |
Carex flacca subsp. serrulata | - | - | - | 59 | 89 | 61 |
Carpinus orientalis | - | - | - | 86 | 89 | 44 |
Cephalanthera longifolia | - | - | - | 50 | 32 | 72 |
Cephalaria flava | - | - | - | 27 | 53 | 28 |
Clinopodium vulgare subsp. orientale | - | - | - | 86 | 74 | 72 |
Cyclamen hederifolium | - | - | - | 50 | 53 | 33 |
Dactylis glomerata subsp. glomerata | - | - | - | 82 | 89 | 100 |
Dioscorea communis | - | - | - | 41 | 42 | 56 |
Euphorbia amygdaloides | - | - | - | 59 | 89 | 72 |
Fagus sylvatica subsp. sylvatica | - | - | - | 45 | 32 | 78 |
Festuca valesiaca | - | - | - | 41 | 37 | 61 |
Fraxinus ornus | - | - | - | 100 | 95 | 83 |
Juniperus oxycedrus subsp. deltoides | - | - | - | 86 | 95 | 89 |
Physospermum cornubiense | - | - | - | 27 | 26 | 39 |
Potentilla micrantha | - | - | - | 55 | 68 | 89 |
Primula veris | - | - | - | 32 | 63 | 33 |
Prunus divaricata | - | - | - | 41 | 26 | 44 |
Rosa arvensis | - | - | - | 59 | 63 | 44 |
Scorzoneroides cichoriacea | - | - | - | 41 | 63 | 44 |
Silene italica subsp. italica | - | - | - | 77 | 74 | 94 |
Veronica vindobonensis | - | - | - | 50 | 47 | 89 |
Viola alba subsp. alba | - | - | - | 91 | 84 | 50 |
Arabis sagittata | - | - | - | 27 | 32 | 17 |
Cornus mas | - | - | - | 45 | 53 | 17 |
Festuca koritnicensis | - | - | - | 41 | 47 | 17 |
Cardamine graeca | - | - | - | 36 | 21 | 33 |
Galium aparine | - | - | - | 55 | 21 | 44 |
Rosa canina/corymbifera | - | - | - | 27 | 11 | 33 |
Muscari neglectum | - | - | - | 32 | 26 | 17 |
Anthoxanthum aristatum | - | - | - | 5 | 11 | 28 |
Quercus petraea subsp. polycarpa | - | - | - | 18 | 11 | 28 |
Campanula trachelium | - | - | - | 14 | 37 | 17 |
Lithospermum purpurocaeruleum | - | - | - | 18 | 47 | 17 |
Sorbus torminalis | - | - | - | 23 | 47 | 33 |
Neottia nidus-avis | - | - | - | 14 | 5 | 28 |
Pteridium aquilinum | - | - | - | 23 | 16 | 44 |
Rubus sanctus | - | - | - | 23 | 11 | 28 |
Verbascum xanthophoeniceum | - | - | - | 14 | 5 | 28 |
Differential taxa | ||||||
Ostrya carpinifolia | p | - | n | 91 | 42 | 28 |
Asplenium trichomanes | p | n | - | 50 | 5 | 28 |
Clematis vitalba | p | n | n | 73 | 16 | 17 |
Ruscus aculeatus | p | - | n | 41 | 16 | 6 |
Poa bulbosa subsp. bulbosa | p | n | - | 36 | 5 | 22 |
Hieracium murorum | p | n | - | 32 | 5 | 22 |
Asplenium onopteris/adiantum-nigrum | p | n | n | 55 | 0 | 17 |
Medicago sativa subsp. falcata | p | - | n | 27 | 11 | 0 |
Viola reichenbachiana/riviniana | p | n | n | 32 | 0 | 0 |
Achillea holosericea | p | - | n | 23 | 11 | 0 |
Solidago virgaurea | p | n | - | 27 | 0 | 6 |
Hedera helix subsp. helix | p | n | p | 41 | 0 | 33 |
Aremonia agrimonoides subsp. agrimonoides | p | n | p | 68 | 11 | 44 |
Quercus pubescens | p | p | n | 95 | 100 | 33 |
Teucrium chamaedrys subsp. chamaedrys | p | p | n | 82 | 95 | 28 |
Sesleria robusta | p | p | n | 41 | 58 | 6 |
Asperula purpurea | p | p | n | 50 | 32 | 6 |
Carex halleriana | p | p | n | 27 | 58 | 0 |
Euonymus verrucosus | p | p | n | 36 | 37 | 0 |
Hippocrepis emerus subsp. emeroides | p | p | n | 59 | 79 | 11 |
Trifolium alpestre | n | p | - | 27 | 89 | 72 |
Galium mollugo aggr. | N | p | - | 27 | 79 | 39 |
Centaurea napulifera | - | p | n | 36 | 74 | 22 |
Orlaya daucoides | - | p | n | 18 | 47 | 11 |
Origanum vulgare subsp. vulgare | - | p | n | 18 | 42 | 11 |
Geranium sanguineum | n | p | n | 5 | 74 | 11 |
Festuca hirtovaginata s.l. | - | p | n | 18 | 37 | 6 |
Polygonatum odoratum | - | p | n | 14 | 42 | 6 |
Veronica jacquinii | n | p | n | 9 | 47 | 0 |
Campanula persicifolia | n | p | n | 9 | 42 | 6 |
Inula conyzae | - | p | n | 18 | 32 | 6 |
Campanula bononiensis | n | p | n | 5 | 47 | 0 |
Tanacetum corymbosum | n | p | - | 0 | 37 | 11 |
Calamintha nepeta | - | p | n | 14 | 26 | 0 |
Astragalus monspessulanus | - | p | n | 18 | 21 | 0 |
Sorbus aria | - | p | n | 18 | 21 | 0 |
Sorbus aucuparia | n | p | - | 0 | 26 | 17 |
Thalictrum minus subsp. saxatile | n | p | n | 5 | 37 | 0 |
Erysimum cuspidatum | - | p | n | 14 | 21 | 0 |
Lamium garganicum | - | p | n | 5 | 21 | 0 |
Achnatherum bromoides | - | p | n | 5 | 21 | 0 |
Allium macedonicum | - | p | n | 5 | 21 | 0 |
Linum flavum subsp. albanicum | n | p | n | 0 | 21 | 0 |
Brachypodium pinnatum | n | p | p | 18 | 84 | 56 |
Vicia tenuifolia subsp. dalmatica | n | p | p | 0 | 21 | 44 |
Chamaecytisus hirsutus | n | - | p | 9 | 32 | 44 |
Festuca heterophylla | n | n | p | 14 | 11 | 56 |
Geum urbanum | n | - | p | 9 | 21 | 39 |
Galium pseudaristatum | - | n | p | 18 | 0 | 50 |
Trifolium ochroleucon | n | n | p | 5 | 11 | 44 |
Cephalanthera rubra | - | n | p | 14 | 5 | 39 |
Silene viridiflora | n | n | p | 0 | 11 | 44 |
Silene coronaria | n | n | p | 0 | 5 | 50 |
Sedum cepaea | n | n | p | 5 | 0 | 50 |
Lathyrus pratensis | n | n | p | 0 | 5 | 44 |
Epipactis helleborine | n | - | p | 0 | 11 | 39 |
Digitalis grandiflora | n | - | p | 0 | 11 | 39 |
Anthemis tinctoria | n | - | p | 0 | 16 | 28 |
Genista carinalis | - | n | p | 14 | 0 | 28 |
Scutellaria columnae | n | n | p | 5 | 0 | 39 |
Lathyrus niger | n | n | p | 0 | 0 | 39 |
Verbascum nigrum subsp. abietinum | n | - | p | 0 | 16 | 22 |
Hieracium lachenalii | n | - | p | 0 | 5 | 28 |
Pilosella leucopsilon | - | n | p | 5 | 0 | 22 |
Hieracium umbrosum | n | n | p | 0 | 0 | 28 |
Silene atropurpurea | - | n | p | 5 | 0 | 22 |
Vicia grandiflora | n | n | p | 0 | 0 | 22 |
Poa nemoralis | - | n | p | 59 | 37 | 100 |
Campanula spatulata | - | n | p | 59 | 26 | 83 |
Luzula forsteri | - | n | p | 41 | 26 | 94 |
Trifolium pignantii | n | n | p | 27 | 16 | 78 |
Quercus frainetto | n | n | p | 9 | 5 | 83 |
Rubus canescens | - | n | p | 27 | 11 | 56 |
Lathyrus laxiflorus | n | n | p | 18 | 11 | 61 |
Hieracium racemosum | - | n | p | 18 | 5 | 39 |
Number of relevés ssigned to each syntaxonomic unit of thermophilous deciduous forests (
Number of relevés | |||
Association | group 1 | group 2 | group 3 |
DrO: Dryopterido pallidae-Ostryetum carpinifoliae | 21 | 19 | 2 |
DiQ: Digitali viridiflorae-Quercetum frainetto | - | - | 8* |
GQp: Genisto carinalis-Quercetum petraeae | - | - | 3 |
VeQ: Verbasco glabrati-Quercetum frainetto | - | - | 5 |
TiC: Tilio tomentosae-Castanetum | 1 | - | - |
sum | 22 | 19 | 18 |
Group 1 consists of Ostrya carpinifolia-Carpinus orientalis mixed stands, with the former taxon being rather dominant in most relevés. Quercus pubescens is a constant species in this group, but it participates in the tree cover with considerably low values. Relevés of this group are assigned to the Dryopterido pallidae–Ostryetum carpinifoliae Bergmeier 1990 association (
Relevés of group 1 should be included in the habitat type 925A (forests of Ostrya, Carpinus orientalis and mixed thermophilous forests) (
Group 2 has a similar character with group 1, but Quercus pubescens is rather dominant when compared with Carpinus orientalis and Ostrya carpinifolia, with the latter having low cover values (<3) in all relevés but three. Relevés of this group are assigned to the Dryopterido pallidae-Ostryetum association Bergmeier 1990 (
Relevés of group 2 should be included in the habitat type 91M0 (Pannonian-Balkanic turkey oak- sessile oak forests) due to the relative dominance of Quercus pubescens. Habitat type 91M0 has replaced the habitat type 924A-eastern Mediterranean and Balkan thermophilous oak-dominated forests (
The updated inventory of typical species that includes differential and constant taxa of group 2 is shown in Suppl. material
Group 3 is clearly differentiated from the previous group. Quercus petraea subsp. polycarpa and Q. frainetto dominate the canopy and it is differentiated by a large number of taxa. When Q. petraea subsp. polycarpa dominates, relevés of this group are assigned to the Genisto carinalis-Quercetum petraeae. When Q. frainetto dominates, relevés of this group are assigned to the Verbasco glabrati-Quercetum frainetto Gamisans et Hebrard 1979, Huetio cynapioidis-Quercetum frainetto Raus ex Raus Bergmeir 2008, or the Digitali viridiflorae-Quercetum frainetto Gamisans et Hebrard 1980 associations (Table
Relevés of group 3 should be included in the habitat type 91M0 (Pannonian-Balkanic turkey oak- sessile oak forests), but a distinct vicariant of the Quercus frainetto-Quercus petraea subsp. polycarpa dominated formations should be formed. The updated inventory of typical species is shown in Suppl. material
Since there is not an existing evaluation protocols for the habitat type 925A or the two vicariants of the 91M0 habitat type for the area, it is not possible to perform any comparisons with existing typical species inventories.
In this paper, we present a thorough methodology that aims to directly bridge some evaluation criteria of the conservation status of habitat types with the phytosociological background of description of habitat types in the 92/43/EEC Directive. Beginning with preferential data sampling, phytosociological analysis of relevés, and concrete, reproducible rules, we provided typical species inventories for all the forest habitat types that occur on Mt. Tzena in Greece. The preferential sampling ensured that degraded localities were avoided, thus typical species did not include differential or indicator taxa of disturbed sites. The latter is among the main properties of typical species. The numerical analyses are reproducible, repeatable in other sites and require data that are typically collected during standard phytosociological studies. The choice of constant and diagnostic/differential species ensures that the defined typical species are directly related with the habitat types, which is yet another important property of typical species. The method we proposed does not address the final property of typical species, which requires that typical species are sensitive to changes of habitat conditions. This can be addressed by considering specific structures that are related to the presence of specific taxa or groups of taxa. The criterion ‘structures and functions’ can include parameters that are directly related with the occurrence and cover of targeted species that indicate changes of the condition of the habitat. For example,
Moreover, this method can be performed at various spatial scales. Large, national data sets can provide typical species inventories of wider regions that share similar phytogeographical conditions, provided that there is a balanced sampling. Typical species inventories that are compiled for wider phytogeographical regions allow for the consideration of ‘dark diversity’ species (
The authors would like to acknowledge the assistance of George Skias in data entry. Additionally, we would like to acknowledge Federico Fernández-González and Olivier Argagnon for their valuable comments that significantly improved the quality and clarity of the manuscript as well as Odysseas Papoulis Xystrakis for proof reading the manuscript.
F.X.: Conceptualization, data analysis, manuscript writing; M.C.: Conceptualization, data collection, data analysis, manuscript proof reading; E.E., D.S and K.T.: Conceptualization, manuscript proof reading.
Figure S1
Data type: maps
Explanation note: Maps including important ecological information for the study area: Substrate, Koeppen climate classification, elevation and habitat types.
Table S1
Data type: vegetation data
Explanation note: Vegetation data.
Table S2
Data type: vegetation table
Explanation note: Vegetation table of beech-dominated stands.
Tables S3 and S4
Data type: tables
Explanation note: Updated inventory both of typical species of beech dominated habitat types in GR1240002 and of typical species of thermophilous forest habitat types in GR1240002.
Table S5
Data type: Vegetation table
Explanation note: Vegetation table of thermophilous forests.