Abstract
Quaking rich fens dominated by boreal semi-aquatic brown-mosses such as Scorpidium scorpioides and Calliergon trifarium are extremely rare in the Carpathians. These fens harbour endangered species persisting at few localities in the region. However, their phytosociological classification has not been sufficiently solved yet, because they lack Sphagnum species as well as calcicole species characteristic for the Caricion davallianae alliance. A recent pan-European synthesis on fen vegetation suggests that these fens belong to the Stygio-Caricion limosae alliance (boreal rich fen vegetation). The isolated occurrence of this alliance southward of the boreal zone and outside the Alps is rather exceptional and might represent a relict from an early post-glacial period. In this study, we compared phytosociological data for the Stygio-Caricion limosae alliance between Northern Europe and the Carpathians plus adjacent regions (the Bohemian Massif, the Dinaric Alps) using NMDS and cluster analysis. We found that the species composition of brown-moss quaking rich fens in Central and Southeastern Europe corresponds well with that in Northern Europe, confirming their assignment to Stygio-Caricion limosae. We further reconstructed the potential past distribution of the alliance in Czech Republic and Slovakia using available floristic and macrofossil data. Macrofossil data suggest that this vegetation type had been much more common in Central Europe and that today it persists only in ancient fens, showing the long-term stability of environmental conditions. The main causes of its present-day rarity are Middle-Holocene woodland phases in fens and recent water table decreases caused by anthropogenic deterioration of the water regime in the landscape.
This is a preview of subscription content, access via your institution.



References
Birks HJB, Walters SM (1973) The flora and vegetation of Barno jezero, Durmitor, Montenegro. Glasn Republ Zavoda Zaštitu Prir Prirodnjačke Zbirke Titogradu 5:5–23
Chytrý M, Hennekens SM, Jiménez-Alfaro B, Knollová I, Dengler J, Jansen F, Landucci F, Schaminée JHJ, Aćić S, Agrillo E, Ambarlı D, Angelini P, Apostolova I, Attorre F, Berg C, Bergmeier E, Biurrun I, Botta-Dukát Z, Brisse H, Campos JA, Carlón L, Čarni A, Casella L, Csiky J, Ćušterevska R, Dajić Stevanović Z, Danihelka J, De Bie E, de Ruffray P, De Sanctis M, Dickoré WB, Dimopoulos P, Dubyna D, Dziuba T, Ejrnæs R, Ermakov N, Ewald J, Fanelli G, Fernández-González F, FitzPatrick Ú, Font X, García-Mijangos I, Gavilán RG, Golub V, Guarino R, Haveman R, Indreica A, Işık Gürsoy D, Jandt U, Janssen JAM, Jiroušek M, Kącki Z, Kavgacı A, Kleikamp M, Kolomiychuk V, Krstivojević Ćuk M, Krstonošić D, Kuzemko A, Lenoir J, Lysenko T, Marcenò C, Martynenko V, Michalcová D, Moeslund JE, Onyshchenko V, Pedashenko H, Pérez-Haase A, Peterka T, Prokhorov V, Rašomavičius V, Rodríguez-Rojo MP, Rodwell JS, Rogova T, Ruprecht E, Rūsiņa S, Seidler G, Šibík J, Šilc U, Škvorc Ž, Sopotlieva D, Stančić Z, Svenning J-C, Swacha G, Tsiripidis I, Turtureanu PD, Uğurlu E, Uogintas D, Valachovič M, Vashenyak Y, Vassilev K, Venanzoni R, Virtanen R, Weekes L, Willner W, Wohlgemuth T, Yamalov S (2016) European Vegetation Archive (EVA): an integrated database of European vegetation plots. Appl Veg Sci 19:173–180
Chytrý M, Tichý L, Holt J, Botta-Dukát Z (2002) Determination of diagnostic species with statistical fidelity measures. J Veg Sci 13:79–90
Coldea G (1990) Munții Rodnei. Studiu geobotanic. Acad. Române, Bucureşti
Coldea G, Sanda V, Popescu A, Ştefan N (1997) Les associations végétales de Roumanie. Tome 1. Les associations herbacées naturelles. Presses Universitaires, Cluj
Dahl E (1956) Rondane. Mountain vegetation in south Norway and its relation to the environment. Skr Norske Vidensk-Akad Oslo, Mat-Naturvidensk Kl 3:1–374
Dahl E (1987) Alpine-subalpine plant communities of South Scandinavia. Phytocoenologia 15:455–484
De Cáceres M, Chytrý M, Agrillo E, Attorre F, Botta-Dukát Z, Capelo J, Czúcz B, Dengler J, Ewald J, Faber-Langendoen D, Feoli E, Franklin SB, Gavilán R, Gillet F, Jansen F, Jiménez-Alfaro B, Krestov P, Landucci F, Lengyel A, Loidi J, Mucina L, Peet RK, Roberts DW, Roleček J, Schaminée JHJ, Schmidtlein S, Theurillat J-P, Tichý L, Walker DA, Wildi O, Willner W, Wiser SK (2015) A comparative framework for broad-scale plot-based vegetation classification. Appl Veg Sci 18:543–560
Dierssen K (1996) Vegetation Nordeuropas. Verlag Eugen Ulmer, Stuttgart
Dítě D, Hájek M, Hájková P (2007) Formal definitions of Slovakian mire plant associations and their application in regional research. Biologia 62:400–408
Dítě D, Hájek M, Svitková I, Košuthová A, Šoltés R, Kliment J (2018) Glacial-relict symptoms in the Western Carpathian flora. Folia Geobot (this issue)
Dítě D, Kubandová M (2005) Blatnica močiarna (Scheuchzeria palustris L.) na Kubínskej holi (severné Slovensko). Bull Slov Bot Spoločn 27:45–48
Dítě D, Peterka T, Dítětová Z, Hájková P, Hájek M (2017) Arcto-alpine species at their niche margin: the Western Carpathian refugia of Juncus castaneus and J triglumis in Slovakia. Ann Bot Fennici 54:67–82
Dítě D, Pukajová D (2002a) Carex limosa L., kriticky ohrozený druh flóry Slovenska. Bull Slov Bot Spoločn 24:65–73
Dítě D, Pukajová D (2002b) Chorologické, ekologické a fytocenologické poznámky k výskytu ostrice výbežkatej (Carex chordorrhiza Ehrh.) na Slovensku. Bull Slov Bot Spoločn 24:61–64
Dítě D, Šoltés R (2010) Rozšírenie druhu Scorpidium scorpioides na Slovensku: minulosť a súčasnosť. Bryonora 46:66–69
Erzberger P, Papp B (2007) New and noteworthy bryophyte records from Montenegro and Serbia. Willdenowia 37:339–351
Fejfar O, Kneblová V, Dohnal Z, Ložek V (1955) Paleontologický a cenologický výzkum kvartérních sedimentů Ostravska v roce 1953. Anthropozoikum 4:241–284
Frahm JP (2012) The phytogeography of European bryophytes. Botanica Serbica 36:23–36
Frey W, Frahm JP, Fischer E, Lobin W (2006) The liverworts, mosses and ferns of Europe. Harley Books, Colchester
Gálová A, Hájková P, Čierniková M, Petr L, Hájek M, Novák J, Rohovec J, Jamrichová E (2016) Origin of a boreal birch bog woodland and landscape development on a warm low mountain summit at the Carpathian-Pannonian interface. The Holocene 26:1112–1125
Granath G, Strengbom J, Rydin H (2010) Rapid ecosystem shifts in peatlands: linking plant physiology and succession. Ecology 91:3047–3056
Hájek M, Dudová L, Hájková P, Roleček J, Moutelíková J, Jamrichová E, Horsák M (2016) Contrasting Holocene environmental histories may explain patterns of species richness and rarity in a Central European landscape. Quatern Sci Rev 133:48–61
Hájek M, Háberová I (2001) Scheuchzerio-Caricetea fuscae R.Tx. 1937. In Valachovič M (ed) Rastlinné spoločenstvá Slovenska. 3 Vegetácia mokradí. Veda, Bratislava, pp 185–274
Hájek M, Horsák M, Hájková P, Dítě D (2006) Habitat diversity of central European fens in relation to environmental gradients and an effort to standardise fen terminology in ecological studies. Perspect Pl Ecol Evol Syst 8:97–114
Hájek M, Jiroušek M, Navrátilová J, Horodyská E, Peterka T, Plesková Z, Navrátil J, Hájková P, Hájek T (2015) Changes in the moss layer in Czech fens indicate early succession triggered by nutrient enrichment. Preslia 87:279–301
Hájková P, Horsák M, Hájek M, Jankovská V, Jamrichová E, Moutelíková J (2015a) Using multi-proxy palaeoecology to test a relict status of refugial populations of calcareous-fen species in the Western Carpathians. The Holocene 25:702–715
Hájková P, Petr L, Horsák M, Rohovec J, Hájek M (2015b) Interstadial inland dune slacks in south-west Slovakia: a multi-proxy vegetation and landscape reconstruction. Quatern Internat 357:314–328
Hájková P, Potůčková A, Hájek M (2017) Zajímavé nálezy vzácných mechorostů ve starých sedimentech 2. Bryonora 59:67–71
Hájková P, Žáčková P, Dudová L, Hájek M (2012) Zajímavé nálezy vzácných mechorostů ve starých sedimentech. Bryonora 50:14–16
Jamrichová E, Petr L, Jiménez- Alfaro B, Jankovská V, Dudová L, Pokorný P, Kołaczek P, Zernitskaya V, Čierniková M, Břízová E, Syrovátka V, Hájková P, Hájek M (2017) Pollen-inferred millennial changes in landscape patterns at a major biogeographical interface within Europe. J Biogeogr 44:2386–2397
Jankovská V (1970) Ergebnisse der Pollen- and Grossrestanalyse des Moors “Velanská cesta” in Südböhmen. Folia Geobot Phytotax 5:43–60
Jankovská V (1980) Paläogeobotanische Rekonstruktion der Vegetationsentwicklung im Becken Třeboňská pánev während des Spätglazials und Holozäns. Academia, Praha
Janovicová K (1998) Zaujímavé bryofloristické nálezy zo Slovenska. Bryonora 21:2–3
Janssen JA, Rodwell JS, García Criado M, Gubbay S, Haynes T, Nieto A, Sanders N, Landucci F, Loidi J, Ssymank A, Tahvanainen T, Valderrabano M, Acosta A, Aronsson M, Arts G, Attorre F, Bergmeier E, Bijlsma RJ, Bioret F, Biţă-Nicolae C, Biurrun I, Calix M, Capelo J, Čarni A, Chytrý M, Dengler J, Dimopoulos P, Essl F, Gardfjell H, Gigante D, Giusso del Galdo G, Hájek M, Jansen F, Jansen J, Kapfer J, Mickolajczak A, Molina JA, Molnár Z, Paternoster D, Piernik A, Poulin B, Renaux B, Schaminée JHJ, Šumberová K, Toivonen H, Tonteri T, Tsiripidis I, Tzonev R, Valachovič M (2016) European Red List of Habitats. Part 2. Terrestrial and freshwater habitats. Publications Office of the European Union, Luxembourg
Janssens JA (1983) A quantitative method for stratigraphic analysis of bryophytes in Holocene peat. J Ecol 71:189–196
Joosten H, Tanneberger F, Moen A (eds) (2017) Mires and peatlands of Europe. Status, distribution and conservation. Schweizerbart Science Publishers, Stuttgart
Juutinen S (2011) The decrease of rich fen bryophytes in springs as a consequence of large-scale environmental loss A 50-year re-sampling study Lindbergia 34:2–8
Kaplan Z, Danihelka J, Štěpánková J, Ekrt L, Chrtek J Jr, Zázvorka J, Grulich V, Řepka R, Prančl J, Ducháček M, Kúr P, Šumberová K, Brůna J (2016) Distributions of vascular plants in the Czech Republic. Part 2. Preslia 88:229–322
Kooijman AM (2012) ‘Poor rich fen mosses’: atmospheric N-deposition and P-eutrophication in base-rich fens. Lindbergia 35:42–52
Kooijman AM, Beltman B, Westhoff V (1994) Extinction and reintroduction of the bryophyte Scorpidium scorpioides in a rich-fen site in the Netherlands. Biol Conservation 69:87–96
Kuneš P, Odgaard Vad B, Gaillard MJ (2011) Soil phosphorus as a control of productivity and openness in temperate interglacial forest ecosystems. J Biogeogr 38:2150–2164
Magyari E, Jakab G, Rudner E, Sümegi P (1999) Palynological and plant macrofossil data on Late Pleistocene short term climatic osscilations in North-east Hungary. Acta Palaeobot 2:491–502
Malmer N (1986) Vegetation gradients in relation to environmental conditions in northwestern European mires. Canad J Bot 64:375–383
Mälson K, Rydin H (2007) The regeneration capabilities of bryophytes for rich fen restoration. Biol Conservation 135:435–442
Migra V, Mičeta K (1997) K výskytu Vignea chordorrhiza (Ehrh. ex L. fil.) Rchb. na Slovensku. Bull Slov Bot Spoločn 19:29–31
Migra V, Šoltés R (1998) Glaciálny relikt Meesia triquetra (Bryophyta) na Kubínskej holi (Oravská Magura). Bull Slov Bot Spoločn 20:48–50
Moen A, Lyngstad A, Øien D (2012) Boreal rich fen vegetation formerly used for haymaking. Nordic J Bot 30:226–240
Mráz P, Ronikier M (2016). Biogeography of the Carpathians: evolutionary and spatial facets of biodiversity. Biol J Linn Soc 119:528–559
Mucina L, Bültmann H, Dierssen K, Theurillat JP, Raus T, Čarni A, Šumberová K, Willner W, Dengler J, García RG, Chytrý M, Hájek M, Di Pietro R, Iakushenko D, Pallas J, Daniëls FJA, Bergmeier E, Guerra AS, Ermakov N, Valachovič M, Schaminée JHJ, Lysenko T, Didukh YP, Pignatti S, Rodwell JS, Capelo j, Weber HE, Solomeshch A, Dimopoulos P, Aguiar C, Hennekens SM, Tichý L (2016) Vegetation of Europe: hierarchical floristic classification of vascular plant, bryophyte, lichen, and algal communities. Appl Veg Sci 19 (suppl. 1):3–264
Navrátilová J, Hájek M, Navrátil J, Hájková P, Frazier RJ (2017) Convergence and impoverishment of fen communities in a eutrophicated agricultural landscape of the Czech Republic. Appl Veg Sci 20:225–235
Nordhagen R (1943) Sikilsdalen og Norges fjellbeiter. En plantesosiologisk monografi. Bergens Mus Skr 22:1–607
Novák FA (1954) Přehled Československé květeny s hlediska ochrany přírody a krajiny. In Veselý J (ed) Ochrana Československé přírody a krajiny. Díl II. Nakladatelství ČSAV, Praha, pp 193–409
Paulissen MPCP, Schaminée JHJ, During HJ, Wieger Wamelink GW, Verhoeven JTA (2014) Expansion of acidophytic late successional bryophytes in Dutch fens between 1940 and 2000. J Veg Sci 25:525–533
Pawlikowski P, Abramczyk K, Szczepaniuk A, Kozub Ł (2013) Nitrogen: phosphorus ratio as the main ecological determinant of the differences in the species composition of brown-moss rich fens in north-eastern Poland. Preslia 85:349–367
Pearson RG (1965) Problems of post-Glacial refugia. Proc R Soc London 161:324–330
Peterka T, Hájek M, Jiroušek M, Jiménez-Alfaro B, Aunina L, Bergamini A, Dítě D, Felbaba-Klushyna L, Graf U, Hájková P, Hettenbergerová E, Ivchenko TG, Jansen F, Koroleva NE, Lapshina ED, Lazarević PM, Moen A, Napreenko MG, Pawlikowski P, Plesková Z, Sekulová L, Smagin VA, Tahvanainen T, Thiele A, Biţa-Nicolae C, Biurrun I, Brisse H, Ćušterevska R, De Bie E, Ewald J, FitzPatrick Ú, Font X, Jandt U, Kącki Z, Kuzemko A, Landucci F, Moeslund JE, Pérez-Haase A, Rašomavičius V, Rodwell JS, Schaminée JHJ, Šilc U, Stančić Z, Chytrý M (2017) Formalized classification of European fen vegetation at the alliance level. Appl Veg Sci 20:124–142
Pokorný P, Chytrý M, Juřičková L, Sádlo J, Novák J, Ložek V (2015) Mid-Holocene bottleneck for central European dry grasslands: Did steppe survive the forest optimum in northern Bohemia, Czech Republic? The Holocene 25:716–726
Rehell S, Virtanen R (2016) Rich-fen bryophytes in past and recent mire vegetation in a successional land uplift area. The Holocene 26:136–146
Řepka R, Grulich V (2014) Ostřice České republiky. Lesnická Práce, Praha
Růžička I (1987) Výsledky záchranného výzkumu ohrožené květeny mizejících rašelinišť a rašelinných luk v okolí Telče. Vlastiv Sborn Vysoč, Odd Věd Přír 8:153–192
Rybníček K (1966) Glacial relics in the bryoflora of the highlands Českomoravská vrchovina (Bohemian-Moravian Highlands); their habitat and cenotaxonomic value. Folia Geobot Phytotax 1:101–119
Rybníček K, Balátová-Tuláčková E, Neuhäusl R (1984) Přehled rostlinných společenstev rašelinišť a mokřadních luk Československa. Stud Českoslov Akad Ved 1984:1–124
Rybníček K, Rybníčková E (1965) Přechodové rašeliniště u Beňadova na Oravě. Biológia 20:373–375
Rybníček K, Rybníčková E (1968) The history of flora and vegetation on the Bláto mire in southeastern Bohemia, Czechoslovakia (palaeoecological study). Folia Geobot Phytotax 3:117–142
Rybníček K, Rybníčková E (1972). Nálezy vzácných rašelinných mechorostů na Oravě. Biológia, 27:795–798
Rybníčková E, Rybníček K (1972) Erste Ergebnisse paläogeobotanischer Untersuchungen des Moores bei Vracov, Südmähren. Folia Geobot Phytotax 7:285–308
Singsaas S (1989) Classification and ordination of the mire vegetation of Stormyra near Tynset, S Norway. Nordic J Bot 9:413–423
Sjörs H (1952) On the relation between vegetation and electrolytes in north Swedish mire waters. Oikos 2:241–258
Sjörs H, Björkbäck F, Nordqvist Y (1965) Northern mires. Acta Phytogeogr Suec 50:180–197
Šmilauer P, Lepš JŠ (2014) Multivariate analysis of ecological data using CANOCO 5. Cambridge University Press, Cambridge
Šoltés R, Dítě D, Mihálik D, Ondreičková K, Hrehová Z, Maximová N, Sedláková B (2015) Seasonal variation in bryophytes cover in the calcareous mire Belianské lúky, Slovakia. Pakistan J Bot 47:255–262
Štechová T, Manukjanová A, Holá E, Kubešová S, Novotný I, Zmrhalová M (2010) Současný stav populací druhů Helodium blandowii (Thuidiaceae) a Scorpidium scorpioides (Calliergonaceae) v České republice. Bryonora 46:24–33
Tichý L (2002) JUICE, software for vegetation classification. J Veg Sci 13:451–453
Tutin TG, Heywood VH, Burges NA, Moore DM, Valentine DH, Walters SM, Webb DA (eds) (1968–1993) Flora Europaea, vols 2–5, 2nd edn. of Vol. 1. Cambridge University Press, Cambridge
Udd D, Mälson K, Sundberg S, Rydin H (2015) Explaining species distributions by traits of bryophytes and vascular plants in a patchy landscape. Folia Geobot 50:161–174
Udd D, Sundberg S, Rydin H (2016) Multi-species competition experiments with peatland bryophytes. J Veg Sci 27:165–175
Vicherová E, Hájek M, Hájek T (2015) Calcium intolerance of fen mosses: physiological evidence, effects of nutrient availability and successional drivers. Perspect Pl Ecol Evol Syst 17:347–359
Vicherová E, Hájek M, Šmilauer P, Hájek T (2017) Sphagnum establishment in alkaline fens: importance of weather and water chemistry. Sci Total Environm 580:1429–1438
Acknowledgements
The research was funded by the Czech Science Foundation (Centre of Excellence Pladias; 14-36079G) and Masaryk University (MUNI/A/1301/2016). The research of PH was partially supported by Czech Academy of Sciences (RVO 67985939). We are grateful to Kamil Rybníček (†) for providing unpublished vegetation plots. Our unpublished data from Romania were collected with the help of Veronika Horsáková and Michal Horsák. Many thanks go to Eva Hettenbergerová for management of the Macrofossil Database of the Czech Republic and Slovakia and for general support during the preparation of the manuscript. The co-ordinating editor and two anonymous reviewers provided important comments on the first draft of the paper.
Author information
Authors and Affiliations
Corresponding author
Electronic supplementary material
ESM 1
(DOC 48854 kb)
Rights and permissions
About this article
Cite this article
Peterka, T., Hájek, M., Dítě, D. et al. Relict occurrences of boreal brown-moss quaking rich fens in the Carpathians and adjacent territories. Folia Geobot 53, 265–276 (2018). https://doi.org/10.1007/s12224-018-9318-3
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s12224-018-9318-3
Keywords
- Classification
- Macrofossil
- Mires
- Plant communities
- Vegetation
- Wetlands