Galium aparine

Categories of substrate humidity do not reflect fine differences of various habitats, on the contrary, they are defined very broadly. These categories can be derived also from the ordinal scale for moisture (1-12) defined by Ellenberg et al. (1991). Plants adapted to growing in dry soils, or soils that frequently dry out, are assigned to a category Dry, which corresponds to the first four degrees of the Ellenberg scale (1-4). Accordingly, plants of mesic habitats are assigned to the category Mesic (5 and 6 of the Ellenberg scale), plants of wet to soaked, poorly aerated soils are in the category Wet (7-9), and finally, aquatic plants are assigned to the category Water (10-12). If available, we used Ellenberg-type indicator values aggregated to broader categories, in other cases, categories were assigned according to the knowledge of local experts.

Data were compiled from several sources available across Europe, namely the indicator-value datasets for Great Britain (Hill et al. 2000), Cantabrian Range in Spain (Jiménez-Alfaro et al. 2021), France (Julve 2015), Germany (Ellenberg et al. 2001, taken from Ellenberg & Leuschner 2010), Switzerland (Landolt et al. 2010), Austria (Karrer 1992), Czech Republic (Chytrý et al. 2018), Hungary (Borhidi 1995), Ukraine (Didukh 2011), Italy (Guarino & La Rosa 2019, with additional corrections by R. Guarino), Southern Aegean region of Greece (Böhling et al. 2002) and European mires (Hájek et al. 2020).

Categories

• Dry
• Mesic
• Wet
• Water

Data source and citation

Axmanová, I. (2022). Substrate humidity relationship. – www.FloraVeg.EU.

Further references

Böhling, N., Greuter, W. & Raus, T. (2002). Zeigerwerte der Gefäßpflanzen der Südägäis (Griechenland) [Indicator values of the vascular plants in the Southern Aegean (Greece)]. Braun-Blanquetia, 32, 1–108.
Borhidi, A. (1995). Social behaviour types, the naturalness and relative ecological indicator values of the higher plants in the Hungarian flora. Acta Botanica Hungarica, 39, 97–181.
Chytrý, M., Tichý, L., Dřevojan, P., Sádlo, J. & Zelený, D. (2018). Ellenberg-type indicator values for the Czech flora. Preslia, 90, 83–103.
Didukh Ya.P. (2011). The ecological scales for the species of Ukrainian flora and their use in synphytoindication. Kyiv: Phytosociocentre.
Ellenberg, H., Weber, H. E., Düll, R., Wirth, V., Werner, W. & Paulißen, D. (1991). Zeigerwerte von Pflanzen in Mitteleuropa. Scripta Geobotanica, 18, 1–248.
Ellenberg, H. & Leuschner, C. (2010). Zeigerwerte der Pflanzen Mitteleuropas. In: Ellenberg H. & Leuschner, C., Vegetation Mitteleuropas mit den Alpen. Stuttgart: Verlag Eugen Ulmer.
French Flora database (baseflor), project of Flore et végétation de la France et du Monde: CATMINAT. Available at http://philippe.julve.pagesperso-orange.fr/catminat.htm [accessed June 2020]
Guarino, R., La Rosa, M. & Pignatti, S. (Eds) (2019). Flora d'Italia, volume 4. Bologna: Edagricole.
Hill, M.O., Roy, D.B., Mountford, J.O., & Bunce, R.G.H. (2000). Extending Ellenberg’s indicator values to a new area: an algorithmic approach. Journal of Applied Ecology, 37, 3–15.
Jiménez-Alfaro, B., Carlón, L., Fernández-Pascual, E., Acedo, C., Alfaro-Saiz, E., Alonso Redondo, R., … Vázquez, V. M. (2021). Checklist of the vascular plants of the Cantabrian mountains. Mediterranean Botany, 42, e74570.
Karrer, G. (1992). Österreichische Waldboden-Zustandsinventur. Teil VII: Vegetationsökologische Analysen. Mitteilungen der forstlichen Bundesversuchsanstalt Wien, 168, 193-242.
Landolt, E., Bäumler, B., Erhardt, A., Hegg, O., Klötzli, F., Lämmler, W., … Wohlgemuth, T. (2010). Flora indicativa – Ökologische Zeigerwerte und biologische Kennzeichen zur Flora der Schweiz und der Alpen. Bern: Haupt Verlag.

Categories of substrate reaction do not reflect fine differences of various habitats, on the contrary, they are defined very broadly. These categories can be derived also from the ordinal scale for reaction (1-9) defined by Ellenberg et al. (1991). Plants adapted to growing in very acidic soils are assigned to a category Acidic, which corresponds to the first four degrees of the Ellenberg scale (1-4). Accordingly, plants of intermediate habitats are assigned to the category Slightly acidic (5 and 6 of the Ellenberg scale), plants of calcium-rich substrates are in the category Alkaline (7-9). If available, we used Ellenberg-type indicator values aggregated to broader categories, in other cases, categories were assigned according to the knowledge of local experts.

Data were compiled from several sources available across Europe, namely the indicator-value datasets for Great Britain (Hill et al. 2000), Cantabrian Range in Spain (Jiménez-Alfaro et al. 2021), France (Julve 2015), Germany (Ellenberg et al. 2001, taken from Ellenberg & Leuschner 2010), Switzerland (Landolt et al. 2010), Austria (Karrer 1992), Czech Republic (Chytrý et al. 2018), Hungary (Borhidi 1995), Ukraine (Didukh 2011), Italy (Guarino & La Rosa 2019, with additional corrections by R. Guarino), Southern Aegean region of Greece (Böhling et al. 2002).

Categories

• Acidic
• Slightly acidic to near-neutral
• Alkaline
• Indifferent

Data source and citation

Axmanová, I. (2022). Substrate reaction relationship. – www.FloraVeg.EU.

Further references

Böhling, N., Greuter, W. & Raus, T. (2002). Zeigerwerte der Gefäßpflanzen der Südägäis (Griechenland) [Indicator values of the vascular plants in the Southern Aegean (Greece)]. Braun-Blanquetia, 32, 1–108.
Borhidi, A. (1995). Social behaviour types, the naturalness and relative ecological indicator values of the higher plants in the Hungarian flora. Acta Botanica Hungarica, 39, 97–181.
Chytrý, M., Tichý, L., Dřevojan, P., Sádlo, J. & Zelený, D. (2018). Ellenberg-type indicator values for the Czech flora. Preslia, 90, 83–103.
Didukh Ya.P. (2011). The ecological scales for the species of Ukrainian flora and their use in synphytoindication. Kyiv: Phytosociocentre.
Ellenberg, H., Weber, H. E., Düll, R., Wirth, V., Werner, W. & Paulißen, D. (1991). Zeigerwerte von Pflanzen in Mitteleuropa. Scripta Geobotanica, 18, 1–248.
Ellenberg, H. & Leuschner, C. (2010). Zeigerwerte der Pflanzen Mitteleuropas. In: Ellenberg H. & Leuschner, C., Vegetation Mitteleuropas mit den Alpen. Stuttgart: Verlag Eugen Ulmer.
French Flora database (baseflor), project of Flore et végétation de la France et du Monde: CATMINAT. Available at http://philippe.julve.pagesperso-orange.fr/catminat.htm [accessed June 2020]
Guarino, R., La Rosa, M. & Pignatti, S. (Eds) (2019). Flora d'Italia, volume 4. Bologna: Edagricole.
Hájek, M., Dítě, D., Horsáková, V., Mikulášková, E., Peterka, T., Navrátilová, J., … Horsák, M. (2020). Towards the pan-European bioindication system: Assessing and testing updated hydrological indicator values for vascular plants and bryophytes in mires. Ecological Indicators, 116, 106527.
Hill, M.O., Roy, D.B., Mountford, J.O., & Bunce, R.G.H. (2000). Extending Ellenberg’s indicator values to a new area: an algorithmic approach. Journal of Applied Ecology, 37, 3–15.
Jiménez-Alfaro, B., Carlón, L., Fernández-Pascual, E., Acedo, C., Alfaro-Saiz, E., Alonso Redondo, R., … Vázquez, V. M. (2021). Checklist of the vascular plants of the Cantabrian mountains. Mediterranean Botany, 42, e74570.
Karrer, G. (1992). Österreichische Waldboden-Zustandsinventur. Teil VII: Vegetationsökologische Analysen. Mitteilungen der forstlichen Bundesversuchsanstalt Wien, 168, 193-242.
Landolt, E., Bäumler, B., Erhardt, A., Hegg, O., Klötzli, F., Lämmler, W., … Wohlgemuth, T. (2010). Flora indicativa – Ökologische Zeigerwerte und biologische Kennzeichen zur Flora der Schweiz und der Alpen. Bern: Haupt Verlag.

Categories of salinity do not reflect fine differences of various habitats, on the contrary, they are defined very broadly. These categories can be derived also from the ordinal scale for reaction (0-9) defined by Ellenberg et al. (1991). Plants adapted to growing in soils without salt content are assigned to a category Non-saline, which corresponds to the first degree of the Ellenberg scale (0) and it is the prevailing category. Plants of slightly saline habitats are assigned to the category Slightly saline or brackish (1-4 of the Ellenberg scale), while plants of salt-rich substrates are in the category Saline (5-9). If available, we used Ellenberg-type indicator values aggregated to broader categories, in other cases, categories were assigned according to the knowledge of local experts.

Data were compiled from several sources available across Europe, namely the indicator-value datasets for Great Britain (Hill et al. 2000), France (Julve 2015), Germany (Ellenberg et al. 2001, taken from Ellenberg & Leuschner 2010), Switzerland (Landolt et al. 2010), Austria (Karrer 1992), Czech Republic (Chytrý et al. 2018), Hungary (Borhidi 1995), Ukraine (Didukh 2011), Italy (Guarino & La Rosa 2019, with additional corrections by R. Guarino), Southern Aegean region of Greece (Böhling et al. 2002).

Categories

• Non-saline
• Slightly saline or brackish
• Saline

Data source and citation

Axmanová, I. (2022). Salinity relationship. – www.FloraVeg.EU.

Further references

Böhling, N., Greuter, W. & Raus, T. (2002). Zeigerwerte der Gefäßpflanzen der Südägäis (Griechenland) [Indicator values of the vascular plants in the Southern Aegean (Greece)]. Braun-Blanquetia, 32, 1–108.
Borhidi, A. (1995). Social behaviour types, the naturalness and relative ecological indicator values of the higher plants in the Hungarian flora. Acta Botanica Hungarica, 39, 97–181.
Chytrý, M., Tichý, L., Dřevojan, P., Sádlo, J. & Zelený, D. (2018). Ellenberg-type indicator values for the Czech flora. Preslia, 90, 83–103.
Didukh Ya.P. (2011). The ecological scales for the species of Ukrainian flora and their use in synphytoindication. Kyiv: Phytosociocentre.
Ellenberg, H., Weber, H. E., Düll, R., Wirth, V., Werner, W. & Paulißen, D. (1991). Zeigerwerte von Pflanzen in Mitteleuropa. Scripta Geobotanica, 18, 1–248.
Ellenberg, H. & Leuschner, C. (2010). Zeigerwerte der Pflanzen Mitteleuropas. In: Ellenberg H. & Leuschner, C., Vegetation Mitteleuropas mit den Alpen. Stuttgart: Verlag Eugen Ulmer.
French Flora database (baseflor), project of Flore et végétation de la France et du Monde: CATMINAT. Available at http://philippe.julve.pagesperso-orange.fr/catminat.htm [accessed June 2020]
Guarino, R., La Rosa, M. & Pignatti, S. (Eds) (2019). Flora d'Italia, volume 4. Bologna: Edagricole.
Hájek, M., Dítě, D., Horsáková, V., Mikulášková, E., Peterka, T., Navrátilová, J., … Horsák, M. (2020). Towards the pan-European bioindication system: Assessing and testing updated hydrological indicator values for vascular plants and bryophytes in mires. Ecological Indicators, 116, 106527.
Hill, M.O., Roy, D.B., Mountford, J.O., & Bunce, R.G.H. (2000). Extending Ellenberg’s indicator values to a new area: an algorithmic approach. Journal of Applied Ecology, 37, 3–15.
Karrer, G. (1992). Österreichische Waldboden-Zustandsinventur. Teil VII: Vegetationsökologische Analysen. Mitteilungen der forstlichen Bundesversuchsanstalt Wien, 168, 193-242.
Landolt, E., Bäumler, B., Erhardt, A., Hegg, O., Klötzli, F., Lämmler, W., … Wohlgemuth, T. (2010). Flora indicativa – Ökologische Zeigerwerte und biologische Kennzeichen zur Flora der Schweiz und der Alpen. Bern: Haupt Verlag.

Ellenberg-type indicator values for European plant species are expert-based rankings of plant species according to their ecological optima on main environmental gradients, using ordinal scales defined by Ellenberg et al. (1991). The indicator values for light are expressed on a scale from 1 to 9. Note that indicator values for trees refer to juvenile individuals in the herb and shrub layers.

Values for individual species were calculated as the means across available national or regional datasets of plant indicator values or were newly assigned based on species co-occurrences in European vegetation plots, see Tichý et al. (2023). Although they have one decimal place, the newly introduced indicator values are compatible with the original Ellenberg values. They can be used for large-scale studies of European flora and vegetation or gap-filling in regional datasets.

Datasets used for compilation include several sources, namely Ellenberg-type indicator values for Great Britain (Hill et al., 2000); France (Julve, 2015); Germany (Ellenberg et al., 2001, taken from Ellenberg & Leuschner, 2010); Czech Republic (Chytrý et al., 2018); Austria (Karrer, 1992); Hungary (Borhidi, 1995); Ukraine (Didukh, 2011) and Italy (Guarino & La Rosa, 2019, a corrected version prepared by R. Guarino, values 10–12 were rescaled to the value 9).

The European values and also the original and taxonomically harmonized regional datasets of Ellenberg-type indicator values are available in the Download section of this website.

Categories

• 1 – deep shade plant, occurring where the incident radiation is less than 1% of that in an open area, rarely at more than 30%
• 2 – transition between values 1 and 3
• 3 – shade plant, usually occurring where the incident radiation is less than 5% of that in an open area, but also at sunnier sites
• 4 – transition between values 3 and 5
• 5 – semi-shade plant, only exceptionally occurring in full light, but usually at more than 10% of the diffuse radiation incident in an open area
• 6 – transition between values 5 and 7; rarely at less than 20% of diffuse radiation incident in an open area
• 7 – half-light plant, mostly occurring at full light, but also in the shade up to about 30% of diffuse radiation incident in an open area
• 8 – light plant, only exceptionally occurring at less than 40% of diffuse radiation incident in an open area
• 9 – full light plant, occurring only in fully irradiated places, not at less than 50% of diffuse radiation incident in an open area

Data source and citation

Tichý L., Axmanová I., Dengler J., Guarino R., Jansen F., Midolo G., … Chytrý M. (2023). Ellenberg-type indicator values for European vascular plant species. Journal of Vegetation Science, 34, e13168. https://doi.org/10.1111/jvs.13168

Further references

Borhidi, A. (1995). Social behaviour types, the naturalness and relative ecological indicator values of the higher plants in the Hungarian flora. Acta Botanica Hungarica, 39, 97–181.
Chytrý M., Tichý L., Dřevojan P., Sádlo J. & Zelený D. (2018). Ellenberg-type indicator values for the Czech flora. Preslia 90: 83–103.
Didukh Ya.P. (2011). The ecological scales for the species of Ukrainian flora and their use in synphytoindication. Kyiv: Phytosociocentre.
Ellenberg H., Weber H. E., Düll R., Wirth V., Werner W. & Paulißen D. (1991). Zeigerwerte von Pflanzen in Mitteleuropa. Scripta Geobotanica, 18, 1–248.
Ellenberg, H. & Leuschner, C. (2010). Zeigerwerte der Pflanzen Mitteleuropas. In: Ellenberg H. & Leuschner, C., Vegetation Mitteleuropas mit den Alpen. Stuttgart: Verlag Eugen Ulmer.
Guarino, R., La Rosa, M. & Pignatti, S. (Eds) (2019). Flora d'Italia, volume 4. Bologna: Edagricole.
Hill, M.O., Roy, D.B., Mountford, J.O., & Bunce, R.G.H. (2000). Extending Ellenberg’s indicator values to a new area: an algorithmic approach. Journal of Applied Ecology, 37, 3–15.
Julve, P. (2015). Baseflor. Index botanique, écologique et chorologique de la flore de France (Baseflor. Botanical, ecological and chorological index of the flora of France). Available at http://philippe.julve.pagesperso-orange.fr/catminat.htm [accessed 2022].
Karrer, G. (1992). Österreichische Waldboden-Zustandsinventur. Teil VII: Vegetationsökologische Analysen (Austrian forest soil status inventory. Part VII: Vegetation ecology analyses). Mitteilungen Forstliche Bundesversuchsanstalt Wien, 168, 193–242.

Ellenberg-type indicator values for European plant species are expert-based rankings of plant species according to their ecological optima on main environmental gradients, using ordinal scales defined by Ellenberg et al. (1991). The indicator values for temperature are expressed on an ordinal scale from 1 to 12. While the original scale proposed by Ellenberg et al. (1991) was defined only up to 9, here we use the extended range defined for the Mediterranean flora (Pignatti et al. 2005, see also Guarino et al. 2019).

Values for individual species were calculated as the means across available national or regional datasets of plant indicator values or were newly assigned based on species co-occurrences in European vegetation plots, see Tichý et al. (2023). Although they have one decimal place, the newly introduced indicator values are compatible with the original Ellenberg values. They can be used for large-scale studies of European flora and vegetation or gap-filling in regional datasets.

Datasets used for compilation include several sources, namely Ellenberg-type indicator values for France (Julve, 2015); Switzerland and the Alps (Landolt et al., 2010); Germany (Ellenberg et al., 2001, taken from Ellenberg & Leuschner, 2010); Czech Republic (Chytrý et al., 2018); Austria (Karrer, 1992); Hungary (Borhidi, 1995) and Italy (Guarino & La Rosa, 2019, a corrected version prepared by R. Guarino).

The European values and also the original and taxonomically harmonized regional datasets of Ellenberg-type indicator values are available in the Download section of this website.

Categories

• 1 – cold indicator, occurring only in high mountain areas, i.e. the alpine and nival belts
• 2 – transition between values 1 and 3
• 3 – cool indicator, occurring mainly in subalpine areas
• 4 – transition between values 3 and 5
• 5 – moderate heat indicator, occurring from lowland to montane belt, mainly in submontane-temperate areas
• 6 – transition between values 5 and 7
• 7 – heat indicator, occurring in relatively warm lowlands
• 8 – transition between values 7 and 9
• 9 – heat indicator, occurring in the Mediterranean and warmest sites in southern Central Europe
• 10 – heat indicator, occurring in warm Mediterranean areas
• 11 – heat indicator, occurring at mesic sites in the southern Mediterranean
• 12 – heat indicator, occurring in warm and semi-desert areas in the southern Mediterranean

Data source and citation

Tichý L., Axmanová I., Dengler J., Guarino R., Jansen F., Midolo G., … Chytrý M. (2023). Ellenberg-type indicator values for European vascular plant species. Journal of Vegetation Science, 34, e13168. https://doi.org/10.1111/jvs.13168

Further references

Borhidi, A. (1995). Social behaviour types, the naturalness and relative ecological indicator values of the higher plants in the Hungarian flora. Acta Botanica Hungarica, 39, 97–181.
Chytrý M., Tichý L., Dřevojan P., Sádlo J. & Zelený D. (2018). Ellenberg-type indicator values for the Czech flora. Preslia 90: 83–103.
Ellenberg H., Weber H. E., Düll R., Wirth V., Werner W. & Paulißen D. (1991). Zeigerwerte von Pflanzen in Mitteleuropa. Scripta Geobotanica, 18, 1–248.
Ellenberg, H. & Leuschner, C. (2010). Zeigerwerte der Pflanzen Mitteleuropas. In: Ellenberg H. & Leuschner, C., Vegetation Mitteleuropas mit den Alpen. Stuttgart: Verlag Eugen Ulmer.
Guarino, R., La Rosa, M. & Pignatti, S. (Eds) (2019). Flora d'Italia, volume 4. Bologna: Edagricole.
Julve, P. (2015). Baseflor. Index botanique, écologique et chorologique de la flore de France (Baseflor. Botanical, ecological and chorological index of the flora of France). Available at http://philippe.julve.pagesperso-orange.fr/catminat.htm [accessed 2022].
Karrer, G. (1992). Österreichische Waldboden-Zustandsinventur. Teil VII: Vegetationsökologische Analysen (Austrian forest soil status inventory. Part VII: Vegetation ecology analyses). Mitteilungen Forstliche Bundesversuchsanstalt Wien, 168, 193–242.
Landolt, E., Bäumler, B., Erhardt, A., Hegg, O., Klötzli, F., Lämmler, W., … Wohlgemuth, T. (2010). Flora indicativa – Ökologische Zeigerwerte und biologische Kennzeichen zur Flora der Schweiz und der Alpen. Bern: Haupt Verlag.
Pignatti, S., Menegoni, P. & Pietrosanti, S. (2005). Valori di bioindicazione delle piante vascolari della Flora d'Italia [Bioindicator values of vascular plants of the Flora of Italy]. Braun-Blanquetia, 39, 1–97.

Ellenberg-type indicator values for European plant species are expert-based rankings of plant species according to their ecological optima on main environmental gradients, using ordinal scales defined by Ellenberg et al. (1991). The indicator value for soil or water reaction are expressed on an ordinal scale from 1 to 9 defined by Ellenberg et al. (1991). In acidic environments, the value can be considered as a proxy for pH, whereas in near-neutral or alkaline environments, it is more a proxy for calcium concentration.

Values for individual species were calculated as the means across available national or regional datasets of plant indicator values or were newly assigned based on species co-occurrences in European vegetation plots, see Tichý et al. (2023). Although they have one decimal place, the newly introduced indicator values are compatible with the original Ellenberg values. They can be used for large-scale studies of European flora and vegetation or gap-filling in regional datasets.

Datasets used for compilation include several sources, namely Ellenberg-type indicator values for Great Britain (Hill et al., 2000); France (Julve, 2015); Germany (Ellenberg et al., 2001, taken from Ellenberg & Leuschner, 2010); Czech Republic (Chytrý et al., 2018); Austria (Karrer, 1992); Hungary (Borhidi, 1995); Italy (Guarino & La Rosa, 2019, a corrected version prepared by R. Guarino).

The European values and also the original and taxonomically harmonized regional datasets of Ellenberg-type indicator values are available in the Download section of this website.

Categories

• 1 – indicator of strong acidity, never occurring in slightly acidic to alkaline conditions
• 2 – transition between values 1 and 3
• 3 – acidity indicator, occurring mainly in acidic conditions, exceptionally in neutral conditions
• 4 – transition between values 3 and 5
• 5 – indicator of moderate acidity, occurring rarely in strongly acidic as well as in neutral to alkaline conditions
• 6 – transition between values 5 and 7
• 7 – indicator of slightly acidic to slightly basic conditions, never occurring in very acidic conditions
• 8 – transition between values 7 and 9, occurring mostly in calcium-rich conditions
• 9 – base and lime indicator, always occurring in calcium-rich conditions

Data source and citation

Tichý L., Axmanová I., Dengler J., Guarino R., Jansen F., Midolo G., … Chytrý M. (2023). Ellenberg-type indicator values for European vascular plant species. Journal of Vegetation Science, 34, e13168. https://doi.org/10.1111/jvs.13168

Further references

Borhidi, A. (1995). Social behaviour types, the naturalness and relative ecological indicator values of the higher plants in the Hungarian flora. Acta Botanica Hungarica, 39, 97–181.
Chytrý M., Tichý L., Dřevojan P., Sádlo J. & Zelený D. (2018). Ellenberg-type indicator values for the Czech flora. Preslia 90: 83–103.
Ellenberg H., Weber H. E., Düll R., Wirth V., Werner W. & Paulißen D. (1991). Zeigerwerte von Pflanzen in Mitteleuropa. Scripta Geobotanica, 18, 1–248.
Ellenberg, H. & Leuschner, C. (2010). Zeigerwerte der Pflanzen Mitteleuropas. In: Ellenberg H. & Leuschner, C., Vegetation Mitteleuropas mit den Alpen. Stuttgart: Verlag Eugen Ulmer.
Guarino, R., La Rosa, M. & Pignatti, S. (Eds) (2019). Flora d'Italia, volume 4. Bologna: Edagricole.
Hill, M.O., Roy, D.B., Mountford, J.O., & Bunce, R.G.H. (2000). Extending Ellenberg’s indicator values to a new area: an algorithmic approach. Journal of Applied Ecology, 37, 3–15.
Julve, P. (2015). Baseflor. Index botanique, écologique et chorologique de la flore de France (Baseflor. Botanical, ecological and chorological index of the flora of France). Available at http://philippe.julve.pagesperso-orange.fr/catminat.htm [accessed 2022].
Karrer, G. (1992). Österreichische Waldboden-Zustandsinventur. Teil VII: Vegetationsökologische Analysen (Austrian forest soil status inventory. Part VII: Vegetation ecology analyses). Mitteilungen Forstliche Bundesversuchsanstalt Wien, 168, 193–242.

Ellenberg-type indicator values for European plant species are expert-based rankings of plant species according to their ecological optima on main environmental gradients, using ordinal scales defined by Ellenberg et al. (1991). The indicator values for nutrients are expressed on an ordinal scale from 1 to 9 defined by Ellenberg et al. (1991). The value is a proxy for the availability of nitrogen or phosphorus and, to some extent, also a proxy for site primary productivity.

Values for individual species were calculated as the means across available national or regional datasets of plant indicator values or were newly assigned based on species co-occurrences in European vegetation plots, see Tichý et al. (2023). Although they have one decimal place, the newly introduced indicator values are compatible with the original Ellenberg values. They can be used for large-scale studies of European flora and vegetation or gap-filling in regional datasets.

Datasets used for compilation include several sources, namely Ellenberg-type indicator values for Great Britain (Hill et al., 2000); France (Julve, 2015); Germany (Ellenberg et al., 2001, taken from Ellenberg & Leuschner, 2010); Czech Republic (Chytrý et al., 2018); Austria (Karrer, 1992); Hungary (Borhidi, 1995); Italy (Guarino & La Rosa, 2019, a corrected version prepared by R. Guarino, values 10–12 were rescaled to the value 9).

The European values and also the original and taxonomically harmonized regional datasets of Ellenberg-type indicator values are available in the Download section of this website.

Categories

• 1 – occurring at nutrient-poorest sites
• 2 – transition between values 1 and 3
• 3 – occurring at nutrient-poor sites more frequently than at average sites and exceptionally at rich sites
• 4 – transition between values 3 and 5
• 5 – occurring at moderately nutrient-rich sites, and less frequently at poor and rich sites
• 6 – transition between values 5 and 7
• 7 – occurring at nutrient-rich sites more often than at average sites and only exceptionally at poor sites
• 8 – pronounced nutrient indicator
• 9 – concentrated at very nutrient-rich sites

Data source and citation

Tichý L., Axmanová I., Dengler J., Guarino R., Jansen F., Midolo G., … Chytrý M. (2023). Ellenberg-type indicator values for European vascular plant species. Journal of Vegetation Science, 34, e13168. https://doi.org/10.1111/jvs.13168

Further references

Borhidi, A. (1995). Social behaviour types, the naturalness and relative ecological indicator values of the higher plants in the Hungarian flora. Acta Botanica Hungarica, 39, 97–181.
Chytrý M., Tichý L., Dřevojan P., Sádlo J. & Zelený D. (2018). Ellenberg-type indicator values for the Czech flora. Preslia 90: 83–103.
Ellenberg H., Weber H. E., Düll R., Wirth V., Werner W. & Paulißen D. (1991). Zeigerwerte von Pflanzen in Mitteleuropa. Scripta Geobotanica, 18, 1–248.
Ellenberg, H. & Leuschner, C. (2010). Zeigerwerte der Pflanzen Mitteleuropas. In: Ellenberg H. & Leuschner, C., Vegetation Mitteleuropas mit den Alpen. Stuttgart: Verlag Eugen Ulmer.
Guarino, R., La Rosa, M. & Pignatti, S. (Eds) (2019). Flora d'Italia, volume 4. Bologna: Edagricole.
Hill, M.O., Roy, D.B., Mountford, J.O., & Bunce, R.G.H. (2000). Extending Ellenberg’s indicator values to a new area: an algorithmic approach. Journal of Applied Ecology, 37, 3–15.
Julve, P. (2015). Baseflor. Index botanique, écologique et chorologique de la flore de France (Baseflor. Botanical, ecological and chorological index of the flora of France). Available at http://philippe.julve.pagesperso-orange.fr/catminat.htm [accessed 2022].
Karrer, G. (1992). Österreichische Waldboden-Zustandsinventur. Teil VII: Vegetationsökologische Analysen (Austrian forest soil status inventory. Part VII: Vegetation ecology analyses). Mitteilungen Forstliche Bundesversuchsanstalt Wien, 168, 193–242.

Ellenberg-type indicator values for European plant species are expert-based rankings of plant species according to their ecological optima on main environmental gradients, using ordinal scales defined by Ellenberg et al. (1991). The indicator value for salinity are expressed on an ordinal scale from 0 to 9 defined by Ellenberg et al. (1991). It is a proxy for the concentration in the environment of soluble salts, including sulphates, chlorides and carbonates of sodium, potassium, calcium and magnesium.

Values for individual species were calculated as the means across available national or regional datasets of plant indicator values or were newly assigned based on species co-occurrences in European vegetation plots, see Tichý et al. (2023). Although they have one decimal place, the newly introduced indicator values are compatible with the original Ellenberg values. They can be used for large-scale studies of European flora and vegetation or gap-filling in regional datasets.

Datasets used for compilation include several sources, namely Ellenberg-type indicator values for Great Britain (Hill et al., 2000); France (Julve, 2015); Germany (Ellenberg et al., 2001, taken from Ellenberg & Leuschner, 2010); Czech Republic (Chytrý et al., 2018); Austria (Karrer, 1992); Hungary (Borhidi, 1995); Italy (Guarino & La Rosa, 2019, a corrected version prepared by R. Guarino, values 10–12 were rescaled to the value 9); South Aegean region of Greece (Böhling et al., 2002) and saline habitats in Central Europe (Dítě et al., 2023).

The European values and also the original and taxonomically harmonized regional datasets of Ellenberg-type indicator values are available in the Download section of this website.

Categories

• 0 – not salt tolerant, glycophyte
• 1 – salt tolerant, mostly on low-salt to salt-free soils, but occasionally on slightly salty soils
• 2 – oligohaline, often on soils with very low salt content
• 3 – β-mesohaline, mostly on soils with low salt content
• 4 – α/β-mesohaline, mostly on soils with low to moderate salt content
• 5 – α-mesohaline, mostly on soils with a moderate salt content
• 6 – α-meso/polyhaline, on soils with moderate to high salt content
• 7 – polyhaline, on soils with a high salt content
• 8 – euhaline, on soils with a very high salt content
• 9 – euhaline to hypersaline, on soils with a very high and in dry periods extremely high salt content

Data source and citation

Tichý L., Axmanová I., Dengler J., Guarino R., Jansen F., Midolo G., … Chytrý M. (2023). Ellenberg-type indicator values for European vascular plant species. Journal of Vegetation Science, 34, e13168. https://doi.org/10.1111/jvs.13168

Further references

Böhling, N., Greuter, W. & Raus, T. (2002). Zeigerwerte der Gefäßpflanzen der Südägäis (Griechenland) [Indicator values of the vascular plants in the Southern Aegean (Greece)]. Braun-Blanquetia, 32, 1–108.
Borhidi, A. (1995). Social behaviour types, the naturalness and relative ecological indicator values of the higher plants in the Hungarian flora. Acta Botanica Hungarica, 39, 97–181.
Chytrý M., Tichý L., Dřevojan P., Sádlo J. & Zelený D. (2018). Ellenberg-type indicator values for the Czech flora. Preslia 90: 83–103.
Dítě, D., Šuvada, R., Tóth, T. & Dítě, Z. (2023). Inventory of halophytes in inland central Europe. Preslia, 95.
Ellenberg H., Weber H. E., Düll R., Wirth V., Werner W. & Paulißen D. (1991). Zeigerwerte von Pflanzen in Mitteleuropa. Scripta Geobotanica, 18, 1–248.
Ellenberg, H. & Leuschner, C. (2010). Zeigerwerte der Pflanzen Mitteleuropas. In: Ellenberg H. & Leuschner, C., Vegetation Mitteleuropas mit den Alpen. Stuttgart: Verlag Eugen Ulmer.
Guarino, R., La Rosa, M. & Pignatti, S. (Eds) (2019). Flora d'Italia, volume 4. Bologna: Edagricole.
Hill, M.O., Roy, D.B., Mountford, J.O., & Bunce, R.G.H. (2000). Extending Ellenberg’s indicator values to a new area: an algorithmic approach. Journal of Applied Ecology, 37, 3–15.
Julve, P. (2015). Baseflor. Index botanique, écologique et chorologique de la flore de France (Baseflor. Botanical, ecological and chorological index of the flora of France). Available at http://philippe.julve.pagesperso-orange.fr/catminat.htm [accessed 2022].
Karrer, G. (1992). Österreichische Waldboden-Zustandsinventur. Teil VII: Vegetationsökologische Analysen (Austrian forest soil status inventory. Part VII: Vegetation ecology analyses). Mitteilungen Forstliche Bundesversuchsanstalt Wien, 168, 193–242.

Following Midolo et al. (2023), the indicator value for disturbance frequency is expressed as the log₁₀ mean inverse of return time (in centuries) of disturbance, which is the mean interval between successive disturbance events. Disturbance frequency refers to all possible types of disturbance that may occur in a given habitat, including anthropogenic and natural disturbance as well as grazing and mowing. Because one habitat can be affected by more than one disturbance type, disturbance frequency values were estimated for the most important disturbance types characterizing each habitat. Data are reported as separate values for disturbance affecting the whole plant community (including all vegetation layers) and values considering the herb layer only. This separation accounts for the fact that disturbance regimes in the tree and shrub layers differ in severity and frequency from the disturbance regimes in the herb layer of the same community. For habitats with herbaceous vegetation only, the whole-community values are equal to the herb-layer values.

Data source and citation

Midolo G., Herben T., Axmanová I., Marcenò C., Pätsch R., Bruelheide H., ... & Chytrý M. (2023). Disturbance indicator values for European plants. Global Ecology and Biogeography, 32, 24–34. https://doi.org/10.1111/GEB.13603

Further references

Herben, T., Chytrý, M., & Klimešová, J. (2016). A quest for species‐level indicator values for disturbance. Journal of Vegetation Science, 27(3), 628-636. https://doi.org/10.1111/jvs.12384>
Herben, T., Klimešová, J., & Chytrý, M. (2018). Effects of disturbance frequency and severity on plant traits: an assessment across a temperate flora. Functional Ecology, 32(3), 799-808. https://doi.org/10.1111/1365-2435.13011>

Following Midolo et al. (2023), the indicator value for disturbance frequency is expressed as the log₁₀ mean inverse of return time (in centuries) of disturbance, which is the mean interval between successive disturbance events. Disturbance frequency refers to all possible types of disturbance that may occur in a given habitat, including anthropogenic and natural disturbance as well as grazing and mowing. Because one habitat can be affected by more than one disturbance type, disturbance frequency values were estimated for the most important disturbance types characterizing each habitat. Data are reported as separate values for disturbance affecting the whole plant community (including all vegetation layers) and values considering the herb layer only. This separation accounts for the fact that disturbance regimes in the tree and shrub layers differ in severity and frequency from the disturbance regimes in the herb layer of the same community. For habitats with herbaceous vegetation only, the whole-community values are equal to the herb-layer values.

Data source and citation

Midolo G., Herben T., Axmanová I., Marcenò C., Pätsch R., Bruelheide H., ... & Chytrý M. (2023). Disturbance indicator values for European plants. Global Ecology and Biogeography, 32, 24–34. https://doi.org/10.1111/GEB.13603

Further references

Herben, T., Chytrý, M., & Klimešová, J. (2016). A quest for species‐level indicator values for disturbance. Journal of Vegetation Science, 27(3), 628-636. https://doi.org/10.1111/jvs.12384>
Herben, T., Klimešová, J., & Chytrý, M. (2018). Effects of disturbance frequency and severity on plant traits: an assessment across a temperate flora. Functional Ecology, 32(3), 799-808. https://doi.org/10.1111/1365-2435.13011>

Following Midolo et al. (2023), the indicator value for disturbance severity is expressed as a continuous value ranging from 0 (no change in biomass) to 1 (complete loss of plant cover). Disturbance severity refers to all possible types of disturbance that may occur in a given habitat, including anthropogenic and natural disturbance as well as grazing and mowing. Because one habitat can be affected by more than one disturbance type, disturbance severity values were estimated for the most important disturbance types characterizing each habitat. Data are reported as separate values for disturbance affecting the whole plant community (including all vegetation layers) and values considering the herb layer only. This separation accounts for the fact that disturbance regimes in the tree and shrub layers differ in severity and frequency from the disturbance regimes in the herb layer of the same community. For habitats with herbaceous vegetation only, the whole-community values are equal to the herb-layer values.

Data source and citation

Midolo G., Herben T., Axmanová I., Marcenò C., Pätsch R., Bruelheide H., ... & Chytrý M. (2023). Disturbance indicator values for European plants. Global Ecology and Biogeography, 32, 24–34. https://doi.org/10.1111/GEB.13603

Further references

Herben, T., Chytrý, M., & Klimešová, J. (2016). A quest for species‐level indicator values for disturbance. Journal of Vegetation Science, 27(3), 628-636. https://doi.org/10.1111/jvs.12384>
Herben, T., Klimešová, J., & Chytrý, M. (2018). Effects of disturbance frequency and severity on plant traits: an assessment across a temperate flora. Functional Ecology, 32(3), 799-808. https://doi.org/10.1111/1365-2435.13011>

Following Midolo et al. (2023), the indicator value for disturbance severity is expressed as a continuous value ranging from 0 (no change in biomass) to 1 (complete loss of plant cover). Disturbance severity refers to all possible types of disturbance that may occur in a given habitat, including anthropogenic and natural disturbance as well as grazing and mowing. Because one habitat can be affected by more than one disturbance type, disturbance severity values were estimated for the most important disturbance types characterizing each habitat. Data are reported as separate values for disturbance affecting the whole plant community (including all vegetation layers) and values considering the herb layer only. This separation accounts for the fact that disturbance regimes in the tree and shrub layers differ in severity and frequency from the disturbance regimes in the herb layer of the same community. For habitats with herbaceous vegetation only, the whole-community values are equal to the herb-layer values.

Data source and citation

Midolo G., Herben T., Axmanová I., Marcenò C., Pätsch R., Bruelheide H., ... & Chytrý M. (2023). Disturbance indicator values for European plants. Global Ecology and Biogeography, 32, 24–34. https://doi.org/10.1111/GEB.13603

Further references

Herben, T., Chytrý, M., & Klimešová, J. (2016). A quest for species‐level indicator values for disturbance. Journal of Vegetation Science, 27(3), 628-636. https://doi.org/10.1111/jvs.12384>
Herben, T., Klimešová, J., & Chytrý, M. (2018). Effects of disturbance frequency and severity on plant traits: an assessment across a temperate flora. Functional Ecology, 32(3), 799-808. https://doi.org/10.1111/1365-2435.13011>

Following Midolo et al. (2023), the indicator value for mowing frequency is expressed as the log₁₀ mean inverse of return time (in centuries) of disturbance, which is the mean interval between successive mowing events.

Data source and citation

Midolo G., Herben T., Axmanová I., Marcenò C., Pätsch R., Bruelheide H., ... & Chytrý M. (2023). Disturbance indicator values for European plants. Global Ecology and Biogeography, 32, 24–34. https://doi.org/10.1111/GEB.13603

Further references

Herben, T., Chytrý, M., & Klimešová, J. (2016). A quest for species‐level indicator values for disturbance. Journal of Vegetation Science, 27(3), 628-636. https://doi.org/10.1111/jvs.12384>
Herben, T., Klimešová, J., & Chytrý, M. (2018). Effects of disturbance frequency and severity on plant traits: an assessment across a temperate flora. Functional Ecology, 32(3), 799-808. https://doi.org/10.1111/1365-2435.13011>

Following Midolo et al. (2023), the indicator value for grazing pressure is expressed as a continuous value ranging from 0 (no change in biomass) to 1 (complete loss of plant cover) caused by grazing.

Data source and citation

Midolo G., Herben T., Axmanová I., Marcenò C., Pätsch R., Bruelheide H., ... & Chytrý M. (2023). Disturbance indicator values for European plants. Global Ecology and Biogeography, 32, 24–34. https://doi.org/10.1111/GEB.13603

Further references

Herben, T., Chytrý, M., & Klimešová, J. (2016). A quest for species‐level indicator values for disturbance. Journal of Vegetation Science, 27(3), 628-636. https://doi.org/10.1111/jvs.12384>
Herben, T., Klimešová, J., & Chytrý, M. (2018). Effects of disturbance frequency and severity on plant traits: an assessment across a temperate flora. Functional Ecology, 32(3), 799-808. https://doi.org/10.1111/1365-2435.13011>

Following Midolo et al. (2023), the indicator value for soil distuance is expressed as a continuous value ranging from 0 (no change in biomass) to 1 (complete loss of plant cover) caused by factor causing plant biomass death/removal from soil turning and furrowing.

Data source and citation

Midolo G., Herben T., Axmanová I., Marcenò C., Pätsch R., Bruelheide H., ... & Chytrý M. (2023). Disturbance indicator values for European plants. Global Ecology and Biogeography, 32, 24–34. https://doi.org/10.1111/GEB.13603

Further references

Herben, T., Chytrý, M., & Klimešová, J. (2016). A quest for species‐level indicator values for disturbance. Journal of Vegetation Science, 27(3), 628-636. https://doi.org/10.1111/jvs.12384>
Herben, T., Klimešová, J., & Chytrý, M. (2018). Effects of disturbance frequency and severity on plant traits: an assessment across a temperate flora. Functional Ecology, 32(3), 799-808. https://doi.org/10.1111/1365-2435.13011>

Diagnostic species are characterized by a concentration of their occurrence in the stands belonging to the target vegetation unit while being rare or absent in other vegetation units. For the European vegetation classes of the EuroVegChecklist (Mucina et al. 2016), the list of these species was compiled from various European literature sources, especially syntaxonomic monographs and revisions containing extensive synthetic phytosociological tables. Expert opinion from EuroVegChecklist authors was used to judge problematic cases. Some species were assigned to more than one class. Unlike for the EUNIS habitat types, no statistical approach was used to determine diagnostic species for European vegetation classes.

Data source and citation

Mucina L., Bültmann H., Dierßen K., Theurillat J.-P., Raus T., Čarni A., … Tichý L. (2016). Vegetation of Europe: Hierarchical floristic classification system of vascular plant, bryophyte, lichen, and algal communities. Applied Vegetation Science, 19(Suppl. 1), 3–264. https://doi.org/10.1111/avsc.12257 (Mucina et al. 2016, version 3, 2024-01-01)

Diagnostic species are characterized by a concentration of their occurrence in the stands belonging to the target habitat type while being rare or absent in other habitat types. For the habitat types of the EUNIS classification (Chytrý et al. 2020), these species were determined based on the calculation of fidelity of each species to a group of vegetation plots representing the target habitat type in a geographically and ecologically stratified selection of plots from the European Vegetation Archive (Chytrý et al. 2016). Fidelity was calculated using the phi coefficient of association (Sokal & Rohlf, 1995; Chytrý et al., 2002) standardized as if each habitat was represented by the same number of plots (Tichý & Chytrý, 2006). The species with a value of phi greater than 0.15 for a particular habitat were considered as diagnostic for this habitat. The statistical significance of the species–habitat association was tested using Fisher's exact test (Sokal & Rohlf, 1995), and if not significant at p < 0.05, the species was excluded from the list of diagnostic species (Tichý & Chytrý, 2006).

Categories

• MA211 Arctic coastal saltmarsh
• MA221 Atlantic saltmarsh driftline
• MA222 Atlantic upper saltmarsh
• MA223 Atlantic upper-mid saltmarsh and saline and brackish reed, rush and sedge bed
• MA224 Atlantic mid-low saltmarsh
• MA225 Atlantic pioneer saltmarsh
• MA232 Baltic coastal meadow
• MA241 Black Sea littoral saltmarsh
• MA251 Mediterranean upper saltmarsh
• MA252 Mediterranean upper-mid saltmarsh and saline and brackish reed, rush and sedge bed
• MA253 Mediterranean mid-low saltmarsh
• N11 Atlantic, Baltic and Arctic sand beach
• N12 Mediterranean and Black Sea sand beach
• N13 Atlantic and Baltic shifting coastal dune
• N14 Mediterranean, Macaronesian and Black Sea shifting coastal dune
• N15 Atlantic and Baltic coastal dune grassland (grey dune)
• N16 Mediterranean and Macaronesian coastal dune grassland (grey dune)
• N17 Black Sea coastal dune grassland (grey dune)
• N18 Atlantic and Baltic coastal Empetrum heath
• N19 Atlantic coastal Calluna and Ulex heath
• N1A Atlantic and Baltic coastal dune scrub
• N1B Mediterranean and Black Sea coastal dune scrub
• N1C Macaronesian coastal dune scrub
• N1D Atlantic and Baltic broad-leaved coastal dune forest
• N1E Black Sea broad-leaved coastal dune forest
• N1F Baltic coniferous coastal dune forest
• N1G Mediterranean coniferous coastal dune forest
• N1H Atlantic and Baltic moist and wet dune slack
• N1J Mediterranean and Black Sea moist and wet dune slack
• N21 Atlantic, Baltic and Arctic coastal shingle beach
• N22 Mediterranean and Black Sea coastal shingle beach
• N31 Atlantic and Baltic rocky sea cliff and shore
• N32 Mediterranean and Black Sea rocky sea cliff and shore
• N33 Macaronesian rocky sea cliff and shore
• N34 Atlantic and Baltic soft sea cliff
• N35 Mediterranean and Black Sea soft sea cliff
• P2N Spring
• P3a Brackish-water vegetation
• P3b Fresh-water small pleustophyte vegetation
• P3c Fresh-water large pleustophyte vegetation
• P3d Fresh-water submerged vegetation
• P3e Fresh-water nymphaeid vegetation
• P3f Oligotrophic-water vegetation
• P3g Dystrophic-water vegetation
• P3h Stonewort vegetation
• Q11 Raised bog
• Q12 Blanket bog
• Q21 Oceanic valley mire
• Q22 Poor fen
• Q23 Relict mire of Mediterranean mountains
• Q24 Intermediate fen and soft-water spring mire
• Q25 Non-calcareous quaking mire
• Q31 Palsa mire
• Q41 Alkaline, calcareous, carbonate-rich small-sedge spring fen
• Q42 Extremely rich moss-sedge fen
• Q43 Tall-sedge base-rich fen
• Q44 Calcareous quaking mire
• Q45 Arctic-alpine rich fen
• Q46 Carpathian travertine fen with halophytes
• Q51 Tall-helophyte bed
• Q52 Small-helophyte bed
• Q53 Tall-sedge bed
• Q54 Inland saline or brackish helophyte bed
• Q61 Periodically exposed shore with stable, eutrophic sediments with pioneer or ephemeral vegetation
• Q62 Periodically exposed shore with stable, mesotrophic sediments with pioneer or ephemeral vegetation
• Q63 Periodically exposed saline shore with pioneer or ephemeral vegetation
• R11 Pannonian and Pontic sandy steppe
• R12 Cryptogam- and annual-dominated vegetation on siliceous rock outcrops
• R13 Cryptogam- and annual-dominated vegetation on calcareous and ultramafic rock outcrops
• R14 Perennial rocky grassland of the Italian Peninsula
• R15 Continental dry rocky steppic grassland and dwarf scrub on chalk outcrops
• R16 Perennial rocky grassland of Central and South-Eastern Europe
• R17 Heavy-metal dry grassland of the Balkans
• R18 Perennial rocky calcareous grassland of subatlantic-submediterranean Europe
• R19 Dry steppic submediterranean pasture of the Amphi-Adriatic region
• R1A Semi-dry perennial calcareous grassland (meadow steppe)
• R1B Continental dry grassland (true steppe)
• R1C Desert steppe
• R1D Mediterranean closely grazed dry grassland
• R1E Mediterranean tall perennial dry grassland
• R1F Mediterranean annual-rich dry grassland
• R1G Iberian oromediterranean siliceous dry grassland
• R1H Iberian oromediterranean basiphilous dry grassland
• R1J Cyrno-Sardean oromediterranean siliceous dry grassland
• R1K Balkan and Anatolian oromediterranean dry grassland
• R1L Madeiran oromediterranean siliceous dry grassland
• R1M Lowland to montane, dry to mesic grassland usually dominated by Nardus stricta
• R1N Open Iberian supramediterranean dry acid and neutral grassland
• R1P Oceanic to subcontinental inland sand grassland on dry acid and neutral soils
• R1Q Inland sanddrift and dune with siliceous grassland
• R1R Mediterranean to Atlantic open, dry, acid and neutral grassland
• R1S Heavy-metal grassland in Western and Central Europe
• R1T Azorean open, dry, acid to neutral grassland
• R21 Mesic permanent pasture of lowlands and mountains
• R22 Low and medium altitude hay meadow
• R23 Mountain hay meadow
• R24 Iberian summer pasture (vallicar)
• R31 Mediterranean tall humid inland grassland
• R32 Mediterranean short moist grassland of lowlands
• R33 Mediterranean short moist grassland of mountains
• R34 Submediterranean moist meadow
• R35 Moist or wet mesotrophic to eutrophic hay meadow
• R36 Moist or wet mesotrophic to eutrophic pasture
• R37 Temperate and boreal moist or wet oligotrophic grassland
• R41 Snow-bed vegetation
• R42 Boreal and Arctic acidophilous alpine grassland
• R43 Temperate acidophilous alpine grassland
• R44 Arctic-alpine calcareous grassland
• R45 Alpine and subalpine calcareous grassland of the Balkans and Apennines
• R51 Thermophilous forest fringe of base-rich soils
• R52 Forest fringe of acidic nutrient-poor soils
• R53 Macaronesian thermophilous forest fringe
• R54 Pteridium aquilinum vegetation
• R55 Lowland moist or wet tall-herb and fern fringe
• R56 Montane to subalpine moist or wet tall-herb and fern fringe
• R57 Herbaceous forest clearing vegetation
• R61 Mediterranean inland salt steppe
• R62 Continental inland salt steppe
• R63 Temperate inland salt marsh
• R64 Semi-desert salt pan
• R65 Continental subsaline alluvial pasture and meadow
• S11 Shrub tundra
• S12 Moss and lichen tundra
• S21 Subarctic and alpine dwarf Salix scrub
• S22 Alpine and subalpine ericoid heath
• S23 Alpine and subalpine Juniperus scrub
• S24 Subalpine genistoid scrub of the Amphi-Adriatic region
• S25 Subalpine and subarctic deciduous scrub
• S26 Subalpine Pinus mugo scrub
• S31 Lowland to montane temperate and submediterranean Juniperus scrub
• S32 Temperate Rubus scrub
• S33 Lowland to montane temperate and submediterranean genistoid scrub
• S34 Balkan-Anatolian submontane genistoid scrub
• S35 Temperate and submediterranean thorn scrub
• S36 Low steppic scrub
• S37 Corylus avellana scrub
• S38 Temperate forest clearing scrub
• S41 Wet heath
• S42 Dry heath
• S43 Macaronesian heath
• S51 Mediterranean maquis and arborescent matorral
• S52 Submediterranean pseudomaquis
• S53 Spartium junceum scrub
• S54 Thermomediterranean arid scrub
• S61 Western basiphilous garrigue
• S62 Western acidophilous garrigue
• S63 Eastern garrigue
• S64 Macaronesian garrigue
• S65 Mediterranean gypsum scrub
• S66 Mediterranean halo-nitrophilous scrub
• S67 Aralo-Caspian semi-desert
• S68 Semi-desert sand dune with sparse scrub
• S71 Western Mediterranean spiny heath
• S72 Eastern Mediterranean spiny heath (phrygana)
• S73 Western Mediterranean mountain hedgehog-heath
• S74 Central Mediterranean mountain hedgehog-heath
• S75 Eastern Mediterranean mountain hedgehog-heath
• S76 Canarian mountain hedgehog-heath
• S81 Canarian xerophytic scrub
• S82 Madeiran xerophytic scrub
• S91 Temperate riparian scrub
• S92 Salix fen scrub
• S93 Mediterranean riparian scrub
• S94 Semi-desert riparian scrub
• T11 Temperate Salix and Populus riparian forest
• T12 Alnus glutinosa-Alnus incana forest on riparian and mineral soils
• T13 Temperate hardwood riparian forest
• T14 Mediterranean and Macaronesian riparian forest
• T15 Broadleaved swamp forest on non-acid peat
• T16 Broadleaved mire forest on acid peat
• T17 Fagus forest on non-acid soils
• T18 Fagus forest on acid soils
• T19 Temperate and submediterranean thermophilous deciduous forest
• T1A Mediterranean thermophilous deciduous forest
• T1B Acidophilous Quercus forest
• T1C Temperate and boreal mountain Betula and Populus tremula forest on mineral soils
• T1D Southern European mountain Betula and Populus tremula forest on mineral soils
• T1E Carpinus and Quercus mesic deciduous forest
• T1F Ravine forest
• T1G Alnus cordata forest
• T1H Broadleaved deciduous plantation of non site-native trees
• T21 Mediterranean evergreen Quercus forest
• T22 Mainland laurophyllous forest
• T23 Macaronesian laurophyllous forest
• T24 Olea europaea-Ceratonia siliqua forest
• T25 Phoenix theophrasti vegetation
• T26 Phoenix canariensis vegetation
• T27 Ilex aquifolium forest
• T28 Macaronesian heathy forest
• T29 Broadleaved evergreen plantation of non site-native trees
• T31 Temperate mountain Picea forest
• T32 Temperate mountain Abies forest
• T33 Mediterranean mountain Abies forest
• T34 Temperate subalpine Larix, Pinus cembra and Pinus uncinata forest
• T35 Temperate continental Pinus sylvestris forest
• T36 Temperate and submediterranean montane Pinus sylvestris-Pinus nigra forest
• T37 Mediterranean montane Pinus sylvestris-Pinus nigra forest
• T38 Mediterranean montane Cedrus forest
• T39 Mediterranean and Balkan subalpine Pinus heldreichii-Pinus peuce forest
• T3A Mediterranean lowland to submontane Pinus forest
• T3B Pinus canariensis forest
• T3C Taxus baccata forest
• T3D Mediterranean Cupressaceae forest
• T3E Macaronesian Juniperus forest
• T3F Dark taiga
• T3G Pinus sylvestris light taiga
• T3H Larix light taiga
• T3J Pinus and Larix mire forest
• T3K Picea mire forest
• T3M Coniferous plantation of non site-native trees
• U21 Boreal and Arctic siliceous scree and block field
• U22 Temperate high-mountain siliceous scree
• U23 Temperate, lowland to montane siliceous scree
• U24 Mediterranean siliceous scree
• U25 Boreal and Arctic base-rich scree and block field
• U26 Temperate high-mountain base-rich scree and moraine
• U27 Temperate, lowland to montane base-rich scree
• U28 Western Mediterranean base-rich scree
• U29 Eastern Mediterranean base-rich scree
• U2A Crimean base-rich scree
• U32 Temperate high-mountain siliceous inland cliff
• U33 Temperate, lowland to montane siliceous inland cliff
• U34 Mediterranean siliceous inland cliff
• U35 Boreal and Arctic base-rich inland cliff
• U36 Temperate high-mountain base-rich inland cliff
• U37 Temperate, lowland to montane base-rich inland cliff
• U38 Mediterranean base-rich inland cliff
• U3A Temperate ultramafic inland cliff
• U3B Mediterranean ultramafic inland cliff
• U3C Macaronesian inland cliff
• U3D Wet inland cliff
• U52 Polar desert
• U61 Subarctic volcanic field
• U62 Mediterranean, Macaronesian and temperate volcanic field
• U71 Unvegetated or sparsely vegetated gravel bar in montane and alpine regions
• U72 Unvegetated or sparsely vegetated gravel bar in the Mediterranean region
• V11 Intensive unmixed crops
• V12 Mixed crops of market gardens and horticulture
• V13 Arable land with unmixed crops grown by low-intensity agricultural methods
• V14 Inundated or inundatable cropland, including rice fields
• V15 Bare tilled, fallow or recently abandoned arable land
• V32 Mediterranean subnitrophilous annual grassland
• V33 Dry Mediterranean land with unpalatable non-vernal herbaceous vegetation
• V34 Trampled xeric grassland with annuals
• V35 Trampled mesophilous grassland with annuals
• V37 Annual anthropogenic herbaceous vegetation
• V38 Dry perennial anthropogenic herbaceous vegetation
• V39 Mesic perennial anthropogenic herbaceous vegetation

Data source and citation

Chytrý, M., Tichý, L., Hennekens, S. M., Knollová, I., Janssen, J. A. M., Rodwell, J. S., … Schaminée, J. H. J. (2020). EUNIS Habitat Classification: expert system, characteristic species combinations and distribution maps of European habitats. Applied Vegetation Science, 23(4), 648–675. https://doi.org/10.1111/avsc.12519 – Version 2021-06-01: https://doi.org/10.5281/zenodo.4812736

Further references

Chytrý, M., Tichý, L., Holt, J., & Botta-Dukát, Z. (2002). Determination of diagnostic species with statistical fidelity measures. Journal of Vegetation Science, 13(1), 79–90. https://doi.org/10.1111/j.1654-1103.2002.tb02025.x
Chytrý, M., Hennekens, S. M., Jiménez-Alfaro, B., Knollová, I., Dengler, J., Jansen, F., … Yamalov, S. (2016). European Vegetation Archive (EVA): an integrated database of European vegetation plots. Applied Vegetation Science, 19(1), 173–180. https://doi.org/10.1111/avsc.12191
Sokal, R. R., & Rohlf, F. J. (1995). Biometry, 3rd edition. New York, NY: Freeman.
Tichý, L., & Chytrý, M. (2006). Statistical determination of diagnostic species for site groups of unequal size. Journal of Vegetation Science, 17(6), 809–818. https://doi.org/10.1111/j.1654-1103.2006.tb02504.x

Constant species are characterized by frequent occurrences in stands belonging to the target vegetation unit, but unlike diagnostic species, they can also commonly occur in other vegetation units. They were determined for the habitat types of the EUNIS classification (Chytrý et al. 2020) based on the calculation of the percentage frequency (constancy) of each species in a group of vegetation plots representing the target habitat type in a geographically and ecologically stratified selection of plots of all habitat types extracted from the European Vegetation Archive (Chytrý et al. 2016). The species with an occurrence frequency in the habitat type higher than 10% were considered as constant taxa.

Categories

• MA211 Arctic coastal saltmarsh
• MA221 Atlantic saltmarsh driftline
• MA222 Atlantic upper saltmarsh
• MA223 Atlantic upper-mid saltmarsh and saline and brackish reed, rush and sedge bed
• MA224 Atlantic mid-low saltmarsh
• MA225 Atlantic pioneer saltmarsh
• MA232 Baltic coastal meadow
• MA241 Black Sea littoral saltmarsh
• MA251 Mediterranean upper saltmarsh
• MA252 Mediterranean upper-mid saltmarsh and saline and brackish reed, rush and sedge bed
• MA253 Mediterranean mid-low saltmarsh
• N11 Atlantic, Baltic and Arctic sand beach
• N12 Mediterranean and Black Sea sand beach
• N13 Atlantic and Baltic shifting coastal dune
• N14 Mediterranean, Macaronesian and Black Sea shifting coastal dune
• N15 Atlantic and Baltic coastal dune grassland (grey dune)
• N16 Mediterranean and Macaronesian coastal dune grassland (grey dune)
• N17 Black Sea coastal dune grassland (grey dune)
• N18 Atlantic and Baltic coastal Empetrum heath
• N19 Atlantic coastal Calluna and Ulex heath
• N1A Atlantic and Baltic coastal dune scrub
• N1B Mediterranean and Black Sea coastal dune scrub
• N1C Macaronesian coastal dune scrub
• N1D Atlantic and Baltic broad-leaved coastal dune forest
• N1E Black Sea broad-leaved coastal dune forest
• N1F Baltic coniferous coastal dune forest
• N1G Mediterranean coniferous coastal dune forest
• N1H Atlantic and Baltic moist and wet dune slack
• N1J Mediterranean and Black Sea moist and wet dune slack
• N21 Atlantic, Baltic and Arctic coastal shingle beach
• N22 Mediterranean and Black Sea coastal shingle beach
• N31 Atlantic and Baltic rocky sea cliff and shore
• N32 Mediterranean and Black Sea rocky sea cliff and shore
• N33 Macaronesian rocky sea cliff and shore
• N34 Atlantic and Baltic soft sea cliff
• N35 Mediterranean and Black Sea soft sea cliff
• P2N Spring
• P3a Brackish-water vegetation
• P3b Fresh-water small pleustophyte vegetation
• P3c Fresh-water large pleustophyte vegetation
• P3d Fresh-water submerged vegetation
• P3e Fresh-water nymphaeid vegetation
• P3f Oligotrophic-water vegetation
• P3g Dystrophic-water vegetation
• P3h Stonewort vegetation
• Q11 Raised bog
• Q12 Blanket bog
• Q21 Oceanic valley mire
• Q22 Poor fen
• Q23 Relict mire of Mediterranean mountains
• Q24 Intermediate fen and soft-water spring mire
• Q25 Non-calcareous quaking mire
• Q31 Palsa mire
• Q41 Alkaline, calcareous, carbonate-rich small-sedge spring fen
• Q42 Extremely rich moss-sedge fen
• Q43 Tall-sedge base-rich fen
• Q44 Calcareous quaking mire
• Q45 Arctic-alpine rich fen
• Q46 Carpathian travertine fen with halophytes
• Q51 Tall-helophyte bed
• Q52 Small-helophyte bed
• Q53 Tall-sedge bed
• Q54 Inland saline or brackish helophyte bed
• Q61 Periodically exposed shore with stable, eutrophic sediments with pioneer or ephemeral vegetation
• Q62 Periodically exposed shore with stable, mesotrophic sediments with pioneer or ephemeral vegetation
• Q63 Periodically exposed saline shore with pioneer or ephemeral vegetation
• R11 Pannonian and Pontic sandy steppe
• R12 Cryptogam- and annual-dominated vegetation on siliceous rock outcrops
• R13 Cryptogam- and annual-dominated vegetation on calcareous and ultramafic rock outcrops
• R14 Perennial rocky grassland of the Italian Peninsula
• R15 Continental dry rocky steppic grassland and dwarf scrub on chalk outcrops
• R16 Perennial rocky grassland of Central and South-Eastern Europe
• R17 Heavy-metal dry grassland of the Balkans
• R18 Perennial rocky calcareous grassland of subatlantic-submediterranean Europe
• R19 Dry steppic submediterranean pasture of the Amphi-Adriatic region
• R1A Semi-dry perennial calcareous grassland (meadow steppe)
• R1B Continental dry grassland (true steppe)
• R1C Desert steppe
• R1D Mediterranean closely grazed dry grassland
• R1E Mediterranean tall perennial dry grassland
• R1F Mediterranean annual-rich dry grassland
• R1G Iberian oromediterranean siliceous dry grassland
• R1H Iberian oromediterranean basiphilous dry grassland
• R1J Cyrno-Sardean oromediterranean siliceous dry grassland
• R1K Balkan and Anatolian oromediterranean dry grassland
• R1L Madeiran oromediterranean siliceous dry grassland
• R1M Lowland to montane, dry to mesic grassland usually dominated by Nardus stricta
• R1N Open Iberian supramediterranean dry acid and neutral grassland
• R1P Oceanic to subcontinental inland sand grassland on dry acid and neutral soils
• R1Q Inland sanddrift and dune with siliceous grassland
• R1R Mediterranean to Atlantic open, dry, acid and neutral grassland
• R1S Heavy-metal grassland in Western and Central Europe
• R1T Azorean open, dry, acid to neutral grassland
• R21 Mesic permanent pasture of lowlands and mountains
• R22 Low and medium altitude hay meadow
• R23 Mountain hay meadow
• R24 Iberian summer pasture (vallicar)
• R31 Mediterranean tall humid inland grassland
• R32 Mediterranean short moist grassland of lowlands
• R33 Mediterranean short moist grassland of mountains
• R34 Submediterranean moist meadow
• R35 Moist or wet mesotrophic to eutrophic hay meadow
• R36 Moist or wet mesotrophic to eutrophic pasture
• R37 Temperate and boreal moist or wet oligotrophic grassland
• R41 Snow-bed vegetation
• R42 Boreal and Arctic acidophilous alpine grassland
• R43 Temperate acidophilous alpine grassland
• R44 Arctic-alpine calcareous grassland
• R45 Alpine and subalpine calcareous grassland of the Balkans and Apennines
• R51 Thermophilous forest fringe of base-rich soils
• R52 Forest fringe of acidic nutrient-poor soils
• R53 Macaronesian thermophilous forest fringe
• R54 Pteridium aquilinum vegetation
• R55 Lowland moist or wet tall-herb and fern fringe
• R56 Montane to subalpine moist or wet tall-herb and fern fringe
• R57 Herbaceous forest clearing vegetation
• R61 Mediterranean inland salt steppe
• R62 Continental inland salt steppe
• R63 Temperate inland salt marsh
• R64 Semi-desert salt pan
• R65 Continental subsaline alluvial pasture and meadow
• S11 Shrub tundra
• S12 Moss and lichen tundra
• S21 Subarctic and alpine dwarf Salix scrub
• S22 Alpine and subalpine ericoid heath
• S23 Alpine and subalpine Juniperus scrub
• S24 Subalpine genistoid scrub of the Amphi-Adriatic region
• S25 Subalpine and subarctic deciduous scrub
• S26 Subalpine Pinus mugo scrub
• S31 Lowland to montane temperate and submediterranean Juniperus scrub
• S32 Temperate Rubus scrub
• S33 Lowland to montane temperate and submediterranean genistoid scrub
• S34 Balkan-Anatolian submontane genistoid scrub
• S35 Temperate and submediterranean thorn scrub
• S36 Low steppic scrub
• S37 Corylus avellana scrub
• S38 Temperate forest clearing scrub
• S41 Wet heath
• S42 Dry heath
• S43 Macaronesian heath
• S51 Mediterranean maquis and arborescent matorral
• S52 Submediterranean pseudomaquis
• S53 Spartium junceum scrub
• S54 Thermomediterranean arid scrub
• S61 Western basiphilous garrigue
• S62 Western acidophilous garrigue
• S63 Eastern garrigue
• S64 Macaronesian garrigue
• S65 Mediterranean gypsum scrub
• S66 Mediterranean halo-nitrophilous scrub
• S67 Aralo-Caspian semi-desert
• S68 Semi-desert sand dune with sparse scrub
• S71 Western Mediterranean spiny heath
• S72 Eastern Mediterranean spiny heath (phrygana)
• S73 Western Mediterranean mountain hedgehog-heath
• S74 Central Mediterranean mountain hedgehog-heath
• S75 Eastern Mediterranean mountain hedgehog-heath
• S76 Canarian mountain hedgehog-heath
• S81 Canarian xerophytic scrub
• S82 Madeiran xerophytic scrub
• S91 Temperate riparian scrub
• S92 Salix fen scrub
• S93 Mediterranean riparian scrub
• S94 Semi-desert riparian scrub
• T11 Temperate Salix and Populus riparian forest
• T12 Alnus glutinosa-Alnus incana forest on riparian and mineral soils
• T13 Temperate hardwood riparian forest
• T14 Mediterranean and Macaronesian riparian forest
• T15 Broadleaved swamp forest on non-acid peat
• T16 Broadleaved mire forest on acid peat
• T17 Fagus forest on non-acid soils
• T18 Fagus forest on acid soils
• T19 Temperate and submediterranean thermophilous deciduous forest
• T1A Mediterranean thermophilous deciduous forest
• T1B Acidophilous Quercus forest
• T1C Temperate and boreal mountain Betula and Populus tremula forest on mineral soils
• T1D Southern European mountain Betula and Populus tremula forest on mineral soils
• T1E Carpinus and Quercus mesic deciduous forest
• T1F Ravine forest
• T1G Alnus cordata forest
• T1H Broadleaved deciduous plantation of non site-native trees
• T21 Mediterranean evergreen Quercus forest
• T22 Mainland laurophyllous forest
• T23 Macaronesian laurophyllous forest
• T24 Olea europaea-Ceratonia siliqua forest
• T25 Phoenix theophrasti vegetation
• T26 Phoenix canariensis vegetation
• T27 Ilex aquifolium forest
• T28 Macaronesian heathy forest
• T29 Broadleaved evergreen plantation of non site-native trees
• T31 Temperate mountain Picea forest
• T32 Temperate mountain Abies forest
• T33 Mediterranean mountain Abies forest
• T34 Temperate subalpine Larix, Pinus cembra and Pinus uncinata forest
• T35 Temperate continental Pinus sylvestris forest
• T36 Temperate and submediterranean montane Pinus sylvestris-Pinus nigra forest
• T37 Mediterranean montane Pinus sylvestris-Pinus nigra forest
• T38 Mediterranean montane Cedrus forest
• T39 Mediterranean and Balkan subalpine Pinus heldreichii-Pinus peuce forest
• T3A Mediterranean lowland to submontane Pinus forest
• T3B Pinus canariensis forest
• T3C Taxus baccata forest
• T3D Mediterranean Cupressaceae forest
• T3E Macaronesian Juniperus forest
• T3F Dark taiga
• T3G Pinus sylvestris light taiga
• T3H Larix light taiga
• T3J Pinus and Larix mire forest
• T3K Picea mire forest
• T3M Coniferous plantation of non site-native trees
• U21 Boreal and Arctic siliceous scree and block field
• U22 Temperate high-mountain siliceous scree
• U23 Temperate, lowland to montane siliceous scree
• U24 Mediterranean siliceous scree
• U25 Boreal and Arctic base-rich scree and block field
• U26 Temperate high-mountain base-rich scree and moraine
• U27 Temperate, lowland to montane base-rich scree
• U28 Western Mediterranean base-rich scree
• U29 Eastern Mediterranean base-rich scree
• U2A Crimean base-rich scree
• U32 Temperate high-mountain siliceous inland cliff
• U33 Temperate, lowland to montane siliceous inland cliff
• U34 Mediterranean siliceous inland cliff
• U35 Boreal and Arctic base-rich inland cliff
• U36 Temperate high-mountain base-rich inland cliff
• U37 Temperate, lowland to montane base-rich inland cliff
• U38 Mediterranean base-rich inland cliff
• U3A Temperate ultramafic inland cliff
• U3B Mediterranean ultramafic inland cliff
• U3C Macaronesian inland cliff
• U3D Wet inland cliff
• U52 Polar desert
• U61 Subarctic volcanic field
• U62 Mediterranean, Macaronesian and temperate volcanic field
• U71 Unvegetated or sparsely vegetated gravel bar in montane and alpine regions
• U72 Unvegetated or sparsely vegetated gravel bar in the Mediterranean region
• V11 Intensive unmixed crops
• V12 Mixed crops of market gardens and horticulture
• V13 Arable land with unmixed crops grown by low-intensity agricultural methods
• V14 Inundated or inundatable cropland, including rice fields
• V15 Bare tilled, fallow or recently abandoned arable land
• V32 Mediterranean subnitrophilous annual grassland
• V33 Dry Mediterranean land with unpalatable non-vernal herbaceous vegetation
• V34 Trampled xeric grassland with annuals
• V35 Trampled mesophilous grassland with annuals
• V37 Annual anthropogenic herbaceous vegetation
• V38 Dry perennial anthropogenic herbaceous vegetation
• V39 Mesic perennial anthropogenic herbaceous vegetation

Data source and citation

Chytrý, M., Tichý, L., Hennekens, S. M., Knollová, I., Janssen, J. A. M., Rodwell, J. S., … Schaminée, J. H. J. (2020). EUNIS Habitat Classification: expert system, characteristic species combinations and distribution maps of European habitats. Applied Vegetation Science, 23(4), 648–675. https://doi.org/10.1111/avsc.12519 – Version 2021-06-01: https://doi.org/10.5281/zenodo.4812736

Further references

Chytrý M., Hennekens S.M., Jiménez-Alfaro B., Knollová I., Dengler J., Jansen F., … Yamalov S. (2016). European Vegetation Archive (EVA): an integrated database of European vegetation plots. Applied Vegetation Science, 19(1), 173–180. https://doi.org/10.1111/avsc.12191

Species association to broadly defined habitats is based on species occurrences reported for finer units, either vegetation types or habitats. We compiled available data from several sources, Sádlo et al. (2007), Mucina et al. (2016), Guarino et al. (2019). Final list of habitats include 18 broad habitats.

Categories

• Marine
• Coastal saltmarsh
• Coastal beach, dune or shingle
• Coastal cliff
• Aquatic
• Spring
• Wetland
• Mire
• Grassland (non-alpine, non-saline)
• Alpine-subalpine and arctic grassland
• Saline vegetation
• Semi-desert
• Heathland
• Oromediterranean scrub
• Scrub
• Forest
• Sparsely vegetated (incl. rock and scree)
• Synanthropic

Data source and citation

Axmanová, I. (2022). Broad habitat. – www.FloraVeg.EU.

Further references

Guarino, R., La Rosa, M. & Pignatti, S. (Eds) (2019). Flora d'Italia, volume 4. Bologna: Edagricole.
Mucina, L., Bültmann, H., Dierßen, K., Theurillat, J.-P., Raus, T., Čarni, A., … Tichý L. (2016). Vegetation of Europe: Hierarchical floristic classification system of vascular plant, bryophyte, lichen, and algal communities. Applied Vegetation Science, 19(Suppl. 1), 3–264. https://doi.org/10.1111/avsc.12257
Sádlo, J., Chytrý, M. & Pyšek, P. (2007). Regional species pools of vascular plants in habitats of the Czech Republic. Preslia, 79, 303–321.