Research Article |
Corresponding author: Naim Berisha ( naim.berisha@uni-pr.edu ) Academic editor: Giovanni Spampinato
© 2023 Naim Berisha, Renata Ćušterevska, Fadil Millaku, Vlado Matevski.
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:
Berisha N, Ćušterevska R, Millaku F, Matevski V (2023) Blysmo compressi-Eriophoretum latifoliae ass. nova, a new association of the Caricion fuscae alliance from the Sharri Mountains. Plant Sociology 60(1): 37-49. https://doi.org/10.3897/pls2023601/04
|
The sedge-moss vegetation of the moderately to low calcium-rich slightly acidic fens of the Caricion fuscae alliance depends on a very specific combination of ecological and climatic conditions to thrive. Until recently, the classification of this vegetation group was complicated by its rarity on the southern edges of its range in Europe. As part of a larger database of phytocenological relevés carried out in Mt. Luboten, we came across an interesting group of 15 relevés on fen vegetation sites. We were curious to know if this plant community was a previously known association or if it might represent something new within this alliance. We compiled a separate dataset at JUICE that includes four plant communities from this alliance, along with our 15 original releves. The classification was based on modified TWINSPAN and beta-flexible clustering as a numerical classification method, with OPTIMCLASS determining the appropriate number of clusters. Five associations were clearly delineated, with the four associations taken from the literature sources clearly grouped individually and a new, fifth association appearing as separate, with completely unique characteristics. This new association: Blysmo compressi-Eriophoretum latifoliae occurs at elevations of ~ 1650 m a.s.l. on NE and NW slopes of the mountain. With this work we offer the description of a new high-mountain fen association. These associations may play an important syntaxonomic role as more Balkan data become available on this alliance. The sedge-moss and fen vegetation in the Balkans is particularly rare and characterised by a very diverse and specific vegetation, so it rightly deserves more attention from vegetation scientists and conservation authorities.
Biodiversity conservation, Kosovo, phytosociology, plant taxonomy, Supervised vegetation classification
Fen plant communities are known to be of great importance for biodiversity, as these natural habitats are among the most endangered in continental Europe. At the same time they act as effective carbon sinks (
From a syntaxonomic point of view, European fens are traditionally assigned to the class Scheuchzerio palustris-Caricetea fuscae Tüxen 1937 (
The investigated area includes subalpine fen vegetation patches in Mt. Luboten, a prominent mountain massif in the Sharri Mountains on the border area between Kosovo and North Macedonia (Figure
The red dot (right image) indicates the location of the study area in Mt. Luboten in Kosovo. The small red dots (left image) in the wider map indicate the distance between Blysmo compressi-Eriophoretum latifoliae ass. nova in Sharri Mts. and Blysmus compressus & Juncus thomasii in Skasmada, Greece.
To clarify the syntaxonomic position of this interesting plant community, which consisted of 15 original relevés, we created a working database on JUICE (
Using OptimClass analysis (
To assess and compare the site conditions of each of the analyzed plant communities in our dataset, we relied on Ellenberg indicator values (
Based on OPTIMCLASS analysis, modified TWINSPAN Classification (
The hierarchial classification of the Scheuchzerio palustris-Caricetea fuscae Tx. 1937 – database relevés in a form of dendrogram of the modified TWINSPAN analysis. We were able to finally distinguish five clusters, that derived from 38 relevés. Cluster 1 = 5 relevés from the Association Blysmus compressus & Juncus thomasii Quezel 1964. Cluster 2 = 15 original relevés from Mt. Luboten (Blysmo compressi-Eriophoretum latifoliae ass. nova); Cluster 3 = 8 relevés from Caricetum macedonicae (Horv. 36)
Analytical table of the association: Blysmo compressi-Eriophoretum latifoliae ass. nova.
Relevé no. | 1 | 2 | 3 | 4 | 5 | 6 | 7* | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | Constancy level |
Original relevé no. | 297 | 298 | 300 | 306 | 311 | 299 | 302 | 309 | 304 | 310 | 301 | 308 | 303 | 305 | 307 | |
Relevé area (m2) | 25 | 25 | 25 | 25 | 25 | 9 | 25 | 25 | 25 | 25 | 100 | 100 | 25 | 25 | 25 | |
Cover in tot. (%) | 98 | 95 | 100 | 95 | 98 | 98 | 98 | 98 | 95 | 95 | 100 | 95 | 98 | 95 | 95 | |
Characteristic and different. species | ||||||||||||||||
Eriophorum latifolium | 3 | 3 | 3 | 3 | 4 | 4 | 4 | 4 | 3 | 3 | 1 | 2 | 4 | 5 | 5 | V |
Juncus conglomeratus | 1 | 2 | 2 | 2 | 3 | 3 | 2 | 1 | 2 | 2 | 1 | 1 | 1 | 2 | 2 | V |
Caltha palustris | 2 | 2 | 3 | 3 | 1 | 1 | 1 | 2 | 2 | 2 | 2 | 1 | 1 | 1 | 1 | V |
Blysmus compressus * | 2 | + | 1 | 1 | 2 | 2 | 3 | 3 | 3 | 3 | 4 | 4 | 1 | + | + | V |
Alchemilla hybrida | 2 | 1 | 2 | 1 | 1 | 1 | 2 | 1 | 2 | 2 | 2 | 1 | 2 | 2 | 1 | V |
Juncus thomasii | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | V |
Cl. Scheuchzerio palustris-Caricetea fuscae, Ord. Caricetalia fuscae | ||||||||||||||||
Parnassia palustris | + | + | + | + | + | + | + | + | + | + | + | + | + | + | + | V |
Gymnadenia frivaldii | + | + | . | + | + | + | + | + | + | . | + | . | + | . | . | IV |
Carex flava | + | + | + | + | + | + | + | . | . | + | + | + | . | . | . | IV |
Pinguicula balcanica | . | + | + | + | + | . | + | + | + | + | . | + | + | + | + | IV |
Narthecium scardicum | . | . | . | . | . | . | . | . | + | + | + | . | . | . | + | II |
All. Caricion fuscae | ||||||||||||||||
Carex echinata | . | + | + | + | + | + | + | + | . | + | + | + | + | . | + | IV |
Carex nigra | + | + | . | + | + | . | + | . | . | + | . | . | . | . | . | II |
Eriophorum angustifolium | . | . | . | . | . | . | . | . | . | . | . | + | . | . | . | I |
Other species | ||||||||||||||||
Geum coccineum | 1 | 1 | 1 | 1 | 1 | 1 | 1 | + | 1 | 1 | 1 | + | 1 | + | + | V |
Athyrium filix-femina | + | + | + | + | . | + | + | . | + | + | + | + | + | + | + | V |
Epilobium montanum | + | + | + | + | + | + | + | . | + | + | + | . | + | + | + | V |
Musci sp. | + | + | + | + | + | + | + | + | . | . | + | + | . | . | . | IV |
Saxifraga aizoides | + | + | + | . | . | + | + | + | + | + | + | + | + | + | . | IV |
Lysimachia atropurpurea | + | + | + | + | . | + | + | . | + | + | + | + | . | + | . | IV |
Trifolium badium | . | . | . | + | + | . | . | + | + | + | + | + | + | + | + | IV |
Deschampsia cespitosa | + | + | . | . | . | + | + | + | + | + | + | + | . | + | + | IV |
Mentha longifolia | + | + | + | + | + | + | + | . | + | + | + | . | . | . | + | IV |
Prunella vulgaris | . | + | . | + | + | + | + | . | + | + | + | . | + | + | + | IV |
Ranunculus montanus | + | + | + | + | . | . | + | + | + | + | . | . | . | . | . | III |
Neotinea maculata | . | . | . | . | . | . | + | + | + | + | + | . | + | + | + | III |
Carex leporina | + | + | + | . | . | . | . | . | + | + | . | . | + | + | + | III |
Veratrum album | . | . | + | . | . | + | . | + | . | . | + | + | + | + | + | III |
Trifolium repens | + | + | + | + | . | + | . | + | . | . | + | + | . | . | . | III |
Stellaria alsine | + | + | . | + | + | . | + | + | . | . | . | . | . | . | + | III |
Veronica beccabunga | . | . | . | . | . | . | + | + | + | + | + | + | . | + | . | III |
Galium palustre | . | . | . | . | . | . | + | + | . | . | + | . | . | + | + | III |
Silene pusilla | . | . | . | + | + | . | . | . | + | + | . | + | + | + | + | III |
Stellaria graminea | . | . | . | + | + | . | + | . | . | + | + | + | . | . | . | II |
Cardamine carnosa | . | . | . | + | + | . | . | . | . | . | + | + | + | . | . | II |
Interpretation of the five clusters obtained is straightforward in that four of them (clusters number one, three, four and five, Figure
The syntaxonomic position of the new Association:
Class: Scheuchzerio palustris-Caricetea fuscae Tx. 1937
Order: Caricetalia fuscae Koch 1926
Alliance: Caricion fuscae Koch 1926
Association: Blysmo compressi-Eriophoretum latifoliae ass. nova
Nomenclatural note
: as explained in the Article 10 (Formation of names of associations and syntaxa of higher ranks) of the Code of Phytosociological Nomenclature (
Blysmo compressi-Eriophoretum latifoliae ass. nova
Nomenclatural type
: Table
Name giving taxa : Eriophorum latifolium and Blysmus compressus.
Diagnostic taxa : Alchemilla hybrida, Juncus conglomeratus, Epilobium montanum, Athyrium filix-femina, Saxifraga aizoides, Lysimachia atropurpurea, Trifolium badium, Caltha palustris, Ranunculus montanus, Neotinea maculata, Carex leporina, Veratrum album, Musci sp., Stellaria alsine, Veronica beccabunga, Pinguicula balcanica and Juncus thomasii.
Constant species (100–55%) : Alchemilla hybrida, Athyrium filix-femina, Blysmus compressus, Caltha palustris, Carex echinata, Deschampsia cespitosa, Epilobium montanum, Eriophorum latifolium, Geum coccineum, Juncus conglomeratus, Juncus thomasii, Lysimachia atropurpurea, Mentha longifolia, Parnassia palustris, Pinguicula balcanica, Prunella vulgaris and Saxifraga aizoides.
As for the floral elements, the Assocation: Blysmo compressi-Eriophoretum latifoliae ass. nova. - plant species of Balkan and Eur-Asiatic origin clearly predominate (Figure
The newly reported high-mountain fen vegetation association Blysmo compressi-Eriophoretum latifoliae occurred at roughly similar elevations compared to the community Blysmus compressus & Juncus thomasii and the association Caricetum macedonicae. However, compared to Pinguiculo-Narthecietum scardici, it grew at lower elevations, while at an average of 300 meters it grew higher than Eriophoro-Caricetum flavae (Figure
Of the four plant communities in the comparative analysis group, Blysmo compressi-Eriophoretum latifoliae has obviousely the greatest similarity to the Blysmus compressus & Juncus thomasii community. However, the difference between these two communities is also obvious. Almost certainly we are dealing with two distinct plant communities. Quézel (1964) reports that this is a plant community that develops along peat bogs rich in Cyperaceae species near streams. And as a particularly species-poor plant community, with fewer than 13 plant species recorded. Blysmo compressi-Eriophoretum latifoliae is also considered species-poor, but has a significantly higher number (34) of plant taxa than the aforementioned plant community. In addition to the differences mentioned, there is the geographical distance of the original record of these two plant communities, which belong to the alliance Caricion fuscae. One on the Greek island of Skasmada and the other in the Sharri Mountains in Kosovo.
Synoptic table in percentage frequency. Plant taxa are sorted according to decreasing fidelity (unstandardized phi-coefficient) to an association. Highlighted in grey are taxa with a statistically significant fidelity to a cluster (Fisher exact test < 0.05).
Associations | 1 | 2 | 3 | 4 | 5 |
---|---|---|---|---|---|
No. of relevés | 5 | 15 | 8 | 5 | 5 |
Alliance diagnostic species ( |
|||||
Plantago lanceolata | 100.0 | 0 | 0 | 0 | 0 |
Carex flacca subsp. flacca | 87.3 | 0 | 0 | 48.0 | 0 |
Anthoxanthum odoratum | 59.0 | 0 | 0 | 0 | 0 |
Parnassia palustris | 0 | 48.0 | 48.0 | 0 | 0 |
Trifolium repens | 0 | 69.1 | 0 | 0 | 0 |
Juncus articulatus | 0 | 0 | 61.8 | 0 | 0 |
Willemetia stipitata subsp. albanica | 0 | 0 | 47.6 | 0 | 53.0 |
Galium palustre | 0 | 0 | 39.7 | 0 | 0 |
Carex nigra | 0 | 0 | 0 | 0 | 45.8 |
Carex echinata | 0 | 0 | 0 | 49.1 | 0 |
Diagnostic species of individual associations | |||||
1. Blysmus compressus & Juncus thomasii Quézel 1964 | |||||
Leontodon hispidus | 61.6 | 0 | 27.4 | 0 | 0 |
Poa nemoralis | 59.0 | 0 | 0 | 0 | 0 |
Epilobium parviflorum | 59.0 | 0 | 0 | 0 | 0 |
Taraxacum laevigatum | 59.0 | 0 | 0 | 0 | 0 |
Carex appropinquata | 59.0 | 0 | 0 | 0 | 0 |
Poa trivialis | 59.0 | 0 | 0 | 0 | 0 |
2. Blysmo compressi-Eriophoretum latifoliae ass. nova | |||||
Alchemilla hybrida | 0 | 100.0 | 0 | 0 | 0 |
Juncus conglomeratus | 0 | 100.0 | 0 | 0 | 0 |
Epilobium montanum | 0 | 91.6 | 0 | 0 | 0 |
Athyrium filix-femina | 0 | 91.6 | 0 | 0 | 0 |
Saxifraga aizoides | 0 | 87.3 | 0 | 0 | 0 |
Lysimachia atropurpurea | 0 | 82.9 | 0 | 0 | 0 |
Trifolium badium | 0 | 78.4 | 0 | 0 | 0 |
Caltha palustris | 0 | 73.7 | 0 | 0 | 0 |
Ranunculus montanus | 0 | 69.1 | 0 | 0 | 0 |
Neotinea maculata | 0 | 69.1 | 0 | 0 | 0 |
Carex leporina | 0 | 69.1 | 0 | 0 | 0 |
Veratrum album | 0 | 69.1 | 0 | 0 | 0 |
Musci | 0 | 65.2 | 0 | 0 | 0 |
Stellaria alsine | 0 | 64.2 | 0 | 0 | 0 |
Veronica beccabunga | 0 | 64.2 | 0 | 0 | 0 |
Pinguicula balcanica | 0 | 61.6 | 0 | 0 | 0 |
Eriophorum latifolium | 0 | 61.2 | 61.2 | 0 | 0 |
Geum coccineum | 0 | 61.2 | 61.2 | 0 | 0 |
Juncus thomasii | 0 | 56.4 | 0 | 0 | 0 |
Cardamine carnosa | 0 | 53.5 | 0 | 0 | 0 |
Gymnadenia frivaldii | 0 | 50.3 | 0 | 0 | 0 |
Stellaria graminea | 0 | 48.1 | 0 | 0 | 0 |
Parnassia palustris | 0 | 48.0 | 48.0 | 0 | 0 |
Deschampsia cespitosa | 0 | 47.0 | 0 | 0 | 0 |
Silene pusilla | 0 | 44.5 | 0 | 0 | 0 |
Blysmus compressus | 0 | 43.9 | 0 | 0 | 0 |
Prunella vulgaris | 0 | 42.0 | 0 | 0 | 0 |
Mentha longifolia | 0 | 35.6 | 0 | 0 | 0 |
3. Caricetum macedonicae (Horv. 36) |
|||||
Carex ferruginea | 0 | 0 | 100.0 | 0 | 0 |
Trifolium hybridum | 0 | 0 | 92.1 | 0 | 0 |
Alchemilla glabra | 0 | 0 | 84.0 | 0 | 0 |
Carex lepidocarpa | 0 | 0 | 75.6 | 0 | 0 |
Equisetum palustre | 0 | 0 | 72.9 | 0 | 0 |
Juncus alpinoarticulatus | 0 | 0 | 71.1 | 0 | 0 |
Campanula abietina | 0 | 0 | 66.7 | 0 | 0 |
Epilobium palustre | 0 | 0 | 62.0 | 0 | 0 |
Luzula multiflora | 0 | 0 | 61.8 | 0 | 0 |
Cynosurus cristatus | 0 | 0 | 61.8 | 0 | 0 |
Cerastium fontanum subsp. vulgare | 0 | 0 | 56.9 | 0 | 0 |
Cardamine raphanifolia | 0 | 0 | 56.9 | 0 | 0 |
Ranunculus breyninus | 0 | 0 | 56.9 | 0 | 0 |
Sagina procumbens | 0 | 0 | 45.9 | 0 | 0 |
Carex paniculata | 0 | 0 | 45.9 | 0 | 0 |
Euphrasia minima | 0 | 0 | 45.9 | 0 | 0 |
Agrostis canina | 0 | 0 | 41.6 | 0 | 0 |
Trifolium pratense | 0 | 0 | 40.7 | 0 | 0 |
4. Eriophoro-Caricetum flavae Ranđelović & Radak 1994 | |||||
Potentilla erecta | 0 | 0 | 0 | 100.0 | 0 |
Hamatocaulis vernicosus | 0 | 0 | 0 | 100.0 | 0 |
Crepis paludosa | 0 | 0 | 0 | 100.0 | 0 |
Ranunculus acris | 0 | 0 | 0 | 87.3 | 0 |
Drepanocladus exannulatus | 0 | 0 | 0 | 87.3 | 0 |
Carex rostrata | 0 | 0 | 0 | 87.3 | 0 |
Equisetum fluviatile | 0 | 0 | 0 | 87.3 | 0 |
Briza media | 0 | 0 | 0 | 73.9 | 0 |
Geum rivale | 0 | 0 | 0 | 59.0 | 0 |
Juncus effusus | 0 | 0 | 0 | 59.0 | 0 |
Filipendula ulmaria | 0 | 0 | 0 | 59.0 | 0 |
Vicia cracca | 0 | 0 | 0 | 59.0 | 0 |
Carex flava | 0 | 0 | 0 | 54.9 | 0 |
Dactylorhiza cordigera | 0 | 0 | 33.8 | 45.9 | 0 |
5. Pinguiculo-Narthecietum scardici |
|||||
Pinguicula leptoceras | 0 | 0 | 0 | 0 | 100.0 |
Crepis aurea subsp. glabrescens | 0 | 0 | 0 | 0 | 87.3 |
Narthecium scardicum | 0 | 0 | 0 | 0 | 85.8 |
Selaginella selaginoides | 0 | 0 | 0 | 0 | 73.9 |
Nardus stricta | 0 | 0 | 27.4 | 0 | 61.6 |
Taraxacum palustre | 0 | 0 | 0 | 0 | 59.0 |
Saxifraga stellaris | 0 | 0 | 0 | 0 | 59.0 |
Soldanella alpina | 0 | 0 | 0 | 0 | 59.0 |
Boxplots showing medians and interquartile ranges of elevation (m a.s.l.) for the five associations in the data set. Explanations: 1 - Blysmus compressus & Juncus thomasii, 2 - Blysmo compressi-Eriophoretum latifoliae ass. nova, 3 - Caricetum macedonicae, 4 - Eriophoro-Caricetum flavae, 5 - Pinguiculo-Narthecietum scardici.
Boxplots showing medians (bold line) and interquartile ranges of plant taxa richness per phytosociological relevé. Explanations: 1 - Blysmus compressus & Juncus thomasii, 2 - Blysmo compressi-Eriophoretum latifoliae ass. nova, 3 - Caricetum macedonicae, 4 - Eriophoro-Caricetum flavae, 5 - Pinguiculo-Narthecietum scardici.
NMDS ordination plot of the five communities with the environmental variables as vectors plotted on the bi-dimensional space. NUTR - Nutrition availability, TEMP - Temperature, EnviFac - Environmental Factor, LIGHT - Light, MOIST - Moisture, REACT - Soil reaction, CONT - Continentality. The grouped numbers indicate plant communities1 - Blysmus compressus & Juncus thomasii, 2 - Blysmo compressi-Eriophoretum latifoliae ass. nova, 3 - Caricetum macedonicae, 4 - Eriophoro-Caricetum flavae, 5 - Pinguiculo-Narthecietum scardici.
The fen and bog vegetation in the Balkans represents the southern limit of distribution for continental Europe. In the syntaxonomic sense, this vegetation group is divided into two main classes: Oxycocco-Sphagnetea Br.-Bl. et Tx. ex Westhoff et al. 1946 and Scheuchzerio palustris-Caricetea fuscae Tx. 1937 (
The fen association Blysmo compressi-Eriophoretum latifoliae represents a rather unique plant community for Kosovo high-mountain vegetation. Based on various comparisons and analyses, this plant community proved to be different from the known plant communities of the same syntaxonomic alliance. Moreover, its floristic composition, the ecology of the habitat where it was thriving, and its general physiognomy do not resemble any of the plant communities we had in the dataset for direct comparison. If we rely on the hierarchical classification in the form of dendrogram, we can clearly see that this plant community is somewhat close only to Blysmus compressus & Juncus thomasii Quezel 1964. But again, even with this plant community, the differences are obvious. Blysmo compressi-Eriophoretum latifoliae is characterized by a greater altitudinal distribution than Blysmus compressus & Juncus thomasii. The average number of plant taxa per relevé is much higher than the former and they share only 6 plant taxa in common, out of 34 plant taxa in total. They have shown completely different preferences based on Ellenberg indicator values. For the plant community Blysmus compressus & Juncus thomasii, Quézel (1964) gives the following two plant species as characteristic ones: Blysmus compressus (L.) Link and Juncus thomasii Ten. While Blysmo compressi-Eriophoretum latifoliae ass. nova, besides Blysmus compressus (L.) Link has other plant species as characteristic ones: Eriophorum latifolium Hoppe and Juncus conglomeratus L. (Table
It is generally accepted that accurate and harmonized syntaxonomic classification of vegetation communities is an elementary requirement for proper conservation of endangered and vulnerable natural habitats (De Cáceres et al. 2015). It is well known too that the classification of fen vegetation varies due to the different classification systems used in European countries (
The new plant association Blysmo compressi-Eriophoretum latifoliae, established in the Mt. Luboten (Sharri Mts.) belongs syntaxonomically to the sedge-moss vegetation of the moderately to low calcium-rich slightly acidic fens of the Alliance: Caricion fuscae Koch 1926. Due to the particular characteristics of the natural habitats in which these communities develop, they are rare and very sensitive throughout the Balkan region. As a result of different classification approaches, disagreements about the affiliation of plant communities to corresponding alliances and, moreover, due to the lack of deeper and more comprehensive studies on this vegetation class, additional and detailed studies are urgently needed.
N.B. and R.Ć. planned and implemented the field sampling, data processing and MS writing. In a later stage, N.B., R.Ć., F.M. and V.M. conducted additional field surveys. F.M. and V.M. commented the obtained results and provided further suggestions. All authors critically revised the final version of the manuscript.
Table