Syntaxonomical classification of wet woodlands with Picea abies in Slovakia
Wet woodlands with domination of Norway spruce are floristically and ecologically distinctive element of coniferous forest vegetation. However, specialized studies on this vegetation are considerably rare. In this survey the syntaxonomical classification of 145 relevés of Sphagnum-rich and other w...
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irk-123456789-1767972021-02-09T01:26:08Z Syntaxonomical classification of wet woodlands with Picea abies in Slovakia Kucera, P. Геоботаніка, екологія, охорона рослинного світу Wet woodlands with domination of Norway spruce are floristically and ecologically distinctive element of coniferous forest vegetation. However, specialized studies on this vegetation are considerably rare. In this survey the syntaxonomical classification of 145 relevés of Sphagnum-rich and other wet woodlands with Picea abies from Slovakia is proposed. Eight plant communities in the rank of association were differentiated. A distinctive feature of the association Sphagno acutifolii-Piceetum Zukrigl 1973 is the co-occurrence of species typical for climax supramontane woodlands on silicate bedrock of the alliance Piceion excelsae Pawłowski ex Pawłowski et al. 1928 and species characteristic for woodlands of the class Vaccinio uliginosi-Pinetea Passarge 1968. Consequently, this association is classified within the order Piceetalia abietis Pawłowski ex Pawłowski et al. 1928 and alliance Piceion excelsae Pawłowski ex Pawłowski et al. 1928. The rest of wet Picea woodland associations are azonal stable communities (Dauergesellschaften) distributed mostly in the montane zone and they differ floristically as well. Therefore those associations are separated in the floristically well distiguishable order Sphagno palustris-Piceetalia P. Kučera 2019 and they are subdivided into three syntaxa in the rank of alliance: (1) alliance Sphagno palustris-Piceion P. Kučera 2019 comprising the communities limited to nutrient-poor habitats with a shallow peat layer or hydromorphic soils with a substantial raw humus layer: Soldanello montanae- Piceetum Volk in Br.-Bl. et al. 1939, Leucobryo glauci-Piceetum abietis Šomšák ex P. Kučera 2019, Sphagno palustris-Piceetum Šomšák 1979, Equiseto sylvatici-Piceetum Šmarda 1950, and the Carex rostrata-Picea abies community; (2) alliance Stellario nemorum-Abietion albae P. Kučera 2019 with the species-rich spring related association Stellario nemorum-Abietetum albae P. Kučera 2019; (3) base rich alliance Valeriano dioicae-Abietion albae P. Kučera 2019 with the association Valeriano dioicae-Abietetum P. Kučera 2019. Formal definitions and description of species composition of the units are given, as well as brief information on their distribution in Slovakia. Several syntaxonomical and nomenclatural notes are provided in the supplement. Вологі ліси з домінуванням ялини звичайної (Picea abies) є флористично та екологічно відмінним елементом рослинності хвойних лісів. Однак спеціальні дослідження цієї рослинності є доволі рідкісними. У нашому дослідженні проведено синтаксономічну класифікацію 145 геоботанічних описів сфагнових та інших вологих лісових масивів із участю Picea abies зі Словаччини. Відмічені вісім рослинних угруповань у ранзі асоціації. Відмінною рисою асоціації Sphagno acutifolii-Piceetum Zukrigl 1973 є трапляння як видів, характерних для клімаксових супрамонтанних лісових масивів на силікатних породах з союзу Piceion excelsae Pawłowski ex Pawłowski et al. 1928, так і видів, характерних для лісових масивів із класу Vaccinio uliginosi-Pinetea Passarge 1968. Отже, ця асоціація належить до порядку Piceetalia abietis Pawłowski ex Pawłowski et al. 1928 та союзу Piceion excelsae Pawłowski ex Pawłowski et al. 1928. Решта асоціацій вологих лісів із участю ялини – це азональні стійкі угруповання (Dauergesellschaften), які поширені переважно в монтанній зоні і є флористично відмінними. Тому ці асоціації відокремлюються у флористично чітко відокремленому порядку Sphagno palustris-Piceetalia P. Kučera 2019 і поділяються на три синтаксони у ранзі союзу: (1) союзу Sphagno palustris- Piceion P. Kučera 2019, що включає угруповання, пов'язані з бідними на поживні речовини оселищами з поверхневим торфовим шаром або гідроморфними ґрунтами із значним шаром гумусу: Soldanello montanae-Piceetum Volk in Br.-Bl. et al. 1939, Leucobryo glauci-Piceetum abietis Šomšák ex P. Kučera 2019, Sphagno palustris-Piceetum Šomšák 1979, Equiseto sylvatici-Piceetum Šmarda 1950 та угруповання Carex rostrata-Picea abies; (2) союзу Stellario nemorum-Abietion albae P. Kučera 2019 з асоціацією Stellario nemorum-Abietetum albae P. Kučera 2019 з різноманітними весняними видами; (3) союзу Valeriano dioicae-Abietion albae P. Kučera 2019 на багатих основами ґрунтах з асоціацією Valeriano dioicae- Abietetum P. Kučera 2019. Подано формальні визначення та описи видового складу цих одиниць, а також стислу інформацію про їхнє поширення у Словаччині. Деякі синтаксономічні та номенклатурні примітки наведено в додатку. 2019 Article Syntaxonomical classification of wet woodlands with Picea abies in Slovakia / P. Kucera // Український ботанічний журнал. — 2019. — Т. 76, № 4. — С. 316-343. — Бібліогр.: 53 назв. — англ. 0372-4123 DOI: https://doi.org/10.15407/ukrbotj76.04.316 http://dspace.nbuv.gov.ua/handle/123456789/176797 en Український ботанічний журнал Інститут ботаніки ім. М.Г. Холодного НАН України |
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| topic |
Геоботаніка, екологія, охорона рослинного світу Геоботаніка, екологія, охорона рослинного світу |
| spellingShingle |
Геоботаніка, екологія, охорона рослинного світу Геоботаніка, екологія, охорона рослинного світу Kucera, P. Syntaxonomical classification of wet woodlands with Picea abies in Slovakia Український ботанічний журнал |
| description |
Wet woodlands with domination of Norway spruce are floristically and ecologically distinctive element of coniferous forest
vegetation. However, specialized studies on this vegetation are considerably rare. In this survey the syntaxonomical classification
of 145 relevés of Sphagnum-rich and other wet woodlands with Picea abies from Slovakia is proposed. Eight plant communities
in the rank of association were differentiated. A distinctive feature of the association Sphagno acutifolii-Piceetum Zukrigl 1973
is the co-occurrence of species typical for climax supramontane woodlands on silicate bedrock of the alliance Piceion excelsae
Pawłowski ex Pawłowski et al. 1928 and species characteristic for woodlands of the class Vaccinio uliginosi-Pinetea Passarge 1968.
Consequently, this association is classified within the order Piceetalia abietis Pawłowski ex Pawłowski et al. 1928 and alliance
Piceion excelsae Pawłowski ex Pawłowski et al. 1928. The rest of wet Picea woodland associations are azonal stable communities
(Dauergesellschaften) distributed mostly in the montane zone and they differ floristically as well. Therefore those associations
are separated in the floristically well distiguishable order Sphagno palustris-Piceetalia P. Kučera 2019 and they are subdivided into
three syntaxa in the rank of alliance: (1) alliance Sphagno palustris-Piceion P. Kučera 2019 comprising the communities limited to
nutrient-poor habitats with a shallow peat layer or hydromorphic soils with a substantial raw humus layer: Soldanello montanae-
Piceetum Volk in Br.-Bl. et al. 1939, Leucobryo glauci-Piceetum abietis Šomšák ex P. Kučera 2019, Sphagno palustris-Piceetum
Šomšák 1979, Equiseto sylvatici-Piceetum Šmarda 1950, and the Carex rostrata-Picea abies community; (2) alliance Stellario
nemorum-Abietion albae P. Kučera 2019 with the species-rich spring related association Stellario nemorum-Abietetum albae P.
Kučera 2019; (3) base rich alliance Valeriano dioicae-Abietion albae P. Kučera 2019 with the association Valeriano dioicae-Abietetum
P. Kučera 2019. Formal definitions and description of species composition of the units are given, as well as brief information on
their distribution in Slovakia. Several syntaxonomical and nomenclatural notes are provided in the supplement. |
| format |
Article |
| author |
Kucera, P. |
| author_facet |
Kucera, P. |
| author_sort |
Kucera, P. |
| title |
Syntaxonomical classification of wet woodlands with Picea abies in Slovakia |
| title_short |
Syntaxonomical classification of wet woodlands with Picea abies in Slovakia |
| title_full |
Syntaxonomical classification of wet woodlands with Picea abies in Slovakia |
| title_fullStr |
Syntaxonomical classification of wet woodlands with Picea abies in Slovakia |
| title_full_unstemmed |
Syntaxonomical classification of wet woodlands with Picea abies in Slovakia |
| title_sort |
syntaxonomical classification of wet woodlands with picea abies in slovakia |
| publisher |
Інститут ботаніки ім. М.Г. Холодного НАН України |
| publishDate |
2019 |
| topic_facet |
Геоботаніка, екологія, охорона рослинного світу |
| url |
http://dspace.nbuv.gov.ua/handle/123456789/176797 |
| citation_txt |
Syntaxonomical classification of wet woodlands with Picea abies in Slovakia / P. Kucera // Український ботанічний журнал. — 2019. — Т. 76, № 4. — С. 316-343. — Бібліогр.: 53 назв. — англ. |
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| fulltext |
316 Ukrainian Botanical Journal, 2019, 76(4)
Геоботаніка, екологія, охорона рослинного світу
Vegetation Science, Ecology, Conserva tion
https://doi.org/10.15407/ukrbotj76.04.316
Syntaxonomical classification of wet woodlands with Picea abies in Slovakia
Peter KUČERA
Comenius University in Bratislava, Botanical Garden, workplace Blatnica
Blatnica 315, SK-038 15 Blatnica pri Martine, Slovakia
peter.kucera@uniba.sk
Kučera P. 2019. Syntaxonomical classification of wet woodlands with Picea abies in Slovakia. Ukrainian Botanical Journal, 76(4):
316–343.
Abstract. Wet woodlands with domination of Norway spruce are floristically and ecologically distinctive element of coniferous forest
vegetation. However, specialized studies on this vegetation are considerably rare. In this survey the syntaxonomical classification
of 145 relevés of Sphagnum-rich and other wet woodlands with Picea abies from Slovakia is proposed. Eight plant communities
in the rank of association were differentiated. A distinctive feature of the association Sphagno acutifolii-Piceetum Zukrigl 1973
is the co-occurrence of species typical for climax supramontane woodlands on silicate bedrock of the alliance Piceion excelsae
Pawłowski ex Pawłowski et al. 1928 and species characteristic for woodlands of the class Vaccinio uliginosi-Pinetea Passarge 1968.
Consequently, this association is classified within the order Piceetalia abietis Pawłowski ex Pawłowski et al. 1928 and alliance
Piceion excelsae Pawłowski ex Pawłowski et al. 1928. The rest of wet Picea woodland associations are azonal stable communities
(Dauergesellschaften) distributed mostly in the montane zone and they differ floristically as well. Therefore those associations
are separated in the floristically well distiguishable order Sphagno palustris-Piceetalia P. Kučera 2019 and they are subdivided into
three syntaxa in the rank of alliance: (1) alliance Sphagno palustris-Piceion P. Kučera 2019 comprising the communities limited to
nutrient-poor habitats with a shallow peat layer or hydromorphic soils with a substantial raw humus layer: Soldanello montanae-
Piceetum Volk in Br.-Bl. et al. 1939, Leucobryo glauci-Piceetum abietis Šomšák ex P. Kučera 2019, Sphagno palustris-Piceetum
Šomšák 1979, Equiseto sylvatici-Piceetum Šmarda 1950, and the Carex rostrata-Picea abies community; (2) alliance Stellario
nemorum-Abietion albae P. Kučera 2019 with the species-rich spring related association Stellario nemorum-Abietetum albae P.
Kučera 2019; (3) base rich alliance Valeriano dioicae-Abietion albae P. Kučera 2019 with the association Valeriano dioicae-Abietetum
P. Kučera 2019. Formal definitions and description of species composition of the units are given, as well as brief information on
their distribution in Slovakia. Several syntaxonomical and nomenclatural notes are provided in the supplement.
Keywords: Abies alba, nomenclature, phytocoenology, Picea abies, plant communities, Western Carpathians
Supplementary Material. Electronic Supplements: A1, Figure 1, A2, А3; B1(Table 1); B2(Table 4), pp. e1—e19, are available in
the online version of this article at: https://ukrbotj.co.ua/76/4/316
Submitted 04 February 2019. Published 02 September 2019
Кучера П. 2019. Синтаксономічна класифікація вологих лісових масивів із участю ялини (Picea abies) у Словаччині.
Український ботанічний журнал, 76(4): 316–343.
Університет ім. Коменського в Братиславі, Ботанічний сад
Блатніца 315, SK-038 15 округа Мартін, Словаччина
Реферат. Вологі ліси з домінуванням ялини звичайної (Picea abies) є флористично та екологічно відмінним елементом
рослинності хвойних лісів. Однак спеціальні дослідження цієї рослинності є доволі рідкісними. У нашому дослідженні
проведено синтаксономічну класифікацію 145 геоботанічних описів сфагнових та інших вологих лісових масивів із
участю Picea abies зі Словаччини. Відмічені вісім рослинних угруповань у ранзі асоціації. Відмінною рисою асоціації
Sphagno acutifolii-Piceetum Zukrigl 1973 є трапляння як видів, характерних для клімаксових супрамонтанних лісових
масивів на силікатних породах з союзу Piceion excelsae Pawłowski ex Pawłowski et al. 1928, так і видів, характерних для
лісових масивів із класу Vaccinio uliginosi-Pinetea Passarge 1968. Отже, ця асоціація належить до порядку Piceetalia abietis
Pawłowski ex Pawłowski et al. 1928 та союзу Piceion excelsae Pawłowski ex Pawłowski et al. 1928. Решта асоціацій вологих
лісів із участю ялини – це азональні стійкі угруповання (Dauergesellschaften), які поширені переважно в монтанній
зоні і є флористично відмінними. Тому ці асоціації відокремлюються у флористично чітко відокремленому порядку
Sphagno palustris-Piceetalia P. Kučera 2019 і поділяються на три синтаксони у ранзі союзу: (1) союзу Sphagno palustris-
Piceion P. Kučera 2019, що включає угруповання, пов'язані з бідними на поживні речовини оселищами з поверхневим
торфовим шаром або гідроморфними ґрунтами із значним шаром гумусу: Soldanello montanae-Piceetum Volk in Br.-Bl.
et al. 1939, Leucobryo glauci-Piceetum abietis Šomšák ex P. Kučera 2019, Sphagno palustris-Piceetum Šomšák 1979, Equiseto
sylvatici-Piceetum Šmarda 1950 та угруповання Carex rostrata-Picea abies; (2) союзу Stellario nemorum-Abietion albae
P. Kučera 2019 з асоціацією Stellario nemorum-Abietetum albae P. Kučera 2019 з різноманітними весняними видами;
(3) союзу Valeriano dioicae-Abietion albae P. Kučera 2019 на багатих основами ґрунтах з асоціацією Valeriano dioicae-
Abietetum P. Kučera 2019. Подано формальні визначення та описи видового складу цих одиниць, а також стислу
інформацію про їхнє поширення у Словаччині. Деякі синтаксономічні та номенклатурні примітки наведено в додатку.
Ключові слова: Західні Карпати, номенклатура, рослинні угруповання, фітоценологія, Abies alba, Picea abies
© 2019 P. Kučera. Published by the M.G. Kholodny Institute of Botany, NAS of Ukraine. This is an open access article under the terms of the
Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0/), which permits use, distribution, and reproduction in
any medium, provided the original work is properly cited
mailto:peter.kucera@uniba.sk
317Український ботанічний журнал, 2019, 76(4)
Introduction
Woodlands with the natural dominance of Picea abies are
distributed in the Western Carpathians predominantly
in the mountains where they form a distinctive belt,
the supramontane altitudinal vegetation zone, mostly
above 1400 m a.s.l. (Domin, 1923; Kučera, 2012). In
syntaxonomy, plant communities of this mountain
forest are classified as the class Piceetea excelsae Klika
19481 in the phytocoenological literature (Hadač et al.,
1969; Šoltés, 1976; Fajmonová, 1978; Matuszkiewicz,
2002; Kučera, 2012).
Apart from the supramontane zone of the Western
Carpathians, Picea abies naturally dominates (or
codominates with Abies alba) on locally distributed
special habitats, especially on more or less ground-
water and/or above-ground-water influenced areas
(Šomšák, 1979, 1983; Šomšák et al., 1993, 1996;
Bujakiewicz, 1981; Majzlanová, 1983; Staszkiewicz,
1993; Kasprowicz, 1996; Parusel, 2007; Wilczek et al.,
2015; and others) in the lower montane elevations of the
Fagus sylvatica-Abies alba forest zone (class Carpino-
Fagetea Jakucs ex Passarge 1968).
Depending on the origin of the habitat, level of the
ground water and time of its influence (or sometimes
overflooding), several types of wet Picea woodlands are
recognized in Slovakia and adjacent countries.
In Austria (Exner, 2007) they are further splitted
between two alliances (order Piceetalia Pawłowski
ex Pawłowski et al. 1928) following the traditional
approach of German-speaking phytocoenologists: (1)
Abieti-Piceion (Br.-Bl. in Br.-Bl. et al. 1939) Soó 19632
with associations Carici brizoidis-Abietetum Trinajstič
1974, Equiseto-Abietetum Moor ex Kuoch 1954; (2)
Piceion excelsae Pawłowski ex Pawłowski et al. 19283
with Equiseto-Piceetum Šmarda 1950, Sphagno-
Piceetum Zukrigl 1973.
In the recent vegetation survey of the Czech Republic
(Chytrý et al., 2013) wet Picea woodlands are classified
and divided into alliances differently (without indication
of the rank of order): (1) Piceion excelsae Pawłowski ex
Pawłowski et al. 1928 containing associations Equiseto-
Piceetum Šmarda 1950, Soldanello montanae-Piceetum
Volk in Br-Bl. et al. 1939; (2) Vaccinio uliginosi-Pinion
sylvestris Passarge 1968 where bog woodland of Vaccinio
1 Syn. Vaccinio-Piceetea Br.-Bl. in Br.-Bl. et al. 1939 pro parte min.
[cf. Braun-Blanquet et al., 1939], nom. inval., Art. 2b, 3m, 25; see
Kučera (2010, 2012), Kučera, Kliment (2011), cf. Theurillat in
Willner et al. (2015).
2 Cf. Kučera (2008a), p. 168.
3 Cf. Kučera, Kliment (2011), p. 88.
uliginosi-Piceetum Schubert 1972 is one of the four
distiguished associations (minority of floristically almost
identical bog woodlands are by a decision delimited
to non-forest vegetation of the alliance Sphagnion
magellanici Kästner et Flössner 1933; Hájková et al.
2011) .
The natural distribution range of Picea abies in Poland
is divided to (1) the outskirts of the hemiboreal forest
zone in northeastern Poland and (2) larger disjunctive
Central European areal associated with Sudetian-
Carpathian mountain ranges (see Szafer, 1959). All
known wet Picea woodlands are classified within
Piceion excelsae Pawłowski ex Pawłowski et al.
1928 (Matuszkiewicz, 2002). Associations Querco-
Piceetum W. Matuszkiewicz et Polakowska 1955
and Sphagno girgensohnii-Piceetum Polakowski
1962 are described from the boreal Picea range. The
association Betulo pubescentis-Piceetum Sokołowski
1980 described by Sokołowski (1980) also from
this region is not recognized in the surveys of
J. Matuszkiewicz (2002) and W. Matuszkiewicz (2014).
Occurrence of the association Bazzanio-Piceetum
(Schmid et Gaisberg 1936) Br.-Bl. et Sissingh in Br-Bl.
et al. 19394 is reported from the mountains of southern
Poland; stands of the association Calamagrostio villosae-
Pinetum Staszkiewicz 1958 known from the Orava-
Nowy Targ Basin north of the Tatras have a special
position in respect of the presence of Picea abies.
Wet Picea woodlands of Ukraine are similarly known
from two parts of the natural Picea abies distribution
areas: (1) the southern limit of the hemiboreal Picea
abies distribution range (Polissya zone) where three
associations are distiguished: Querco-Piceetum
W. Matuszkiewicz et Polakowska 1955, Sphagno
girgensohnii-Piceetum Polakowski 1962 and Betulo
pubescentis-Piceetum Sokołowski 1980; and (2) the
Eastern Carpathians with associations Bazzanio-
Piceetum (Schmid et Gaisberg 1936) Br.-Bl. et Sissingh
in Br-Bl. et al. 1939, Equiseto-Piceetum Šmarda
1950 and Sphagno-Piceetum Zukrigl 1973 (Budzhak,
Onyshchenko, 2004; Shelyag-Sosonko et al., 2006;
Solomakha, 2008; Solomakha et al., 2016).
A comprehensive evaluation of wet Picea woodlands
of Slovakia has not been published so far and the
syntaxonomical survey of the class Piceetea excelsae
Klika 1948 (Kučera, 2012) was focused on the forest
communities of the supramontane vegetation zone.
However, basic ecological differentiation of Sphagnum-
rich Norway spruce communities (Sphagno-Piceetum
auct.) was presented (Kučera, 2012, p. 249–251).
4 See below subchapter Soldanello montanae-Piceetum.
318 Ukrainian Botanical Journal, 2019, 76(4)
Traditionally, the following associations were
distinguished in Slovakia: Bazzanio-Abietetum (Kuoch
1954) Ellenberg et Klötzli 1972 and Bazzanio-Piceetum
(Schmid et Gaisberg 1936) Br.-Bl. et Sissingh in Br-Bl.
et al. 1939,1 Equiseto-Abietetum Moor 1952, Sphagno
palustris-Piceetum Šomšák 1979, "Leucobryo-Piceetum
Stefanovič 1961"2 (Kontriš, 1981; Majzlanová, 1983,
1993; Šomšák, 1979, 1983; Šomšák et al., 1993,
1996; Kubíček, Šomšák, 1993; Kubíček et al., 1997a,
b) and "Sphagno-Piceetum Hartm."3 (Staszkiewicz,
1993). Many phytocoenological relevés of wet Picea
communities from Slovakia remain unpublished
(master's theses, dissertations, research reports). Some
of them were recently included in the descriptions of
new associations Leucobryo glauci-Piceetum abietis
Šomšák ex P. Kučera 2019 and Stellario nemorum-
Abietetum albae P. Kučera 2019 (Kučera, 2019), all of
the sources are here cited within the particular syntaxa.
The aim of this study is to present formal descriptions
and differentiation of the plant communities of wet
woodlands with Picea abies (and Abies alba) based on
both published and unpublished relevés from Slovakia.
Material and methods
The initial set of phytocoenological relevés of wet
woodlands with Picea abies (especially Sphagnum-
rich wet woodlands) was prepared using the Turboveg
for Windows database software (Hennekens, 2016;
cf. Hennekens, Schaminée, 2001) from the dataset
provided for the prepared monograph Plant communities
of Slovakia, Forest and shrub vegetation (Valachovič
et al., in prep.) stored in Centrálna databáza… (2016).
The detailed description of the next methodological
steps due to the length of the paper are provided in
Electronic Supplement A1 where a dendrogram used
as the basis for syntaxa classification is also given. The
resulting formal characteristic species combinations
of the distiguished associations are given in Electronic
Supplement A2.
Nomenclature of the vascular plants and bryophytes
follows the lists of Marhold et al. (1998) and Kubinská,
Janovicová (1998); if otherwise then with an author
citation, the name Orthodicranum undulatum is given
according to Šomšák's (1976) original data (= Dicranum
bergeri Blandow ?). Syntaxa nomenclature rules are
applied in accordance with the International Code of
Phytosociological Nomenclature (Weber et al., 2000).
1 See below subchapter Soldanello montanae-Piceetum.
2 See below subchapter Leucobryo glauci-Piceetum.
3 Probably Sphagno-Piceetum (Tüxen 1937) Hartmann 1953.
Results and discussion
Division of the class Piceetea excelsae into basic floristic-
ecological groups
Until present, natural Central European Picea abies
woodlands constituting the class Piceetea excelsae
Klika 1948 (syn. Vaccinio-Piceetea Br.-Bl. in Br.-Bl.
et al. 1939 nom. inval., see above) were usually divided
into two orders following the strong differences in the
species composition of communities on the two main
habitat types (Hadač et al., 1969; Kučera, 2010, 2012):
(1) Athyrio-Piceetalia sensu auct. non Hadač 19624
(= Cortuso matthioli-Piceetalia P. Kučera nom. prov.)
on carbonates and (2) Piceetalia excelsae Pawłowski
ex Pawłowski et al. 1928 on silicate bedrock (granites,
quartzites etc.).
The number of species positively differentiating
the supramontane Picea forests of the order Piceetalia
excelsae in Slovakia is very small and they have weak
and very small differential value: Calamagrostis villosa,
Dryopteris dilatata, Vaccinium myrtillus, Avenella
flexuosa. On the contrary, supramontane Picea
forests of the order Cortuso-Piceetalia in Slovakia
are characterized by numerous species and with
a considerable higher differential value: Valeriana
tripteris, Primula elatior, Phyteuma spicatum, Cortusa
matthioli, Polygonatum verticillatum, Mycelis muralis,
Cirsium erisithales, Calamagrostis varia etc. (see Kučera,
2012, tab. 3, columns 10–11).
Comparison of these two units with the collected
relevés of wet woodlands with Picea abies revealed strong
floristic individuality of the wet woodlands expressed by
presence of the species group specific to wet woodlands:
Equisetum sylvaticum, Luzula pilosa, Caltha palustris,
Deschampsia cespitosa, Potentilla erecta, Lysimachia
vulgaris, Polytrichum commune, Sphagnum palustre
agg. etc. (Tab. 1, col. 1 – in Electronic Supplement
B1). The significance of differences in the plant
species composition is equivalent to the phytocoenotic
differential value of the order Cortuso-Piceetalia.
Therefore, the presented wet woodlands with Picea
abies (and Abies alba) are here evaluated as a separate
unit in the rank of order – Sphagno palustris-Piceetalia
P. Kučera 2019 ordo nov. (see below). The arrangement
of Picea abies woodlands of the class Piceetea excelsae
Klika 1948 into three orders reflects prime floristic
differences based on the major ruling ecological patterns
applicable on the continental scale.
4 The question of incorrect syntaxonomical use of the validly published
name Athyrio-Piceetalia Hadač 1962 in the most of geobotanical and
syntaxonomical studies will be discussed in another paper.
319Український ботанічний журнал, 2019, 76(4)
Each of the orders Piceetalia excelsae Pawłowski ex
Pawłowski et al. 1928 and Cortuso matthioli-Piceetalia
P. Kučera nom. prov. (= Athyrio-Piceetalia sensu auct.
non Hadač 1962) comprises only one alliance of
Picea woodlands in the Western Carpathians (Kučera,
2012). It is a result of the regional floristic uniformity
(phytogeography) of the subordinated associations.
On the contrary, associations of the order Sphagno
palustris-Piceetalia P. Kučera 2019 can be grouped into
three superior units, each of them with a specific set
of species reflecting distinctive ecological conditions
(Table 2). In total, eight types of wet woodlands
with Picea abies (and Abies alba) from Slovakia are
recognized in this study (Table 3).
A syntaxonomic overview of Picea abies wet woodland
communities (class Piceetea excelsae Klika 1948) in
Slovakia is provided below:
Piceetalia excelsae Pawłowski ex Pawłowski et al. 1928
Piceion excelsae Pawłowski ex Pawłowski et al. 1928
Sphagno acutifolii-Piceetum Zukrigl 1973
Sphagno palustris-Piceetalia abietis P. Kučera 2019 ordo
nov.
Sphagno palustris-Piceion abietis P. Kučera 2019 all.
nov.
Soldanello montanae-Piceetum Volk in Br.-Bl. et al.
1939
Carex rostrata-Picea abies community
Leucobryo glauci-Piceetum abietis Šomšák ex
P. Kučera 2019
Sphagno palustris-Piceetum Šomšák 1979
Equiseto sylvatici-Piceetum Šmarda 1950
(Calamagrostio villosae-Pinetum Staszkiewicz 1958)
Stellario nemorum-Abietion albae P. Kučera 2019 all.
nov.
Stellario nemorum-Abietetum albae P. Kučera 2019
(Petasito albi-Piceetum Samek 1961)
Valeriano dioicae-Abietion albae P. Kučera 2019 all. nov.
Valeriano dioicae-Abietetum P. Kučera 2019 ass. nov.
Description of syntaxa of wet woodlands with Picea
abies from Slovakia
I. Piceetalia excelsae Pawłowski ex Pawłowski et al.
1928
This order comprises natural plant communities
forming in the Western Carpathians a separate altitu-
dinal vegetation zone of climax Picea abies woodland,
in the Tatras also with Pinus cembra and Larix decidua
(Kučera, 2012, 2017; Zięba et al., 2018).
On the bog ecotones, a series of vegetation types
between communities of the classes Oxycocco-
Sphagnetea Br.-Bl. et Tx. ex Westhof et al. 1964 and
Piceetea excelsae develops, regularly including spatially
more or less developed forest (or krummholz-forest)
bog communities of the class Vaccinio uliginosi-Pinetea
Passarge 1968.
Bogside ecotones or groundwater-influenced
habitats confined to gentle (moderate) slopes adjacent
to mountain plateaus of some West Carpathians
mountain ranges bear forest communities (stable
communities, Dauergesellschaften) with subsiding
occurrence of Eriophorum vaginatum, Carex nigra and
selected Sphagnum species other than S. girgensohnii. At
the same time they still support the constant presence
of species characteristic to climax supramontane
Picea forests on acid soils, i.e. Athyrium distentifolium,
Dryopteris expansa, D. dilatata, Homogyne alpina,
Polytrichum formosum. Therefore these phytocoenoses
are here classified as a peripheral member of the
alliance (I. A.) Piceion excelsae Pawłowski ex Pawłowski
et al. 1928 close to the class Vaccinio uliginosi-Pinetea
Passarge 1968. This group is positively differentiated
by Juncus filiformis and Sphagnum capillifolium in the
frame of the evaluated relevé set from Slovakia (Tab. 3).
Until present, only six relevés were published
from Slovakia, although the community has wider
distribution (Kučera, in prep.). Within the group of wet
Picea communities, their species composition has close
relations to the association Sphagno acutifolii-Piceetum
Zukrigl 1973 s. str. (i.e. in the sense of the lectotype
relevé).
1. Sphagno acutifolii-Piceetum Zukrigl 1973
Original diagnosis: Zukrigl (1973), p. 152, tab. 6.
Nomenclatural type: Zukrigl (1973), p. 152, tab 6,
rel. 2, lectotype; Willner, Zukrigl (1999), p. 154.
Characteristic species combination: see Electronic
Supplement A2.
Data: Kučera (2005): p. 65, rel. 3; Kučera (2012):
p. 311–312, rel. 74–77, p. 317, rel. 97.
Relevés from Slovakia presented here correspond
to the association Sphagno acutifolii-Piceetum Zukrigl
1973 (cf. Electronic Supplement A3, section I). Stands
of this association in the Western Carpathians are
dominated by Picea abies, which is in accordance with
the community distribution mostly in the supramontane
altitudinal vegetation zone. Sometimes Sorbus
aucuparia is admixed, partial Pinus mugo occurrence is
connected with adjacent krummholz stands.
320 Ukrainian Botanical Journal, 2019, 76(4)
No.
of relevés
A B C
89 36 13
Trees and shrubs
E
3
Abies alba –6 6278 –23
Fagus sylvatica –. 4325 –.
Pinus sylvestris –31 –. 4762
Alnus glutinosa –15 –. 3838
Alnus incana –21 –3 3846
Betula pendula –17 –. 3738
E
2
Picea abies 3194 –64 –69
Fagus sylvatica –. 5742 –.
Sorbus aucuparia –8 3756 –31
Lonicera xylosteum –. 3417 –.
Acer pseudoplatanus –. 3114 –.
Salix caprea –1 2914 –.
Sambucus racemosa –1 2511 –.
Frangula alnus –11 –. 4946
Viburnum opulus –. –. 4123
Lonicera nigra –3 1144 3762
E
1
Betula pubescens 3118 –3 –.
Salix aurita 2811 –. –.
Fagus sylvatica –4 3322 –.
Lonicera xylosteum –. 3114 –.
Abies alba –22 2572 2169
Daphne mezereum –. –. 4831
Lonicera nigra –16 –11 4354
Viburnum opulus –. –. 4123
Sorbus aucuparia –47 –56 4092
Frangula alnus –12 –. 3331
Betula pendula –7 –. 3123
Ribes petraeum –1 –. 3115
Differential field layer species (E
1
)
Vaccinium vitis-idaea 4694 –25 –69
Carex canescens 3631 –3 –8
Ranunculus flammula 3316 –. –.
Juncus effusus 3316 –. –.
Melampyrum sylvaticum 3316 –. –.
Agrostis canina 3225 –. –8
Potentilla erecta 3138 –. –23
Agrostis stolonifera 3118 –3 –.
Carex echinata 3030 –. –15
Trientalis europaea 2912 –. –.
Lysimachia vulgaris 2936 –. –23
Valeriana simplicifolia 2610 –. –.
Table 2. Differential table of alliances of the order Sphagno palustris-Piceetalia abietis P. Kučera 2019 with fidelity
(φ (× 100) ≥ 25) and constancy (%) in the exponent
A – Sphagno palustris-Piceion abietis P. Kučera 2019
B – Stellario nemorum-Abietion albae P. Kučera 2019
C – Valeriano dioicae-Abietion albae P. Kučera 2019
No.
of relevés
A B C
89 36 13
Stellaria nemorum –1 8378 –.
Chrysosplenium alternifolium –2 6969 –8
Petasites albus –1 6867 –8
Lysimachia nemorum –1 6250 –.
Geranium robertianum –. 5539 –.
Cardamine trifolia –. 5539 –.
Adenostyles alliariae –. 5539 –.
Luzula luzulina –. 5236 –.
Homogyne alpina –21 5158 –.
Impatiens noli-tangere –. 5144 –8
Urtica dioica –8 5044 –.
Rubus hirtus –. 5033 –.
Gentiana asclepiadea –3 4967 –31
Galium odoratum –. 4831 –.
Prenanthes purpurea –1 4661 –31
Dryopteris dilatata –1 4631 –.
Phegopteris connectilis –3 4331 –.
Milium effusum –. 4325 –.
Ranunculus lanuginosus –. 4022 –.
Ranunculus platanifolius –. 4022 –.
Oxalis acetosella –54 39100 –77
Senecio ovatus –19 3783 1769
Phyteuma spicatum –. 3719 –.
Rubus idaeus –35 3783 –54
Chaerophyllum hirsutum –11 3578 2369
Luzula sylvatica –3 3522 –.
Cardamine flexuosa –. 3417 –.
Calamagrostis epigejos –. 3417 –.
Poa remota –. 3417 –.
Veronica anagallis-aquatica –. 3417 –.
Equisetum sylvaticum –72 32100 –77
Dryopteris filix-mas –12 3253 –31
Cicerbita alpina –. 3114 –.
Geum rivale –2 3039 –23
Carex sylvatica –. 3031 –15
Calamagrostis arundinacea –22 2961 –38
Deschampsia cespitosa –33 2953 –15
Ranunculus repens –17 2539 –15
Doronicum austriacum –1 2511 –.
Rubus saxatilis –1 –. 9392
Valeriana dioica –4 –. 7977
Polygonatum verticillatum –2 –6 7069
Caltha palustris –38 –19 67100
Crepis paludosa –35 –28 65100
Luzula pilosa 965 –11 59100
Cirsium oleraceum –. –. 5438
321Український ботанічний журнал, 2019, 76(4)
No.
of relevés
A B C
89 36 13
Clematis alpina –. –. 5438
Maianthemum bifolium –53 –44 51100
Filipendula ulmaria –3 –14 5054
Thalictrum aquilegiifolium –. –. 4831
Carex alba –. –. 4831
Galium schultesii –. –3 4431
Fragaria vesca –3 –25 4254
Solidago virgaurea –10 –31 4162
Melica nutans –. –. 4123
Astrantia major –. –. 4123
Paris quadrifolia –6 –. 4031
Equisetum palustre –3 –3 4031
Dactylorhiza maculata –1 –. 3923
Carex digitata –. –. 3315
Actaea spicata –. –. 3315
Carex remota –4 –8 3331
Epipactis palustris –1 –. 3115
Bistorta major –2 –. 2915
Polygonatum multiflorum –. –3 2815
Valeriana tripteris –. –3 2815
Angelica sylvestris –1 –3 2615
Differential ground layer species (E
0
)
Polytrichum commune 3861 –36 –8
Lepidozia reptans 3131 –. –15
Pohlia nutans 3024 –. –8
Sphagnum recurvum agg. 2811 –. –.
Chiloscyphus pallescens 2610 –. –.
Herzogiella seligeri 259 –. –.
Plagiomnium affine –12 5472 –23
Cirriphyllum piliferum –1 5136 –.
Plagiothecium undulatum –2 3925 –.
Plagiomnium undulatum –4 3331 –8
Plagiothecium curvifolium –24 3144 –8
Plagiomnium rostratum –. 3114 –.
Conocephalum conicum –. 3114 –.
Thuidium tamariscinum –. 2811 –.
Trichocolea tomentella –1 –. 3923
Eurhynchium angustirete –9 –. 3731
Tetraphis pellucida –11 –. 3431
Vaccinium myrtillus is the dominant species of the
field layer, constantly accompanied by Homogyne
alpina, Avenella flexuosa, V. vitis-idaea, more frequent
are also species Calamagrostis villosa, Dryopteris dilatata,
Athyrium distentifolium. The differential attribute
against the other communities of the alliance Piceion
excelsae Pawłowski ex Pawłowski et al. 1928 (especially
Vaccinio myrtilli-Piceetum Šoltés 1976) is occurrence
of Eriophorum vaginatum and Juncus filiformis, Carex
canescens, C. nigra, (Nardus stricta); however, also
C. echinata or C. pauciflora could be present.
Ground layer is defined by Polytrichum commune,
P. formosum, Sphagnum capillifolium, Dicranum
scoparium, less frequent are S. girgensohnii and
Plagiothecium curvifolium. Other peat moss species were
also recorded (in one of total six relevés): S. rubellum
and S. russowii.
Phytocoenological records of the association
Sphagno acutifolii-Piceetum come from Martinské hole
(Veterné hole Mts) and Kubínska hoľa Mt. (Oravská
Magura Mts). Stands are distributed above 1 390 m
a.s.l., only ocassionally below 1 300 m (Kubínska hoľa
Mt.) and then in more species-rich variant.
Variability of the association in Slovakia is poorly
known as only six relevés were published. However,
some differences could be identified in the species
composition either of field layer or ground layer (see
Kučera, 2012, rel. 74–77 vs. Kučera, 2005, rel. 3 vs.
Kučera, 2012, rel. 97).
Nomenclatural and syntaxonomical note on the name
type "Sphagno-Piceetum"
As shown in Electronic Supplement A3, section I
(Willner, 2007; Kučera, 2012; Chytrý et al., 2013),
application of names with the species combination
Sphagnum-Picea should strictly follow determination of
the validly published original diagnosis of a particular
syntaxon. At the same time, their careful consideration
is needed because they could label different syntaxa. For
example, the proposal of Chytrý et al. (2013) to reject
Sphagno-Piceetum (Tüxen 1937) Hartmann 1953 as a
nomen ambiguum (cf. Art. 36) does not solve problems
of later multiple descriptions of "Sphagno-Piceetum"
syntaxa from which several are not homonyms what
Chytrý et al. (2013) stated.
Selected cases are briefly discussed in Electronic
Supplement A3; however, the necessary nomenclatural
proposals are given here to assure their effective
publication in the respect of the current version of the
ICPN (Weber et al. 2000, Art. 1):
A) Completion of the name Piceetum excelsae
sphagnetosum Tüxen 1937 (see Tüxen 1937, p. 123)
according to ICPN Rec. 10C:
Table 2. Continuation
322 Ukrainian Botanical Journal, 2019, 76(4)
Table 3. Differential table of all associations of the wet woodlands with Picea abies with constancy (%) and fidelity
(φ (× 100) ≥ 25) in the exponent
I – order Piceetalia excelsae Pawłowski ex Pawłowski et al. 1928
A – alliance Piceion excelsae Pawłowski ex Pawłowski et al. 1928
1 – Sphagno acutifolii-Piceetum Zukrigl 1973
II – order Sphagno palustris-Piceetalia abietis P. Kučera 2019
B – alliance Sphagno palustris-Piceion abietis P. Kučera 2019
2 – Soldanello montanae-Piceetum Volk in Br.-Bl. et al. 1939
3 – Carex rostrata-Picea abies community (with cover-abundance values in italics)
4 – Leucobryo glauci-Piceetum abietis Šomšák ex P. Kučera 2019
5 – Sphagno palustris-Piceetum Šomšák 1979
6 – Equiseto sylvatici-Piceetum Šmarda 1950
C – alliance Stellario nemorum-Abietion albae P. Kučera 2019
7 – Stellario nemorum-Abietetum albae P. Kučera 2019
D – alliance Valeriano dioicae-Abietion albae P. Kučera 2019
8 – Valeriano dioicae-Abietetum P. Kučera 2019
Field and ground layer species with constancy less than 10 % in a single column are omitted (except Sphagnum subsp.) as well as
one relevé records of Salix sp. (E
1
), Athyrium sp., Taraxacum sect. Ruderalia, Ranunculus acris (in group No. 1).
Order I II II II
Alliance A B C D
Group No. 1 2 3 4 5 6 7 8
No. of relevés 6 24 1 15 37 13 36 13
Differential tree and shrub species
E
3
Sorbus aucuparia 33 41 4 – . .– .– 8– 6– .–
Abies alba . – 17 – . .– .– 8– 7864 23–
Fagus sylvatica . – .– . .– .– .– 2547 .–
Pinus sylvestris .– 17– . 4717 32– 38– .– 6230
Alnus incana .– 4– . 7– 3214 38– 3– 4629
Alnus glutinosa .– .– . 20– 19– 23– .– 3828
E
2
Pinus mugo 3355 .– . .– .– .– .– .–
Pinus sylvestris .– 827 . .– .– .– .– .–
Alnus incana .– .– . .– 2210 4640 .– 23–
Fagus sylvatica 17– .– . .– .– .– 4249 .–
Lonicera xylosteum .– .– . .– .– .– 1738 .–
Acer pseudoplatanus .– .– . .– .– .– 1435 .–
Sorbus aucuparia 17– 4– . 7– .– 38– 5633 31–
Salix caprea .– .– . .– .– 8– 1426 .–
Viburnum opulus .– .– . .– .– .– .– 2345
Frangula alnus .– 4– . 13– 11– 23– .– 4638
Lonicera nigra .– .– . .– 3– 15– 4429 6247
E
1
Pinus mugo 3355 .– . .– .– .– .– .–
Salix aurita .– .– . 7– 2439 .– .– .–
Corylus avellana .– .– . .– 827 .– .– .–
Lonicera xylosteum .– .– . .– .– .– 1435 .–
Fagus sylvatica .– 17– . .– .– .– 2230 .–
Daphne mezereum .– .– . .– .– .– .– 3153
Viburnum opulus .– .– . .– .– .– .– 2345
Ribes petraeum .– .– . .– 3– .– .– 1533
Sorbus aucuparia 67– 21– . 60– 54– 62– 56– 9228
Frangula alnus .– 4– . 20– 19– .– .– 3127
Lonicera nigra .– 4– . .– 19– 4628 11– 5436
Abies alba 17– 38– . 7– 11– 46– 7230 6927
323Український ботанічний журнал, 2019, 76(4)
Order I II II II
Alliance A B C D
Group No. 1 2 3 4 5 6 7 8
No. of relevés 6 24 1 15 37 13 36 13
Other tree and shrub species
E
3
Picea abies 100– 100– 2n/a 100– 100– 100– 100– 100–
Betula pubescens .– 29– . 7– 3014 23– .– 31–
Betula pendula .– 17– . 7– 14– 3825 .– 3825
Larix decidua .– .– . 13– 3– .– .– 8–
Salix cinerea .– 4– . .– 3– .– .– .–
Salix ×multinervis .– .– . .– 3– .– .– .–
Frangula alnus .– .– . .– 3– .– .– .–
Salix pentandra .– .– . .– 3– .– .– .–
E
2
Picea abies 50– 9616 2n/a 93– 9515 92– 64– 69–
Abies alba .– 8– . .– .– 15– 1713 15–
Alnus glutinosa .– .– . 13– 5– 15– .– 8–
Betula pubescens .– 4– . .– 5– 15– .– .–
Sambucus racemosa .– 4– . .– .– .– 1125 .–
Salix cinerea .– 4– . .– 3– 8– .– .–
Pinus ×celakovskiorum A. et Gr. .– 4– . .– .– .– .– .–
Sambucus nigra .– 4– . .– .– .– .– .–
Salix aurita .– .– . .– 3– .– .– .–
Juniperus communis .– .– . .– 3– .– .– .–
Larix decidua .– .– . .– 3– .– .– .–
Betula pendula .– .– . .– .– 8– .– .–
Padus avium .– .– . .– .– .– .– 8–
E
1
Picea abies 67– 92– . 93– 89– 100– 78– 100–
Alnus incana .– 8– . 13– 2724 8– 3– 8–
Betula pubescens .– 12– . 7– 2420 23– 3– .–
Acer pseudoplatanus .– 17– . .– .– 15– 11– 15–
Betula pendula .– 8– . 7– 3– 15– .– 2323
Alnus glutinosa .– .– . 2021 5– 8– .– 15–
Salix caprea .– .– . 7– 3– 8– 8– .–
Rosa pendulina .– .– . .– 5– .– 6– 15–
Salix aurita .– .– . 7– 8– .– .– .–
Salix cinerea .– 4– . .– .– 8– .– 8–
Pinus sylvestris .– 4– . 7– .– 8– .– .–
Padus avium .– .– . .– .– 8– .– 8–
Sambucus racemosa .– .– . .– .– .– 6– .–
Ribes uva-crispa .– .– . .– 3– .– .– .–
Salix silesiaca .– .– +n/a .– .– .– 3– .–
Differential field layer species (E
1
)
Juncus filiformis 8387 .– . .– 5– .– .– .–
Eriophorum vaginatum 8381 173 . .– .– .– .– .–
Dryopteris dilatata 8372 4– . .– .– .– 3115 .–
Athyrium distentifolium 6772 .– . .– 5– .– 6– .–
Dryopteris expansa 5066 .– . .– .– .– 3– .–
Nardus stricta 6763 12– . .– 14– .– 3– .–
Homogyne alpina 10057 42– 2n/a .– 16– 23– 5821 .–
Carex nigra 6750 12– +n/a 7– 279 .– 3– 15–
324 Ukrainian Botanical Journal, 2019, 76(4)
Order I II II II
Alliance A B C D
Group No. 1 2 3 4 5 6 7 8
No. of relevés 6 24 1 15 37 13 36 13
Carex canescens 6738 21– . 20– 4921 15– 3– 8–
Avenella flexuosa 8335 21– . 6721 41– 15– 42– 23–
Listera cordata .– 1229 . .– .– .– 3– .–
Thelypteris palustris .– 827 +n/a .– .– .– .– .–
Melampyrum sylvaticum .– 4– . 4749 14– 8– .– .–
Luzula luzuloides .– 4– . 4732 3216 .– 22– 15–
Calluna vulgaris .– 4– . 2030 .– 8– .– .–
Agrostis canina .– 4– . 7– 5459 .– .– 8–
Viola palustris .– .– . .– 4154 .– .– 8–
Ranunculus flammula .– .– . 7– 3550 .– .– .–
Juncus effusus .– 4– . .– 3244 8– .– .–
Potentilla erecta .– 12– . 20– 6543 31– .– 23–
Agrostis stolonifera .– 4– . 7– 3541 8– 3– .–
Carex rostrata .– .– 2n/a .– 1638 .– .– .–
Valeriana simplicifolia .– .– . .– 2236 8– .– .–
Ajuga reptans .– .– . 7– 2432 .– 3– 8–
Carex pallescens .– .– . .– 1131 .– .– .–
Moneses uniflora .– .– . .– 1629 8– .– .–
Senecio "nemorensis" .– 4– . .– 1429 .– .– .–
Galium palustre .– 4– . .– 1927 8– 3– .–
Peucedanum palustre .– .– . .– 827 .– .– .–
Galium uliginosum .– .– . .– 827 .– .– .–
Melampyrum pratense .– .– . .– 827 .– .– .–
Orthilia secunda .– 8– . 33– 5126 31– .– 46–
Trientalis europaea .– 8– . .– 11– 3845 .– .–
Veratrum album subsp. lobelianum .– 12– +n/a .– 8– 3827 22– 23–
Stellaria nemorum .– .– . .– 3– .– 7885 .–
Chrysosplenium alternifolium .– .– . .– 5– .– 6973 8–
Petasites albus .– 4– . .– .– .– 6772 8–
Lysimachia nemorum .– .– . .– 3– .– 5066 .–
Adenostyles alliariae .– .– . .– .– .– 3959 .–
Cardamine trifolia .– .– . .– .– .– 3959 .–
Geranium robertianum .– .– . .– .– .– 3959 .–
Impatiens noli-tangere .– .– . .– .– .– 4458 8–
Luzula luzulina .– .– . .– .– .– 3657 .–
Prenanthes purpurea .– .– . .– 3– .– 6157 31–
Gentiana asclepiadea .– 4– . .– 3– 8– 6756 31–
Rubus hirtus .– .– . .– .– .– 3355 .–
Galium odoratum .– .– . .– .– .– 3152 .–
Milium effusum .– .– . .– .– .– 2547 .–
Urtica dioica .– .– . .– 14– 15– 4445 .–
Ranunculus lanuginosus .– .– . .– .– .– 2244 .–
Ranunculus platanifolius .– .– . .– .– .– 2244 .–
Phyteuma spicatum .– .– . .– .– .– 1941 .–
Geum rivale .– .– . .– 5– .– 3941 23–
Carex sylvatica .– .– . .– .– .– 3140 15–
Veronica anagallis-aquatica .– .– . .– .– .– 1738 .–
Cardamine flexuosa .– .– . .– .– .– 1738 .–
Calamagrostis epigejos .– .– . .– .– .– 1738 .–
325Український ботанічний журнал, 2019, 76(4)
Order I II II II
Alliance A B C D
Group No. 1 2 3 4 5 6 7 8
No. of relevés 6 24 1 15 37 13 36 13
Poa remota .– .– . .– .– .– 1738 .–
Phegopteris connectilis .– .– . .– 3– 15– 3138 .–
Rubus idaeus .– 12– . 20– 49– 54– 8337 54–
Oxalis acetosella .– 50– . 40– 51– 85– 10035 77–
Cicerbita alpina .– .– . .– .– .– 1435 .–
Dryopteris filix-mas .– 8– . 7– 14– 23– 5335 31–
Luzula sylvatica .– 12– . .– .– .– 2232 .–
Epilobium montanum .– .– . .– 8– .– 3331 31–
Cardamine amara .– .– . .– 11– .– 2528 15–
Poa palustris .– .– . .– 3– .– 1727 8–
Symphytum tuberosum .– .– . .– .– .– 827 .–
Sanicula europaea .– .– . .– .– .– 827 .–
Rubus saxatilis .– .– . .– 3– .– .– 9294
Valeriana dioica .– .– . 7– 5– 8– .– 7775
Polygonatum verticillatum .– .– . .– 3– 8– 6– 6971
Clematis alpina .– .– . .– .– .– .– 3859
Cirsium oleraceum .– .– . .– .– .– .– 3859
Filipendula ulmaria .– .– . .– 8– .– 14– 5456
Crepis paludosa .– 4– . 20– 5414 54– 28– 10053
Thalictrum aquilegiifolium .– .– . .– .– .– .– 3153
Carex alba .– .– . .– .– .– .– 3153
Caltha palustris .– 8– . 33– 5413 54– 19– 10052
Galium schultesii .– .– . .– .– .– 3– 3150
Fragaria vesca .– .– . .– 5– 8– 2514 5449
Melica nutans .– .– . .– .– .– .– 2345
Astrantia major .– .– . .– .– .– .– 2345
Solidago virgaurea .– .– . 13– 8– 31– 3110 6241
Maianthemum bifolium .– 8– . 67– 6813 77– 44– 10039
Carex remota .– .– . .– 11– .– 8– 3138
Dactylorhiza maculata .– .– . .– .– 8– .– 2337
Carex digitata .– .– . .– .– .– .– 1537
Actaea spicata .– .– . .– .– .– .– 1537
Equisetum palustre .– .– . .– 3– 15– 3– 3137
Epipactis palustris .– .– . .– 3– .– .– 1533
Valeriana tripteris .– .– . .– .– .– 3– 1533
Polygonatum multiflorum .– .– . .– .– .– 3– 1533
Paris quadrifolia .– .– . .– 5– 2321 .– 3133
Angelica sylvestris .– .– . .– 3– .– 3– 1530
Dentaria glandulosa .– .– . .– .– .– 1923 2329
Bistorta major .– .– . 7– 3– .– .– 1526
Calamagrostis arundinacea .– .– . 6735 24– 8– 6130 38–
Lysimachia vulgaris .– 4– . 20– 5430 6237 .– 23–
Equisetum sylvaticum 17– 42– . 60– 86– 10027 10027 77–
Chaerophyllum hirsutum .– .– . 7– 19– 15– 7847 6939
Senecio ovatus .– .– . 13– 24– 46– 8343 6931
Luzula pilosa .– 4– . 9332 8626 8525 11– 10037
Other field layer species (E
1
)
Vaccinium myrtillus 100– 100– +n/a 100– 97– 100– 75– 100–
Vaccinium vitis-idaea 83– 10021 . 93– 9213 92– 25– 69–
326 Ukrainian Botanical Journal, 2019, 76(4)
Order I II II II
Alliance A B C D
Group No. 1 2 3 4 5 6 7 8
No. of relevés 6 24 1 15 37 13 36 13
Calamagrostis villosa 83– 62– 2n/a 60– 81– 10024 61– 69–
Dryopteris carthusiana .– 42– . 67– 7619 77– 58– 46–
Athyrium filix-femina .– 8– . 67– 7318 62– 7520 69–
Deschampsia cespitosa 17– 8– 1n/a 13– 5423 38– 5322 15–
Hieracium murorum .– 12– . 40– 16– 31– 5323 38–
Myosotis palustris spojene .– 4– . 7– 4122 15– 36– 31–
Carex echinata 33– 4– 1n/a 40– 4622 23– .– 15–
Ranunculus repens .– .– . 7– 27– 31– 3924 15–
Gymnocarpium dryopteris .– 4– . 7– 22– 23– 19– 31–
Veronica officinalis .– .– . 13– 3222 23– 14– 15–
Lycopodium annotinum 33– 3825 . 13– .– .– 8– 15–
Glyceria nemoralis .– .– . .– 2423 .– 19– 15–
Mycelis muralis .– .– . .– 19– 8– 17– 23–
Anemone nemorosa .– .– . .– 3– 15– 1918 15–
Huperzia selago .– 4– . 7– 5– 8– 6– 2324
Cirsium palustre .– .– . 7– 1618 8– 3– 8–
Glyceria fluitans .– .– . 13– 1623 .– 3– .–
Oxycoccus palustris .– 12– . .– 8– .– .– .–
Soldanella hungarica .– .– . .– 1118 8– 3– .–
Pyrola rotundifolia .– .– . .– 1121 .– .– 8–
Calamagrostis canescens .– .– . .– 8– .– .– 15–
Prunella vulgaris .– .– . .– 3– 8– 3– 15–
Doronicum austriacum .– .– . .– .– 8– 1121 .–
Alchemilla sp. 17– .– . .– .– .– 6– .–
Rumex alpinus 17– .– . .– .– .– 3– .–
Carex pauciflora 17– 4– . .– .– .– .– .–
Hypericum maculatum .– 4– +n/a .– .– 8– .– .–
Differential ground layer species (E
0
)
Sphagnum capillifolium 8358 12– . 7– 16– 23– .– 23–
Sphagnum rubellum 3355 .– . .– .– .– .– .–
Lophocolea heterophylla 3343 .– . 7– .– 8– .– .–
Polytrichum formosum 8341 25– . 47– 22– 8– 22– 38–
Calypogeia azurea 3340 4– . .– .– 8– .– 8–
Dicranum fuscescens 1738 .– . .– .– .– .– .–
Polytrichum alpinum 1738 .– . .– .– .– .– .–
Barbilophozia floerkei 1738 .– . .– .– .– .– .–
Sphagnum fuscum 1738 .– . .– .– .– .– .–
Barbilophozia attenuata 1738 .– . .– .– .– .– .–
Pleuridium subulatum 1738 .– . .– .– .– .– .–
Polytrichum commune 10037 7517 4n/a 47– 57– 62– 36– 8–
Leucobryum glaucum .– .– . 10084 11– 8– .– 15–
Orthodicranum undulatum .– .– . 1334 .– .– .– .–
Hylocomium splendens .– 8– . 5332 4624 15– 19– 8–
Brachythecium starkei .– .– . .– 1430 .– 3– .–
Lepidozia reptans .– .– . 33– 4930 38– .– 15–
Sphagnum palustre agg. .– 50– 1n/a 27– 7028 46– 3– 62–
Chiloscyphus pallescens .– .– . 7– 1927 8– .– .–
Sphagnum quinquefarium .– .– . .– 827 .– .– .–
Rhodobryum roseum .– .– . .– 827 .– .– .–
327Український ботанічний журнал, 2019, 76(4)
Order I II II II
Alliance A B C D
Group No. 1 2 3 4 5 6 7 8
No. of relevés 6 24 1 15 37 13 36 13
Cephalozia bicuspidata .– .– . .– 827 .– .– .–
Calliergon cordifolium .– .– . .– 827 .– .– .–
Plagiochila asplenioides .– .– . 7– 4326 23– 17– 38–
Lophocolea bidentata .– .– . .– 1914 3841 .– 8–
Sphagnum recurvum agg. .– 4– . .– 1617 2329 .– .–
Bazzania trilobata .– 2916 . .– 5– 3827 .– 31–
Cirriphyllum piliferum .– .– . .– 3– .– 3655 .–
Plagiomnium affine .– 4– . 20– 8– 31– 7248 23–
Plagiothecium undulatum .– 8– . .– .– .– 2539 .–
Plagiomnium rostratum .– .– . .– .– .– 1435 .–
Conocephalum conicum .– .– . .– .– .– 1435 .–
Plagiomnium undulatum .– .– . .– 5– 15– 3132 8–
Thuidium tamariscinum .– .– . .– .– .– 1131 .–
Trichocolea tomentella .– .– . .– .– 8– .– 2337
Eurhynchium angustirete .– .– . 7– 14– 15– .– 3130
Tetraphis pellucida .– .– . 7– 1912 15– .– 3128
Other ground layer species (E
0
)
Dicranum scoparium 83– 67– . 93– 81– 92– 61– 85–
Sphagnum girgensohnii 67– 8322 2bn/a 47– 65– 38– 53– 46–
Pleurozium schreberi 33– 54– . 8021 658 69– 11– 69–
Plagiothecium curvifolium 67– 8– . 27– 24– 46– 4411 8–
Rhytidiadelphus triquetrus .– 4– . 27– 4624 31– 6– 38–
Rhizomnium punctatum .– .– . 13– 27– 23– 3620 23–
Sphagnum squarrosum .– 17– +n/a .– 3017 31– 6– 23–
Pohlia nutans 17– 8– . 20– 3522 23– .– 8–
Calypogeia integristipula 33– .– . 20– 227 8– .– 23–
Dicranum montanum 33– 4– . .– 14– 15– .– 15–
Mnium sp. .– .– . 7– 2222 15– .– 8–
Climacium dendroides .– .– . .– 2222 15– .– 15–
Dicranella heteromalla 17– 4– . 2722 8– .– .– 15–
Sphagnum sp. .– 8– . 20– 3– 8– .– 15–
Herzogiella seligeri .– .– . 7– 1416 15– .– .–
Pellia sp. .– .– . 7– 1116 8– .– .–
Sphagnum russowii 17– .– . .– 8– 8– .– .–
Dicranum polysetum .– 4– . .– 1124 .– .– .–
Plagiothecium laetum 17– .– . 7– .– .– 3– 8–
Barbilophozia lycopodioide 17– 4– . .– .– .– .– 8–
Sphagnum magellanicum .– .– 1n/a 7– 3– .– .– .–
Sphagnum flexuosum .– .– 1n/a .– 3– 8– .– .–
Sphagnum subnitens .– 4– . .– .– .– .– .–
Sphagnum fallax .– .– . .– 3– .– .– .–
Sphagnum obtusum .– .– . .– 3– .– .– .–
Sphagnum cuspidatum .– .– . .– 3– .– .– .–
Sphagnum riparium .– .– . .– .– 8– .– .–
Sphagnum teres .– .– . .– .– 8– .– .–
Other species in the column 3 only:
Agrostis tenuis +, Eriophorum angustifolium +, Drepanocladus fluitans +.
328 Ukrainian Botanical Journal, 2019, 76(4)
Piceetum excelsae sphagnetosum quinquefarii Tüxen
1937
Consequently, the association name published by
Hartmann (1953) is to be completed as follows:
Sphagno quinquefarii-Piceetum (Tüxen 1937)
Hartmann 1953; see Hartmann (1953), Anhang,
p. XIII.
B) Lectotypes for the association Sphagno
girgensohnii-Piceetum Polakowski 1962 nom. cons.
propos. and its subunits (ICPN Def. VIII, Art. 19):
Sphagno girgensohnii-Piceetum Polakowski 1962
nom. cons. propos.
Nomenclatural type: Polakowski (1962), tab. 4,
rel. 29; lectotypus hoc loco.
Sphagno girgensohnii-Piceetum lycopodietosum
annotini Polakowski 1962
Nomenclatural type: Polakowski (1962), tab. 4,
rel. 29; lectotypus hoc loco.
Sphagno girgensohnii-Piceetum vaccinietosum myrtilli
Polakowski 1962 (Art. 14)
Nomenclatural type: Polakowski (1962), tab. 4,
rel. 54; lectotypus hoc loco.
II. Sphagno palustris-Piceetalia P. Kučera ordo nov.
hoc loco
Original diagnosis: Sphagno palustris-Piceion abietis
P. Kučera 2019 all. nov., Sphagno girgensohnii-Piceion
(Kielland-Lund 1981) P. Kučera 2019 stat. nov.,
Stellario nemorum-Abietion albae P. Kučera 2019 all.
nov., Valeriano dioicae-Abietion albae P. Kučera 2019
all. nov.
Nomenclatural type: Sphagno palustris-Piceion
P. Kučera 2019 all. nov., holotypus hoc loco.
Differential species (φ (× 100) ≥ 25) (145 relevés,
Tab. 1 – in Electronic Supplement B1):
E
3
: Pinus sylvestris (42), Abies alba (38), Alnus
incana (36), Betula pendula (31), Alnus glutinosa (29),
B. pubescens (27),
E
2
: Picea abies (48), Alnus incana (29), Frangula
alnus (28),
E
1
: Abies alba (47), Picea abies (43), Alnus incana
(29) Frangula alnus (27), Salix aurita (26),
Equisetum sylvaticum (82), Luzula pilosa (63),
Caltha palustris (52), Deschampsia cespitosa (45),
Dryopteris carthusiana (43), Potentilla erecta (43),
Lysimachia vulgaris (42), Maianthemum bifolium
(39), Myosotis palustris agg. (38), Carex echinata (38),
Carex canescens (37), Ranunculus repens (34), Agrostis
canina (33), Orthilia secunda (32), Vaccinium vitis-
idaea (32), Lysimachia nemorum (30), Athyrium filix-
femina (30), Carex nigra (30), Glyceria nemoralis (29),
Agrostis stolonifera (29), Veronica officinalis (28), Viola
palustris (28), Filipendula ulmaria (27), Impatiens noli-
tangere (26), Juncus effusus (26), Cardamine trifolia
(26), Ranunculus flammula (26), Valeriana dioica (26),
Petasites albus (25),
E
0
: Polytrichum commune (55), Sphagnum palustre
agg. (54), S. girgensohnii (48), S. squarrosum (33),
Leucobryum glaucum (31), Plagiomnium affine (30),
Pohlia nutans (30), Pleurozium schreberi (28).
The order Sphagno palustris-Piceetalia P. Kučera
2019 constitutes a separate unit of the class Piceetea
excelsae Klika 1948 alongside with the orders Cortuso-
Piceetalia (Athyrio-Piceetalia sensu auct. non Hadač
1962) and Piceetalia excelsae Pawłowski ex Pawłowski
et al. 1928. It comprises all types of wet woodlands with
dominating trees Picea abies (occasionally Abies alba),
its peripheral phytocoenoses mediate connections to
the classes Vaccinio uliginosi-Pinetea Passarge 1968 and
Alnetea glutinosae Br.-Bl. et Tüxen ex Westhoff et al.
1946.
Formal floristic differentiation of the order Sphagno
palustris-Piceetalia in Slovakia consists for the great part
of species exclusive to wet woodlands within the class
Piceetea excelsae Klika 1948: Pinus sylvestris, Alnus
glutinosa + A. incana, Betula pubescens, Equisetum
sylvaticum, Caltha palustris, Deschampsia cespitosa,
Potentilla erecta, Lysimachia vulgaris, Sphagnum palustre
agg. etc. (see Tab. 1 – in Electronic Supplement B1).
On the base of floristic differences which reflect
specific ecological conditions of the seven differentiated
communities from Slovakia (Tab. 3; Tab. 4 – in
Electronic Supplement B2), three subunits of the order
Sphagno palustris-Piceetalia are distinguished in this
study (Tab. 2) and they are given the rank of alliance:
(1) Sphagno palustris-Piceion P. Kučera 2019 all. nov.
with oligotrophic communities on nutrient-poor shal-
low peat layers of histosols or other poor hydromophic
soils (gleysols, stagnosols) with raw humus layer
(Sphagno palustris-Piceetum Šomšák 1979, Equiseto
sylvatici-Piceetum Šmarda 1950, Soldanello montanae-
Piceetum Volk in Br.-Bl. et al. 1939 and Carex rostrata-
Picea abies community) or on temporarily flooded
humic podzols on the margin of mires (Leucobryo
glauci-Piceetum abietis Šomšák ex P. Kučera 2019);
(2) Stellario nemorum-Abietion albae P. Kučera 2019
all. nov. with communities on minerotrophic habitats
of spring areas and small creeks on gentle (moderate)
slopes or wet flatlands (Stellario nemorum-Abietetum
albae P. Kučera 2019);
329Український ботанічний журнал, 2019, 76(4)
(3) Valeriano dioicae-Abietion albae P. Kučera 2019
all. nov. with communities on base-rich minerotrophic
habitats – fens and other water-influenced habitats
(Valeriano dioicae-Abietetum P. Kučera 2019 ass. nov.).
Still problematic is the classification of the
associations Querco-Piceetum W. Matuszkiewicz
et Polakowska 1955, Betulo pubescentis-Piceetum
Sokołowski 1980 and Sphagno girgensohnii-Piceetum
Polakowski 1962 described from the northeastern
Poland as they come from a different phytogeographical
region of the outskirts of the boreal Picea abies
distribution range. This difference is expressed for
example by partial presence of Quercus robur, a species
exotic to communities of the aforementioned three
alliances. Especially the last mentioned association is
similar to Soldanello montanae-Piceetum Volk in Br.-Bl.
et al. 1939.
Scandinavian wet woodlands with Picea abies were
included by Kielland-Lund (1981, 1994) into the
separate suballiance Sphagno-Piceenion Kielland-Lund
1981 (association Chamaemoro-Piceetum Kielland-
Lund 1962). Although they are parallel to communities
of the alliance Sphagno palustris-Piceion P. Kučera
2019, they form a distinct phytogeographical group
of phytocoenoses (occurrence of the species Rubus
chamaemorus L., Carex vaginata, Linnaea borealis,
Calamagrostis purpurea (Trin.) Trin.). Therefore they are
here classified as a separate unit in the rank of alliance:
Sphagno girgensohnii-Piceion (Kielland-Lund 1981)
P. Kučera 2019 stat. nov. hoc loco
Raised name: Sphagno [girgensohnii]-Piceenion
Kielland-Lund 1981; Kielland-Lund (1981), p. 150;
cf. Art. 3g Example 2, Recomm. 10C.
II. B. Sphagno palustris-Piceion abietis P. Kučera all.
nov. hoc loco
Original diagnosis: Sphagno palustris-Piceetum
Šomšák 1979, Soldanello montanae-Piceetum Volk in
Br.-Bl. et al. 1939, Equiseto sylvaticae-Piceetum Šmarda
1950, Betulo pubescentis-Abietetum albae Lemée ex
Thébaud 2008, Leucobryo glauci-Piceetum abietis
Šomšák ex P. Kučera 2019.
Nomenclatural type: Sphagno palustris-Piceetum
Šomšák 1979; Šomšák (1979), p. 23–29, tab. 3,
holotypus hoc loco.
Differential species (φ (× 100) ≥ 25) (89 relevés,
Tab. 2):
E
3
: –,
E
2
: Picea abies (31),
E
1
: Betula pubescens (31), Salix aurita (28),
Vaccinium vitis-idaea (46), Carex canescens (36),
Ranunculus flammula (33), Juncus effusus (33),
Melampyrum sylvaticum (33), Agrostis canina (32),
Potentilla erecta (31), Agrostis stolonifera (31), Carex
echinata (30), Trientalis europaea (29), Lysimachia
vulgaris (29), Valeriana simplicifolia (26),
E
0
: Polytrichum commune (38), Lepidozia reptans
(31), Pohlia nutans (30), Sphagnum recurvum agg. (28),
Chiloscyphus pallescens (26), Herzogiella seligeri (25).
The alliance comprises the majority of known types of
wet woodlands with Picea abies (Tab. 4 – in Electronic
Supplement B2). From the view of natural altitudinal
vegetation zonation they mostly represent habitats
with the extragradal natural occurrence of Picea abies
(Kučera, 2019).
Picea abies is the canopy dominant in the wind
undisturbed stands, Abies alba was a natural component
of some communities being a competitor of Norway
spruce. Fagus sylvatica occurred occasionally on dryer
habitats. Depending on the type of habitat and the
succession stage other tree species could be admixed
(or temporarily dominating): Pinus sylvestris, Betula
pendula, B. pubescens, Alnus incana, A. glutinosa.
The common character of phytocoenoses is the
dominance of Polytrichum commune and Sphagnum
species (especially S. girgensohnii, S. centrale,
S. palustre), in seasonally dry habitats adjoining some
mires they are replaced by Leucobryum glaucum.
Dicranum scoparium and Pleurozium schreberi are their
frequent companions. Bazzania trilobata is considered
as the character species of the association Soldanello
montanae-Piceetum Volk in Br.-Bl. et al. 1939; however,
the species is not frequent in available relevé data from
Slovakia and it is frequently present also in the relevés
assigned to Equiseto sylvatici-Piceetum Šmarda 1950.
Field layers of all communities are dominated usually
by constant Vaccinium myrtillus and V. vitis-idaea,
less frequently accompanied by Calamagrostis villosa,
Oxalis acetosella and Equisetum sylvaticum; Dryopteris
carthusiana is frequently present. Species Luzula pilosa,
Athyrium filix-femina and Maianthemum bifolium for
the most part positively differentiate the group of
communities Leucobryo glauci-Piceetum abietis Šomšák
ex P. Kučera 2019, Sphagno palustris-Piceetum Šomšák
1979, Equiseto sylvatici-Piceetum Šmarda 1950 against
species-poor Soldanello montanae-Piceetum Volk in
Br.-Bl. et al. 1939.
330 Ukrainian Botanical Journal, 2019, 76(4)
Depending on a particular community/community
group, other following species/species groups have
differential value:
(1) Eriophorum vaginatum, Oxycoccus palustris,
(Carex nigra);
(2) Lycopodium annotinum;
(3) Caltha palustris, Lysimachia vulgaris, Crepis
paludosa, Deschampsia cespitosa;
(4) Agrostis canina (+ stolonifera), Potentilla erecta,
Viola palustris, Glyceria nemoralis (+ fluitans); Juncus
effusus, Ranunculus flammula.
The alliance Sphagno palustris-Piceion P.
Kučera 2019 also includes Calamagrostio villosae-
Pinetum Staszkiewicz 1958 (original form of
the name "Pineto-Calamagrostidetum villosae";
cf. W. Matuszkiewicz, 1981; and others) described by
Staszkiewicz (1958) from remarkable wet woodlands
of Nowy Targ surroundings (northerly of the Tatras),
eastwards of the Slovak-Polish state border dividing
the Orava region. A speciality of the recorded stands
(Staszkiewicz, 1958; Staszkiewicz, Szeląg, 2003) is the
dominance of Pinus sylvestris over (sometimes missing)
Picea abies (successional stage in part of localities?).
Species-poor composition with mostly constantly
present Carex nigra indicates presumable classification
of this community into Soldanello montanae-Piceetum
caricetosum fuscae Kasprowicz ex P. Kučera 2019.
The association Betulo pubescentis-Abietetum albae
Lemée ex Thébaud 2008 was described from France
(Thébaud 2006, p. 78) and documented from the
Massif Central (a mountain range outside the natural
occurrence of Picea abies) and from the Vosges (cf. Boeuf
et al. 2014). This plant community is syntaxonomically
close to Soldanello-Piceetum Volk in Br.-Bl. et al. 1939,
but its species composition reflects more nutrient
habitat conditions.
2. Soldanello montanae-Piceetum Volk in Br.-Bl.
et al. 1939
Original diagnosis: Braun-Blanquet et al. (1939),
p. 31–32.
Nomenclatural type: Petermann and Seibert (1979),
tab. 1, rel. b2 (Aufnahme Nr. 621), neotypus hoc loco.
Pseud.: Bazzanio-Piceetum (= Mastigobryo-
Piceetum) sensu auct. non (Schmid et Gaisberg 1936)
Br.-Bl. et Sissingh in Br.-Bl. et al. 1939.
Characteristic species combination: see Electronic
Supplement A2.
In respect of the number of recorded species this
association is a species-poor wet Picea woodland
without own "character species". However, the
community represents a distinct separate syntaxon
within the alliance Sphagno palustris-Piceion P.
Kučera 2019. It differs by (almost total) absence of
species frequently found in the remaining asociations
of the alliance: Luzula pilosa, Athyrium filix-femina,
Maianthemum bifolium, or Deschampsia cespitosa,
Caltha palustris, Crepis paludosa, Lysimachia vulgaris,
Senecio ovatus.
The canopy and understorey are dominated by
Picea abies, sporadically Betula pubescens and Pinus
sylvestris were recorded with higher cover-abundance
values. Recorded presence of Abies alba (cf. also
Braun-Blanquet et al., 1939; Trautmann, 1952, tab. 2),
especially in the field layer, as well as data of Kasprowicz
(1996) from the Polish part of Orava suggest that silver
fir occurrence in the stands was strongly reduced by
human influence. Even Fagus sylvatica grows here
marginally. Questionable is the possibility of the total
replacement of P. abies with A. alba in localities distant
from the continuous natural areal of Norway spruce.
The field layer is mostly dominated by
Vaccinium myrtillus, V. vitis-idaea is a constant
companion. Calamagrostis villosa and Oxalis acetosella
are sometimes admixed with higher cover, similarly also
Lycopodium annotinum, Homogyne alpina. Equisetum
sylvaticum is either absent or (sub-)dominant compound
of the phytocoenoses.
The name-giving species of the association,
Soldanella montana Willd., is known in Slovakia
undoubtly from only one locality in the Pieniny Mts only,
other localities are under consideration (Kochjarová
et al., 2016). In the relevés of the order Sphagno
palustris-Piceetalia P. Kučera 2019 from Slovakia, only
S. hungarica was recorded until present, mostly within
the association Sphagno palustris-Piceetum Šomšák
1979.
Sphagnum girgensohnii is mostly dominat species of
the ground layer, in records from Slovakia also
occasionally S. palustre agg. (incl. S. centrale). Numerous
other peat moss species were recorded. Constant species
are also Polytrichum commune, Dicranum scoparium and
Pleurozium schreberi; data on Polytrichum formosum
are partly doubtful (cf. Staszkiewicz, 1993). Bazzania
trilobata, a character liverwort species of this association
(Braun-Blanquet et al. 1939, Trautmann 1952), growing
abundantly in this community also in the Western
Carpathians (Bujakiewic, 1981; Kasprowicz, 1996;
Parusel, 2007) was recorded less frequently in Slovakia.
331Український ботанічний журнал, 2019, 76(4)
The association was recorded in Slovakia only at the
foothills southerly and northerly of the Tatras and in the
Babia Hora Mt. surroundings, one relevé is from the
Tichá Dolina in the Tatras (Kobzáková, 1987).
Although Sodanello montanae-Piceetum is a very
species-poor community, floristic variability allow
differentiation of the ecologically interpretable
subcommunities. Part of them were described under
the name Mastigobryo-Piceetum/Bazzanio-Piceetum
(Schmid et Gaisberg 1936) Br.-Bl. et Sissingh in
Br.-Bl. et al. 1939 due to inadequate identification
of Soldanello-Piceetum with Bazzanio-Piceetum.
On the other side, misidentification of Soldanello-
Piceetum with Lophozio-Piceetum Volk in Br.-Bl. et al.
1939 (see Oberdorfer, 1957) led to differentiation of
subassociation barbilophozietosum which actually does
not belong to Soldanello-Piceetum (see below). Selected
syntaxonomical notes on the association are provided in
Electronic Supplement A3, section II.
2a. Soldanello montanae-Piceetum sphagnetosum
recurvi (Trautmann 1952) P. Kučera 2019 comb. nov.
hoc loco
Basionym (in the original form of the name):
Mastigobryo-Piceetum (Schmid et Gaisberg 1936)
Br.-Bl. et Sissingh in Br.-Bl. et al. 1939 Subass. von
Sphagnum recurvum Trautmann 1952; Trautmann
(1952), p. 292.
Original diagnosis: Trautmann (1952), p. 292–293,
tab. 2, second column (Subassoziation von Sphagnum
recurvum).
Nomenclatural type: not established yet. Trautmann
(1952) published a synoptic table of relevés from
the Bavarian Forest, thus no relevé is available for
lectotypification in his work. Relevés from that region
and most probably identical with Trautmann's (1952)
data were published by Hartmann, Jahn (1967) in
tab. 3a; however, they were published under the name
"Mastigobryo-Piceetum, Subassoziation nach Carex
brizoides und Equisetum sylvaticum" nom. illeg. (Def. V),
thus cannot be used directly for neotypification
(cf. Art. 21).
The subassociation is characterized by intensified
presence of Carex brizoides and Equisetum sylvaticum
(Trautmann 1952) and by (mostly) absence of the
species characteristic for the other subassociations (e.g.
Lycopodium annotinum, Homogyne alpina, Oxycoccus
palustris).
Kasprowicz (1996, tab. 1, rel. 11) published a relevé
with dominance of Carex brizoides from the Polish part
of Orava, slightly differing by occurrence of Agrostis
stolonifera and Calamagrostis canescens. No data on the
syntaxon were published from Slovakia; however, stands
with Carex brizoides were observed in Orava flatlands
(Bernátová, Kučera, 2007–2008, not.).
This subassociation could be confused with the
association Carici brizoidis-Abietetum Trinajstić 1974.
Though, the latter unit represent stands of different
species-richer montane wet woodland (with Carex
remota) first described from the Dinarides (cf. Trinajstić,
1974).
The name-giving species of the subassociation –
Sphagnum recurvum P. Beauv. – has not been found
in Europe (Flatberg, 1992). Possible renaming of the
subassociation (Art. 43) (i.e. most probably S. fallax;
if the species identity was identified correctly within
S. recurvum agg.) should be based on reevaluation of
the original material of Trautmann (1952) and other
documented relevés of this subassociation from his
study area (the Bavarian Forest).
2b. Soldanello montanae-Piceetum homogynetosum
alpinae (Trautmann 1952) P. Kučera 2019 comb. nov.
hoc loco
Basionym (in the original form of the name):
Mastigobryo-Piceetum (Schmid et Gaisberg 1936)
Br.-Bl. et Sissingh in Br.-Bl. et al. 1939 Subass. von
Homogyne alpina Trautmann 1952; Trautmann (1952),
p. 292.
Original diagnosis: Trautmann (1952), p. 292–293,
tab. 2, third column (Subassoziation von Homogyne
alpina).
Nomenclatural type: not established yet. Trautmann
(1952) published a synoptic table of relevés from the
Bavarian Forest (Bavaria), thus no relevé is available
for lectotypification in his work. Similar relevés were
partly published by Neuhäuslová, Eltsova (2002a: tab.
1, 2002b: tab. 1.2) but the neotypification was not
accomplished because: (1) these relevés come from the
southern Bohemia (bordering to Bavaria) and (2) they
were published under the names "Soldanello-Piceetum"
resp. "Bazzanio-Piceetum typicum var. lycopodiosum" (cf.
Art. 21).
Syn.: Soldanello montanae-Piceetum equisetetosum
Oberdorfer 1957 nom. superfluum (Art. 29c).
Oberdorfer (1957, p. 382) transferred Trautmann's
(1952) relevés of the subassociation Mastigobryo-
Piceetum homogynetosum alpinae Trautmann 19521
1 Trautmann (1952) considered Mastigobryo-Piceetum and Soldanello-
Piceetum for synonyms.
332 Ukrainian Botanical Journal, 2019, 76(4)
(synoptic table) into the association Soldanello-
Piceetum, however without retaining the original epithet
(cf. Art. 26).
Data: Šoltés (1989): tab. 7a, rel. 2–4; Staszkiewicz
(1993): tab. 1, rel. 10–11; Šomšák et al. (1996): tab. 3,
rel. 1 + tab. 4, rel. 3; Kubíček et al. (1997a): tab. 1, rel. 1;
Vačko (2000), p. 60 (rel. 1).
Stands of the subassociation are usually recognizable
by presence (or dominance) of Homogyne alpina, Oxalis
acetosella, less frequent species Equisetum sylvaticum
and Lycopodium annotinum could be codominant.
Vaccinium myrtillus (dominant), V. vitis-idaea are
constantly present, but the latter species usually reaches
only small cover-abundance values (+, 1) in comparison
to the following subassociation. Dryopteris filix-mas,
Athyrium filix-femina, Hieracium murorum, Luzula
sylvatica and other more nutrient demanding species
usually positively differentiate this subunit against the
subassociation bazzanietosum (see below).
Sphagnum girgensohnii is the most frequently
dominating peat moss species documented in the
available relevés, sometimes accompanied (or altered)
especially by S. palustre and S. squarrosum in Slovakian
relevés. Other less constant mosses are Polytrichum
commune, Dicranum scoparium, Bazzania trilobata was
recorded sporadically in Slovakia.
This subcommunity was found on the southern
(Šoltés, 1989) and northern foothills of the Tatras
(Šomšák et al., 1996; Kubíček et al., 1997a; Vačko, 2000)
as well as in the Oravské Beskydy Mts (Staszkiewicz,
1993).
2c. Soldanello montanae-Piceetum bazzanietosum
trilobatae Petermann et Seibert ex P. Kučera 2019
subas. nov. hoc loco
Validated name: Soldanello montanae-Piceetum
bazzanietosum trilobatae Petermann et Seibert 1979,
nom. inval. (Art. 3o).
Original diagnosis: Petermann and Seibert (1979),
tab. 1.
Nomenclatural type: Petermann and Seibert (1979),
tab. 1, rel. a11 (Aufnahme Nr. 1209), holotypus hoc loco.
Data: Šoltés (1989): tab. 7a, rel. 5; Viceníková
(1991): tab. 4, rel. 5; Šomšák et al. (1996): p. 74 +
tab. 3, rel. 3–5; Kuderavá et al. (2000): p. 211, rel. 2.
This subassociation comprises the most species-
poor phytocoenoses of the association. More nutrient-
demanding species are absent, species composition of
the field layer often consists only of Vaccinium myrtillus,
V. vitis-idaea with less frequent Calamagrostis villosa,
Lycopodium annotinum (Dryopteris carthusiana, Oxalis
acetosella, Avenella flexuosa).
The ground layer is dominated by peat moss species,
in Slovakia S. girgensohnii and less frequently S. palustre
agg., further are present Polytrichum commune
(subdominant), Dicranum scoparium, Pleurozium
schreberi, Bazzania trilobata.
This subcommunity was found on the southern
(Šoltés, 1989; Viceníková, 1991) and northern foothills
of the Tatras (Šomšák et al., 1996) as well as in the
flatlands of Orava region (Kuderavá et al., 2000) where
it has wider distribution (Kučera, 2010, not.).
Only some smaller portion of the original relevés of
Soldanello montanae-Piceetum bazzanietosum trilobatae
published by Petermann, Seibert (1979, tab. 1) belong
to this unit because (1) the authors did not recognize
Soldanello montanae-Piceetum homogynetosum alpinae
(or even Equiseto-Piceetum Šmarda 1950, Petasito albi-
Piceetum Samek 1961) and part of the relevés represent
secondary communities of originally mixed woodlands
with Fagus sylvatica (similar to [1] Bazzanio-Piceetum
(Schmid et Gaisberg 1936) Br.-Bl. et Sissingh in Br.-
Bl. et al. 1939, or to [2] Luzulo nemorosae-Piceetum
(Schmid et Gaisberg 1936) Br.-Bl. et Sissingh in Br.-Bl.
et al. 1939; cf. Schmid, Gaisberg, 1936, tab. III; Kučera,
2009b, p. 27–28; Kučera, 2010, p. 834; Kučera, 2012,
p. 241–242).
2d. Soldanello montanae-Piceetum caricetosum fuscae
Kasprowicz ex P. Kučera 2019 subass. nov. hoc loco
Validated name and basionym: Bazzanio-Piceetum
caricetosum fuscae Kasprowicz 1996 nom. inval.
(Art. 5); Kasprowicz (1996), tab. 1, p. 150.
Original diagnosis: Kasprowicz (1996), tab. 1,
p. 150–152.
Nomenclatural type: Kasprowicz (1996), tab. 1,
rel. 2; lectotypus hoc loco.
Data: Bujakiewicz (1981): tab. 14, rel. 2; Kobzáková
(1987): p. 61; Staszkiewicz (1993): tab. 1, rel. 9;
Viceníková (1998): tab. 4, rel. 3, 6 + tab. 14, rel. 31;
Šomšák et al. (1996): tab. 3, rel. 2; Kubíček et al.
(1997b), tab. 1, rel. 3.
Transitional phytocoenoses of the association, often
on bogside ecotones, characterised by occurrence of
Carex nigra (cf. Kasprowicz, 1996) and/or Eriophorum
vaginatum, Oxycoccus palustris, occasionally also
Vaccinium uliginosum; marginal phytocoenoses with
only Carex nigra (Bujakiewicz, 1981; Kobzáková, 1987:
with C. pauciflora), C. echinata (Viceníková, 1998,
tab. 4, rel. 6) are probably to place here. When the
333Український ботанічний журнал, 2019, 76(4)
population of Pinus mugo (s. str.) is/was present at the
site, hybrid series of Pinus ×celakovskiorum Asch. &
Graebn. (= P. mugo × P. sylvestris; Businský, 1998) could
be present.
In Slovakia, Sphagnum girgensohnii was recorded
as the dominant of the ground layer, accompanied by
S. palustre agg., S. capillifolium. Frequently occurring
Polytrichum commune, Pleurozium schreberi, Dicranum
scoparium were recorded only sporadically.
This subcommunity was recorded in the Tatras
(Kobzáková, 1987) and on their southern (Viceníková,
1998) and northern foothills (Šomšák et al., 1996;
Kubíček et al., 1997b) as well as in the Oravské
Beskydy Mts and their foothills (Bujakiewicz, 1981;
Staszkiewicz, 1993).
Stands published under the name Calamagrostio
villosae-Pinetum Staszkiewicz 1958 eastwards of the
Slovak-Polish border presumably belong into this
subassociation, propably as a separate Pinus sylvestris
variant.
3. Carex rostrata-Picea abies community
With one relevé Šoltés (1989, tab. 7a, rel. 11)
documented a plant community with herb layer
dominants Calamagrostis villosa, Homogyne alpina
and Carex rostrata, whereas dominating mosses were
Polytrichum commune and Sphagnum girgensohnii.
Significant feature of the overall species composition is
lack of species specific for below presented associations
of the alliance, i.e. Leucobryo-Piceetum Šomšák ex
P. Kučera 2019, Sphagno palustris-Piceetum Šomšák
1979 and Equiseto-Piceetum Šmarda 1950, while at
the same time differing from subunits of Soldanello
montanae-Piceetum (see above and Tab. 2).
A somewhat similar community was published by
Zukrigl (1973, p. 152, tab. 6, rel. 1) within Sphagno
acutifolii-Piceetum Zukrigl1973; however, the
latter author documented a non-forest community
transitional towards the class Scheuchzerio-Caricetea
nigrae Tüxen 1937.
As the Carex rostrata-Picea wet woodland data
consist of only one relevé, this "group" was excluded
from fidelity calculations for associations presented in
this survey.
4. Leucobryo glauci-Piceetum abietis Šomšák ex
P. Kučera 2019
Original diagnosis: Kučera (2019), tab. 1, p. 28–31.
1 The correct cover-abundance value for Picea abies (E
3
) is "3" (cf.
cover value of tree layer in relevé = "40 %") (R. Šoltés, in e-mail).
Nomenclatural type: Kučera (2019), tab. 1, rel. 9,
holotype.
Characteristic species combination: see Electronic
Supplement A2.
Data: for published relevés see Kučera (2019, tab.
1) = Šomšák (1976): tab. 4, rel. 1, 2, 4; Ferančíková
(1994), tab. 1, rel. 2–6, tab. 3, rel. 4; Maťová (1994),
p. 44; Viceníková (1998), tab. 7, rel. 1–2, tab. 15, rel.
90–91; Novotková (1999), tab. 1, rel. 3.
The association Leucobryo glauci-Piceetum comprises
specific stands located on plains of dryer margins of
mires where water from snow melting and rains remains
considerable long above the soil surface (Šomšák, 1976;
Šomšák et al., 1993), though, growth of Picea abies
trees is not limited by water regime (Šomšák, 1976).
This wet woodland was formerly labelled by a nomen
fictum "Leucobryo-Piceetum Stefanović 1961" (see
Šomšák et al., 1993) in the theses of Šomšák (1976)
and his students (Feračíková, 1994; Maťová, 1994;
Viceníková, 1998) (see nomenclatural notes: Kučera,
2012, p. 247).
The floristic characteristics of the association
published Kučera (2019); however, the recorded overall
species composition, especially considering bryophytes,
is biased towards the field expertise of relevant author:
more detailed species records are given in the relevés
of Šomšák (1976; collaboration with J. Foltínová)
(cf. Kučera, 2019, tab. 1).
Stands of this plant community are characterized
by dominance of Picea abies; this species is also
a determining component of the understorey. Of
other trees a higher constancy is reached only by Pinus
sylvestris.
The most characteristic feature of phytocoenoses
of this association is the dominance of Leucobryum
glaucum in the ground layer. Also other bryophytes
are constant: Sphagnum spp. usually with the high
dominance (most frequently S. girgensohnii, S. palustre
agg.), Dicranum scoparium, Pleurozium schreberi, less
frequent Hylocomium splendens, Polytrichum commune
and others.
The field layer consists of stable association of
Vaccinium myrtillus, Luzula pilosa, V. vitis-idaea, Avenella
flexuosa, Calamagrostis arundinacea, Maianthemum
bifolium, Dryopteris carthusiana, Athyrium filix-femina,
Calamagrostis villosa as well as Equisetum sylvaticum
and Melampyrum sylvaticum.
Till present, Leucobryo glauci-Piceetum was found
only in the Popradská kotlina – in the glacifluvial terrain
southerly of the Tatras (Šomšák, 1976; Ferančíková,
334 Ukrainian Botanical Journal, 2019, 76(4)
1994; Maťová, 1994; Viceníková, 1998; Novotková,
1999).
Two subassociations were distinguished within the
unit (Kučera, 2019): typicum and agrostietosum caninae.
5. Sphagno palustris-Piceetum Šomšák 1979
Original diagnosis: Šomšák (1979), tab. 3, p. 26–28.
Nomenclatural type: Šomšák (1979), tab. 3, rel. 13,
holotype (ut "neotype").
Characteristic species combination: see Electronic
Supplement A2.
Data: Šomšák (1976): tab. 4, rel. 3; Šomšák (1979):
tab. 3, rel. 1–11, 13; Šomšák (1980): p. 20; Kontriš
(1981): p. 23, rel. above; Marková (1991): tab. 5,
rel. 1–3; Viceníková (1991): tab. 4, rel. 3; Kubíček
and Šomšák (1993): tab. 2, rel. 9; Ferančíková (1994):
p. 44 + tab. 3, rel. 1–3; Holotová (1994): tab. 3,
rel. 2; Maťová (1994): tab. 3, rel. 1–2 + tab. 4, rel.
1–2; Viceníková (1998): tab. 4, rel. 1–2, 4–5 + tab. 8,
rel. 1–3; Dítě (2003, not.): 1 relevé.
This wet Picea woodland is the most species-rich
association of the alliance Sphagno palustris-Piceion
P. Kučera 2019; however, the recorded overall species
composition, especially considering bryophytes, is
author biased (cf. Šomšák, 1979 vs. most of the later
relevés; see above). It is floristically and geographically
related to Eriophoro vaginati-Betuletum sensu Šomšák
1979 (Šomšák, 1979 and works of his students; Šomšák
et al., 1993) belonging to the class Vaccinio uliginosi-
Pinetea Passarge 1968.
The dominant tree is Picea abies, admixed are
Pinus sylvestris, Betula pubescens or Alnus incana
(A. glutinosa), sporadically B. pendula. Norway spruce
dominates also the understorey; however, more species
could participate in the species composition: Sorbus
aucuparia, Salix aurita, Frangula alnus and others.
The herb layer is usually dominated by combination
or one of the following species: Vaccinium myrtillus,
Equisetum sylvaticum, Calamagrostis villosa, Luzula
pilosa, higher cover-abundance values could be reached
by Caltha palustris subsp. laeta, Oxalis acetosella and
other species. More species have high constancy (see
above). The characteristic attribute of this association is
combination of species Agrostis canina (+ A. stolonifera),
Potentilla erecta, Viola palustris, Glyceria nemoralis
(+ G. fluitans), Juncus effusus, Ranunculus flammula,
Moneses uniflora, Ajuga reptans, Soldanella hungarica
and Carex nigra.
The ground layer is dominated by either Sphagnum
palustre agg. (centrale + palustre) or S. girgensohnii;
however also other peat moss species were recorded
(S. squarrosum, S. fallax, S. capillifolium etc.). Dicranum
scoparium is the constant species, accompanied by
Pleurozium schreberi and Polytrichum commune. In
the relevés of Šomšák (1979; collaboration with
J. Foltínová as bryophyte specialist), other moss species
reach relatively high constancy: Lepidozia reptans,
Hylocomium splendens, Plagiochila asplenioides,
Pohlia nutans, Plagiothecium curvifolium, Chiloscyphus
pallecens, Lophocolea bidentata, Sphagnum squarrosum,
S. fallax [ut S. recurvum].
Till the present, the association Sphagno palustris-
Piceetum Šomšák 1979 was recorded only in the
southern foothills of the Tatras, exceptionally on
foothills of the Babia hora Mt. in the northermost part
of Slovakia (Šomšák, 1980).
Exner (2007) included the unit Sphagno palustris-
Piceetum Šomšák 1979 into the association Equiseto-
Piceetum Šmarda 1950; however, the presented relevé
data (synoptic table) of the latter association from
Austria (Willner et al., 2007, tab. 39) do not indicate
representation of wet woodlands equivalent to Sphagno
palustris-Piceetum Šomšák 1979. Moreover, the
syntaxon bearing the name Sphagno palustris-Piceetum
Šomšák 1979 is not freely interchangeable with other
syntaxa with name combination "Sphagno-Piceetum“
(cf. Chytrý et al., 2013, p. 432) because of its distinct
original diagnosis.
6. Equiseto sylvatici-Piceetum Šmarda 1950
Original diagnosis: Šmarda (1950), p. 147–148.
Nomenclatural type: Šmarda (1950), p. 147, rel. 2,
lectotype; Jirásek (1996), p. 239.
Characteristic species combination: see Electronic
Supplement A2.
Data: Šomšák (1976): tab. 4, rel. 5; Šomšák (1979):
tab. 3, rel. 12; Marková (1991): tab. 4, rel. 3; + p. 51
(rel. 4); Viceníková (1991): p. 24 + tab. 4, rel. 2;
Holotová (1994): tab. 3, rel. 1 + p. 45; Maťová (1994):
tab. 4, rel. 3; Kučerová (1996): tab. 6, rel. 1–2; Šomšák
et al. (1996): tab. 2, rel. 1–2.
Association Equiseto sylvatici-Piceetum Šmarda 1950
represent a moderately species-rich wet woodland of the
alliance Sphagno palustris-Piceion P. Kučera 2019, with
a position between Soldanello montanae-Piceetum Volk
in Br.-Bl. et al. 1939 and Sphagno palustris-Piceetum
Šomšák 1979. Lectotypification by Jirásek (1996)
determined a clear differentiation of the unit from the
very species-poor wet woodlands of the association
335Український ботанічний журнал, 2019, 76(4)
Soldanello montanae-Piceetum Volk in Br.-Bl. et al. 1939
(cf. Braun-Blanquet et al., 1939; Trautmann, 1952).
The dominant tree species is again Picea abies in
the relevés recorded from Slovakia. Locally other
species could reach high cover in the canopy (partly the
influence of the past wind disturbance): Pinus sylvestris,
Betula pendula (B. pubescens), Alnus incana. P. abies
dominates also in lower vegetation layers, admixed are
A. incana, Sorbus aucuparia, Abies alba, Lonicera nigra.
Presence of S. aucuparia and A. alba support the idea
of the human-driven decline of silver fir at least in a
part of the stands (cf. Jirásek, 1996). Questionable is
the possibility of the total replacement of P. abies with
A. alba in localities distant from the continuous natural
areal of Norway spruce.
Constant and at the same time dominant species
of the field layer are Calamagrostis villosa, Equisetum
sylvaticum, Vaccinium myrtillus, sporadic (co-)do-
minants could be Oxalis acetosella, Veratrum album
subsp. lobelianum. Other constant species are
Vaccinium vitis-idaea, Luzula pilosa, Oxalis acetosella,
high frequency have also Dryopteris carthusiana,
Maianthemum bifolium, Lysimachia vulgaris, Athyrium
filix-femina.
The most frequent ground layer species are Dicranum
scoparium, Pleurozium schreberi, Polytrichum commune.
The prevailing part of relevés assigned here to this
association show only presence data for bryophytes.
Though, from the data on the total ground layer cover
and from the rest of the relevés it could be deduced that
the dominating species for almost every relevé is from
the genus Sphagnum. Most frequently recorded were
Sphagnum palustre agg. and S. girgensohnii, with the
decreasing constancy also: S. squarrosum, S. recurvum
agg., S. capillifolium and only once each S. russowii,
S. riparium, S. flexuosum and S. teres. Higher cover-
abundance values were also reached probably by
Polytrichum commune. Bazzania trilobata was recorded
in almost half of the relevés from Slovakia.
Association Equiseto sylvatici-Piceetum Šmarda 1950
was documented in Slovakia mostly in the southern
foothills of the Tatras, Šomšák et al. (1996) published
two records from the northern foothills of the Tatras.
Jirásek (1996, 2002) proposed differentiation of two
subassociations: deschampsietosum cespitosae (with two
of the total three original relevés of Šmarda (1950)) and
typicum (with two variants). Relevé data from Slovakia
do not reproduce the proposed division (cf. Jirásek,
1996, tab. 3).
II. C. Stellario nemorum-Abietion albae P. Kučera
2019 all. nov. hoc loco
Original diagnosis: Stellario nemorum-Abietetum
albae P. Kučera 2019 (Kučera 2019, tab. 2), Petasito
albi-Piceetum Samek 1961 (Samek 1961, p. 75, tab. II,
rel. 3, 5, 9, 11, 18), Carici remotae-Abietetum Husová
1998 (Husová, 1998, tab. 1).
Nomenclatural type: Stellario nemorum-Abietetum
albae P. Kučera 2019 (Kučera, 2019, p. 27–44, tab. 2),
holotypus hoc loco.
Differential species (φ (× 100) ≥ 25) (36 relevés,
Tab. 2):
E
3
: Abies alba (62), Fagus sylvatica (43)
E
2
: Fagus sylvatica (57), Sorbus aucuparia (37),
Lonicera xylosteum (34), Acer pseudoplatanus (31), Salix
caprea (29), Sambucus racemosa (25)
E
1
: Fagus sylvatica (33), Lonicera xylosteum (31),
Abies alba (25)
Stellaria nemorum (83), Chrysosplenium alternifolium
(69), Petasites albus (68), Lysimachia nemorum
(62), Geranium robertianum (55), Cardamine trifolia
(55), Adenostyles alliariae (55), Luzula luzulina
(52), Homogyne alpina (51), Impatiens noli-tangere
(51), Urtica dioica (50), Rubus hirtus (50), Gentiana
asclepiadea (49), Galium odoratum (48), Prenanthes
purpurea (46), Dryopteris dilatata (46), Phegopteris
connectilis (43), Milium effusum (43), Ranunculus
lanuginosus (40), R. platanifolius (40), Oxalis acetosella
(39), Senecio ovatus (37), Phyteuma spicatum (37),
Rubus idaeus (37), Chaerophyllum hirsutum (35), Luzula
sylvatica (35), Cardamine flexuosa (34), Calamagrostis
epigejos (34), Poa remota (34), Veronica anagallis-
aquatica (34), Equisetum sylvaticum (32), Dryopteris
filix-mas (32), Cicerbita alpina (31), Geum rivale (30),
Carex sylvatica (30), Calamagrostis arundinacea (29),
Deschampsia cespitosa (29), Ranunculus repens (25),
Doronicum austriacum (25).
E
0
: Plagiomnium affine (54), Cirriphyllum piliferum
(51), Plagiothecium undulatum (39), Plagiomnium
undulatum (33), Plagiothecium curvifolium (31),
Plagiomnium rostratum (31), Conocephalum conicum
(31), Thuidium tamariscinum (28).
This alliance unites wet woodland types considerably
differing from the communities of the alliance
Sphagno palustris-Piceion P. Kučera 2019. The species
composition (see Tabs 2–4 in Electronic Supplement
B2) and habitat type show its affinities to spatially
adjacent Fagus-Abies woodlands of the class Carpino-
Fagetea Jakucs ex Passarge 1968 while constant presence
of Equisetum sylvaticum (dominance in the association
336 Ukrainian Botanical Journal, 2019, 76(4)
Stellario nemorum-Abietetum albae P. Kučera 2019) –
together with Polytrichum commune, Sphagnum
girgensohnii, Deschampsia cespitosa – are attributes of
the order Sphagno palustris-Piceetalia P. Kučera 2019.
Group of hygrophiles Chaerophyllum hirsutum,
Chrysosplenium alternifolium, Impatiens noli-tangere,
Lysimachia nemorum, Stellaria nemorum, Petasites albus
growing together with species as Gentiana asclepiadea,
Prenanthes purpurea, Fragaria vesca, Plagiomnium
affine etc. (Kučera, 2019, tab. 2), and constantly
without Carex echinata, Lysimachia vulgaris, Trientalis
europaea (Vaccinium vitis-idaea), justify the separation
of the spring-related association Stellario nemo-
rum-Abietetum albae P. Kučera 2019 from the alliance
Sphagno palustris-Piceion Kučera 2019 into a separate
unit of the rank of alliance.
Association Petasito albi-Piceetum Samek 1961 (non
Petasito [albae?]-Piceetum Zupančič 1999 nom. illeg.
(Art. 31), or nom. inval. (Art. 3g)1) is a related unit to
Stellario-Abietetum P. Kučera 2019 (cf. Kučera, 2019)
and belongs to the alliance Stellario nemorum- Abietetum
albae P. Kučera 2019 as well. It was described from the
southern Bohemia (Czech Republic) by Samek (1961).
Petasito albi-Piceetum Samek 1961 was not recognized
or at least mentioned as a synonym neither in the
new vegetation survey of the Czech Republic (Chytrý
et al., 2013) nor in the older unfinished series edited
by J. Moravec (Jirásek, 2002). Records of Petasito albi-
Piceetum Samek 1961 are not known from Slovakia till
present.
Association Carici remotae-Abietetum Husová 1998
represents another wet woodland belonging to the
alliance Stellario nemorum-Abietion P. Kučera 2019.
However, only relevé data phytocoenotically close to
the nomenclatural type of the association should be
considered (cf. Husová 1998, tab. 1).
In the alliance should be included the association
Chaerophyllo hirsuti-Abietetum albae Boeuf et Simler
in Boeuf 2011 and probably also the phytocoenoses
delimited in the subassociation Carici pendulae-
Abietetum albae caricetosum brizoidis Boeuf 2011 nom.
inval. (Art. 3i) (cf. Boeuf, 2010). The subassociation
Carici pendulae-Abietetum albae typicum Boeuf 2011
nom. inval. (Art. 3i) most probably belongs to the water
influenced Abies alba woodlands of the class Carpino-
Fagetea Jakucs ex Passarge 1968.
1 The only place in the whole monograph of Zupančič (1999) where
the taxon name is specified is photo appendix (p. 221) in the end of
book. This unit syntaxonomically belongs to the class Carpino-Fagetea
Jakucs ex Passarge 1968.
7. Stellario nemorum-Abietetum albae P. Kučera 2019
Original diagnosis: Kučera (2019), tab. 2, p. 32–41.
Nomenclatural type: Kučera (2019), tab. 2, rel. 8,
holotypus
Syn.: Equiseto sylvatici-Abietetum sensu auct. slov.
non Moor 1952.
Characteristic species combination: see Electronic
Supplement A2.
Data for the respective subassociations:
calamagrostietosum – for published relevés see
Kučera (2019, tab. 2) = Majzlanová (1982): tab. 13,
rel. 15–21 and Majzlanová (s. d.): tab. 2, rel. 2, 3, 5, 10;
crepidetosum – for published relevés see Kučera
(2019, tab. 2) = Šomšák (1983): tab. 2, col. VI, rel. 1–4
and Majzlanová (s. d.): tab. 2, rel. 12; Šomšák et al.
(1996): tab. 4, rel. 1, 2; Kubíček et al. (1997b): tab. 1,
rel. 2; Kučera (2012): p. 295, rel. 31 + p. 296, rel. 32;
petasitetosum – for published relevés see Kučera
(2019, tab. 2) = Majzlanová (1982): tab. 13, rel. 1–14
and Majzlanová (s. d.): tab. 2, rel. 6.
The floristic composition of the association was
described by Kučera (2019): stands of the relevés
recorded until present are determined by Picea abies
and Abies alba. These two tree species are accompanied
by Fagus sylvatica, sometimes Sorbus aucuparia was
recorded. The understorey consists of the all four
mentioned species, the most constant shrub species are
Lonicea nigra and L. xylosteum.
The field layer is dominated by Oxalis acetosella
and Equisetum sylvaticum, higher cover-abundance
values are reached also by Stellaria nemorum, Petasites
albus, rarely Chaerophyllum hirsutum. Except Oxalis
and Equisetum, constant species are Senecio ovatus,
S. nemorum, Ch. hirsutum, Rubus idaeus, Chrysosplenium
alternifolium, Vaccinium myrtillus, Gentiana asclepiadea,
Athyrium filix-femina and Calamagrostis arundinacea.
The most abundant ground layer species is
Plagiomnium affine accompanied by Dicranum
scoparium and/or Sphagnum girgensohnii, the less
frequent bryophytes are Plagiothecium curvifolium,
Cirriphyllum piliferum and Polytrichum commune.
Stands of Stellario nemorum-Abietetum are bound
to habitat of spring areas and other more wet localities
(Majzlanová, 1983, 1993; Šomšák, 1983; Šomšák et al.,
1996). Until present, they were recorded prevailingly
in the flysch mountains of northern Slovakia: Oravská
Magura, Oravské Beskydy (Šomšák, 1983) and
Skorušinské vrchy (Majzlanová, 1982). A separate
variant of higher elevations was recorded in the Veľká
Fatra Mts at spring habitats on granodiorites (Kučera,
337Український ботанічний журнал, 2019, 76(4)
2012). Transitional phytocoenoses were found at the
northern foothills of the Tatras near Podspády (cf.
Šomšák et al., 1996, tab. 4, rel. 1, 2; Kubíček et al.,
1997b, tab. 1, rel. 2).
The association Stellario nemorum-Abietetum albae
P. Kučera 2019 is a newly described unit from Slovakia.
Its spatial distribution in the northern regions of the
country bordering to Poland suggests a high probability
of its occurrence in Poland as well as in the Carpathians’
mountain ranges of Moravia and Silesia, and also in
Ukraine (most probably also in Romania). It is possible
that some phytocoenoses from Switzerland included
by Kuoch (1954) into the subassociation Equiseto-
Abietetum hylocomietosum Kuoch 1954 belongs to the
alliance Stellario nemorum-Abietion albae P. Kučera
2019 or even to the association Stellario nemorum-
Abietetum P. Kučera 2019 (see below notes to Equiseto-
Abietetum).
Kučera (2019) differentiated three subassociations
within the association Stellario nemorum-Abietetum:
calamagrostietosum villosae Majzlanová ex P. Kučera
2019, crepidetosum paludosae P. Kučera 2019 and
petasitetosum albi Majzlanová ex P. Kučera 2019
II. D. Valeriano dioicae-Abietion albae P. Kučera
2019 all. nov. hoc loco
Original diagnosis: Equiseto sylvatici-Abietetum albae
Moor 1952 (Moor, 1952, p. 66–72, tab. 5), Valeriano
dioicae-Abietetum P. Kučera 2019 ass. nov.
Nomenclatural type: Equiseto sylvatici-Abietetum
albae Moor 1952 (Moor, 1952), holotypus hoc loco.
Characteristic species: Cirsium oleraceum, Filipendula
ulmaria, Fragaria vesca, Rubus saxatilis, Valeriana
dioica.
Differential species (derived from the relevé group of
this alliance from Slovakia) (φ (× 100) ≥ 25) (13 relevés,
Tab. 2):
E
3
: Pinus sylvestris (47), Alnus glutinosa (38),
A. incana (38), Betula pendula (37),
E
2
: Frangula alnus (49), Viburnum opulus (41),
Lonicera nigra (37),
E
1
: Daphne mezereum (48), Lonicera nigra (43),
Viburnum opulus (41), Sorbus aucuparia (40), Frangula
alnus (33), Betula pendula (31), Ribes petraeum (31),
Rubus saxatilis (93), Valeriana dioica (79),
Polygonatum verticillatum (70), Caltha palustris (67),
Crepis paludosa (65), Luzula pilosa (59), Cirsium
oleraceum (54), Clematis alpina (54), Maianthemum
bifolium (51), Filipendula ulmaria (50), Thalictrum
aquilegiifolium (48), Carex alba (48), Galium schultesii
(44), Fragaria vesca (42), Solidago virgaurea (41), Melica
nutans (41), Astrantia major (41), Paris quadrifolia (40),
Equisetum palustre (40), Dactylorhiza maculata (agg.)
(39), Carex digitata (33), Actaea spicata (33), Carex
remota (33), Epipactis palustris (31), Bistorta major (29),
Polygonatum multiflorum (28), Valeriana tripteris (28),
Angelica sylvestris (26),
E
0
: Trichocolea tomentella (39), Eurhynchium
angustirete (37), Tetraphis pellucida (34).
This alliance comprises wet woodland types with
Abies alba and Picea abies (and probably constant
presence of Fagus sylvatica) on base-rich habitats which
determine the presence of species group Valeriana
dioica, Cirsium oleraceum, Filipendula ulmaria, with
Equisetum sylvaticum, Caltha palustris, Crepis paludosa,
Deschampsia cespitosa along with calciphilous plants as
Aconitum vulparia, Carex alba, C. flacca, Kuoch (1954)
recorded also Bellidiastrum michelii and Calamagrostis
varia. Overall floristic composition of the alliance is
poorly known as this type of communities was never
properly recognized.
Tree species composition could also contain Fraxinus
excelsior, Acer pseudoplatanus, Sorbus aria (Moor,
1952), Alnus incana (+ glutinosa) and other species
in various proportion, depending on the successional
stage of a particular stand (wind and other type of
natural disturbance). Shrub species Lonicera nigra,
L. xylosteum, Viburnum spp. are present.
It is questionable if Picea abies could extinct
locally on the habitats under consideration during the
Holocene due to interspecific competition, especially
in the mountain ranges without development of the
natural Picea altitudinal vegetation zone and other
types of refugia. However, artificially forced expansion
of Norway spruce (plantations) could result in the
reintroduction of this species to localities where it
would be naturally absent.
The association Equiseto sylvatici-Abietetum
originally described by Moor (1952) and classified
within the alliance Valeriano dioicae-Abietion albae P.
Kučera 2019 has a distinctive set of constant species
Equisetum sylvaticum, Cirsium oleraceum, Crepis
paludosa, Deschampsia cespitosa, Lysimachia nemorum,
Carex flacca, Hordelymus europaeus, C. sylvatica,
Primula elatior, Knautia maxima along with other less
constant species Aconitum vulparia, Valeriana dioica,
Caltha palustris, Filipendula ulmaria, V. officinalis,
Equisetum arvense, Athyrium filix-femina, Fragaria vesca
etc.; with constant presence of Fagus sylvatica in the
canopy.
338 Ukrainian Botanical Journal, 2019, 76(4)
Relevés from Slovakia assigned to this alliance do not
strictly reproduce the described floristic characteristics
of Equiseto sylvatici-Abietetum Moor 1952. Although
the Slovak phytocoenoses are similar to the latter unit
with their species composition, e.g. by occurrence of
Caltha palustris, Cirsium oleraceum, Crepis paludosa,
Valeriana dioica, they lack species Aconitum vulparia,
Carex flacca, Geranium sylvaticum, Hordelymus
europaeus, Lonicera alpigena, Primula elatior, Salix
appendiculata. The combination of the constant species
Valeriana dioica, Crepis paludosa, Caltha palustris with
Cirsium oleraceum, Filipendula ulmaria constitute a
specific relevé group including at the same time species
Thalictrum aquilegiifolium, Clematis alpina, Equisetum
sylvaticum, Chaerophyllum hirsutum, Rubus saxatilis or
Polygonatum verticillatum; partly the Carex alba group
(see below).
Although it is clear that some of the recorded relevés
document transitional phytocoenoses (e.g. Viceníková,
1991, tab. 4, rel. 1: with Viola palustris, Molinia caerulea,
Poa palustris, Carex flava, Anemone nemorosa etc.), this
group as an unit differs noticeably from all of the above
described associations. Taking into account certain lack
of knowledge about the plant community corresponding
to Moor (1952, tab. 5) Equiseto-Abietetum, presented
Slovak relevés are included into the new association
Valeriano dioicae-Abietetum P. Kučera 2019 ass. nov.
For nomenclatural purposes a lectotypification of the
association Equiseto sylvatici-Abietetum is here given:
Equiseto sylvatici-Abietetum Moor 1952
Original diagnosis: Moor (1952), tab. 5.
Nomenclatural type: Moor (1952), tab. 5, rel. 1,
lectotypus hoc loco.
Nomenclatural and syntaxonomical note to the
association is given in Electronic Supplement A3,
section III.
8. Valeriano dioicae-Abietetum P. Kučera 2019 ass.
nov. hoc loco
Original diagnosis: Šomšák (1979, relevé on the
p. 25), the below specified nomenclatural type and
Kučera (in prep.; most of the included relevés are found
in various unpublished theses).
Nomenclatural type: Marková (1991), tab. 4, rel. 1
(unpublished master's thesis), locality: Western
Carpathians, Popradská kotlina Basin, near Tatranská
Kotlina, forest stand 1356a, altitude 760 m a.s.l., 6. 8.
1989.
E
3
(cover 80%): Abies alba 2, Picea abies 2;
E
0
(cover 5%): Lonicera nigra 1;
E
1
(cover 95%): Abies alba 2, Picea abies 2, Lonicera
nigra 1, Padus avium +, Sorbus aucuparia +,
Equisetum sylvaticum 3, Oxalis acetosella 3,
Maianthemum bifolium 2, Rubus saxatilis 2, Valeriana
tripteris 2, Caltha palustris subsp. laeta 1, Hieracium
murorum 1, Luzula pilosa 1, Polygonatum verticillatum
1, Rubus idaeus 1, Senecio ovatus 1, Solidago virgaurea
1, Vaccinium myrtillus 1, Ajuga reptans r, Athyrium
filix-femina +, Astrantia major +, Carex alba +, Clematis
alpina +, Dryopteris carthusiana +, D. filix-mas +,
Fragaria vesca +, Galium schultesii +, Gymnocarpium
dryopteris +, Melica nutans r, Mycelis muralis +,
Valeriana dioica +, Calamagrostis arundinacea r, Cirsium
oleraceum r, Crepis paludosa r, Dentaria glandulosa r,
Filipendula ulmaria r, Geum rivale r, Orthilia secunda r,
Polygonatum multiflorum r,
E
0
(cover 45 %, only presence of the species is noted):
Dicranum scoparium, Plagiomnium affine, Polytrichum
formosum.
Characteristic species combination: see Electronic
Supplement A2.
Data: Šomšák (1979): p. 25; Marková (1991): tab.
4, rel. 1–2, 4–5; Viceníková (1991): tab. 4, rel. 1, 4, 6
+ tab. 5, rel. 1–2; Viceníková (1998): tab. 14, rel. 30,
44, 45.
Documented relevés show domination of Picea abies
in the canopy; Abies alba is codominant in a small part
of relevés. I assume that the latter species was constantly
present (and dominant) in all of the documented stands
which come from the foothills of the Belianske Tatry
Mts and close vicinity, and that Fagus sylvatica was also
present there (cf. Kučera, 2008b, 2009). High frequence
of Pinus sylvestris is partly the result of the historical
human influence on the forest in the region. Other tree
species were documented less frequently (Alnus incana
+ glutinosa, Betula pubescens + pendula). Most frequent
young trees are Picea abies, Sorbus aucuparia and Abies
alba, from shrubs Lonicera nigra. Daphne mezereum,
Viburnum opulus, Rosa pendulina, even Ribes petraeum,
are sporadic.
A set of constant herb layer species comprise
Crepis paludosa, Caltha palustris, Vaccinium myrtillus,
Luzula pilosa, Maianthemum bifolium, Rubus saxatilis,
Equisetum sylvaticum, Valeriana doica and Oxalis
acetosella; overall species composition is rich.
Two subunits are differentiated here, at present in the
rank of variant regarding the lack of knowledge of the
overall variability of this plant community:
(1) Carex alba variant (4 relevés): with exclusive
species group Carex alba (constant), Astrantia major,
339Український ботанічний журнал, 2019, 76(4)
Galium schultesii, Melica nutans, Actea spicata,
Mycelis muralis, Valeriana tripteris, Carex sylvatica,
with other species concentrated here – Dryopteris filix
mas, Prenanthes purpurea, Gymnocarpium dryopteris,
Hieracium murorum. The role of Sphagnum subsp. is
insufficiently known.
(2) Calamagrostis villosa variant (9 relevés): with
Vaccinium vitis-idaea and Calamagrostis villosa
(exceptionally also in the previous variant), less
frequent (sporadic) species Epilobium montanum,
Lysimachia vulgaris, Potentilla erecta, Huperzia selago,
Avenella flexuosa, Dactylorhiza maculata agg. Peat moss
species (Sphagnum girgensohnii, S. palustre agg., or
others) could reach high cover-abundance values. Also
Bazzania trilobata was recorded in the stands of this
variant.
Characterization of the ground layer is difficult:
except one relevé of Šomšák (1979, p. 25), the rest
of total 13 relevés come from theses whose authors
recorded only presence data of bryophytes. Constant is
only Dicranum scoparium, other more frequent species
are Pleurozium schreberi, Rhytidiadelphus triquetrus,
Plagiochila asplenioides and Polytrichum formosum.
From the cover data of the ground layer in the relevés
could be expected that species of genus Sphagnum
(S. palustre agg., S. girgensohnii, occassionally
S. capillifolium, S. squarrosum) dominate in the stands of
Calamagrostis villosa-variant of this unit (with Bazzania
trilobata present infrequently).
Relevés of this wet woodland were recorded only on
the south-eastern foothils of the easternmost part of the
Tatras – the carbonate Belianske Tatry Mts as well as in
the close adjacent area to the south-west (surroundings
of Kežmarské Žľaby).
Conclusions
The presented phytocoenological revision of the
wet woodlands with Picea abies in Slovakia revealed
that diversity of the respective plant communities
(6 associations and 1 community) is comparable to
other Norway spruce-dominated communities which
are syntaxonomically classified within the orders
Piceetalia excelsae Pawłowski ex Pawłowski et al. 1928
and Cortuso-Piceetalia (Athyrio-Piceetalia sensu auct.
non Hadač 1962) of the supramontane altitudinal
vegetation zone of the Western Carpathians.
Except for one association (Sphagno acutifolii-
Piceetum Zukrigl 1973), the wet woodlands under
consideration form a specific floristic and ecological
unit of azonal Picea abies (and Abies alba) wet woodlands
distributed in the mostly lower altitudes of the montane
Fagus-Abies zone, here classified within the newly
proposed order Sphagno palustris-Piceetalia P. Kučera
2019. Plant communities of this order distinguished
from the available Slovak relevé data are according
to the distinct floristic differences splitted into three
alliances: Sphagno palustris-Piceion P. Kučera 2019,
Stellario nemorum-Abietion albae P. Kučera 2019 and
Valeriano dioicae-Abietion albae P. Kučera 2019.
Surveys from other European countries (e.g. Moor,
1952; Kielland-Lund 1981; Husová, 1998; Exner, 2007;
Chytrý et al., 2013; Boeuf et al., 2014;) indicate that the
proposed syntaxonomical system of the order Sphagno
palustris-Piceetalia P. Kučera 2019 is applicable for the
whole European region.
Acknowledgements
I would like to thank anonymous reviewers for their
comments on the first version of the manuscript, and R. Bouef
(Brumath), V. Hayova (Kyiv) and one of the reviewers for their
help with literature. This study was partially supported by the
Slovak grant agency VEGA, project No. 2/0119/19.
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Recommended for publication by D.V. Dubyna
e1
Electronic Supplement to: Kučera. Syntaxonomical classification of wet woodlands with Picea abies in Slovakia
Ukrainian Botanical Journal, 2019, 76(4)
Supplement A1: Detailed description of methods and the resulting dendrogram
The initial set of phytocoenological relevés of wet woodlands with Picea abies (especially Sphagnum-rich wet
woodlands) was prepared using the Turboveg for Windows database software (Hennekens, 2016; cf. Hennekens,
Schaminée, 2001) from the dataset provided for the prepared monograph Plant communities of Slovakia, Forest and
shrub vegetation (Valachovič et al., in prep.) stored in Centrálna databáza… (2016).
Subsequently, the following data were deleted: duplicitous relevés, non-forest relevés, relevés of Alnus incana
and A. glutinosa phytocoenoses (class Alnetea glutinosae Br.-Bl. et Tx. ex Westhoff et al. 1946), relevés belonging
to (krummholz-)forest peatland communities of the class Vaccinio uliginosi-Pinetea sylvestris Passarge 1968
(stable communities with open canopy cover and Oxycoccus palustris agg., Vaccinium uliginosum, Ledum palustre,
Eriophorum vaginatum etc.) as well as Picea abies-Sphagnum spp./Pinus cembra-Sphagnum spp. relevés classified
as climax woodlands of the class Piceetea excelsae Klika 1948 (cf. Kučera, 2012, 2017). Finally, the dataset was
actualised with missing relevés of Šomšák (1976, 1979, 1983) (cf. Kučera, 2019). For purposes of this paper all
available relevés irrespective of their plot size are used due to small number of recorded relevés.
A subfinal dataset of 153 relevés was exported for further modifications in the JUICE software package (Tichý,
2016; cf. Tichý, 2002). Using JUICE, taxa with unequal taxonomic rank were merged to the nearest mutual rank
(Caltha palustris – C. palustris subsp. laeta; Cardamine amara – C. amara subsp. opicii; Galeobdolon luteum agg. –
G. luteum – G. montanum; Myosotis palustris agg. – M. scorpioides) as well as Sphagnum palustre and S. centrale as
the latter species was not recognized in older works. Other Sphagnum species were retained, including infrequent
data on "Sphagnum sp." for preservation of the information on the Sphagnum taxa presence.
For the statistical comparison of higher syntaxa (Tab. 1) other merged taxa were set: Aconitum firmum s. l.,
Alchemilla spp., Campanula rotundifolia agg., Cardaminospis arenosa agg., Chiloscyphus polyanthos s. l., Cladonia
squamosa (incl. var.), Crepis jacquinii, Luzula luzuloides, Pellia spp., Primula elatior (incl. subsp. tatrensis), Senecio
nemorensis agg., Soldanella hungarica (incl. "montana", subsp. hungarica), Solidago virgaurea (incl. subsp. minuta),
Sorbus aucuparia (incl. subsp. glabrata).
Data on taxa of the Dryoteris carthusiana group were retained for the purpose of maintaining the important
floristic information for higher syntaxa comparison: undetermined "Dryopteris carthusiana agg." occurred only two
times in the whole dataset. Similarly, other species were not merged with very sporadic data on the undetermined
taxa: Aconitum sp. (1×), Glyceria sp. (1×), Juncus sp. (1×), Lepidozia sp. (1×), Myosotis sp. (1×), Polytrichum sp.
(1×), Potentilla sp. (2×), Salix sp. (E
1
, 5×),* Soldanella sp. (4×), Sphagnum sp. (10×).
The subfinal dataset was further analysed within JUICE and basic types of communities were identified.
Consequently, relevés which represent secondary Norway spruce stands or transitional phytocoenoses to A. glutinosa
communities were excluded (8 relevés: Majzlanová, 1982, tab. 13, rel. 22; Majzlanová, s. d., tab. 2, rel. 4, 7, 8, 9;
Kobzáková, 1987, p. 64; Viceníková, 1991, tab. 5, rel. 3; Olekšák, 1995, tab. 6). Thus, the final dataset of wet Picea
woodlands counted 145 relevés.
Statistical analysis of the final dataset was performed by the software package SYN-TAX 2000 (Podani, 2001a).
Hierarchical clustering was executed using the coefficient Podani's discordance (see more Podani, 2001b), using
also variants without E
3
+E
2
species and also without E
0
species (i.e. with E
1
species only). However, the final relevé
classification follows a manual rearrangement of several relevés to reach more compact species delimitation of
basic units (associations) (fig. 1). The main reason for this modification is a supposed author bias (see the next four
paragraphs and fig. 1):
– Although relevé groups from the opposite parts of diversity range (Sphagno acutifolii-Piceetum, Solda-
nello-Piceetum, (Leucobryo-Piceetum) vs. Stellario-Abietetum, Equiseto-Abietetum[-Carex alba subunit]) were
constantly or with only insignificant variation reproduced as distinctive separate units within the performed variants
of the numerical classification, the most of remaining relevés (Sphagno palustris-Piceetum, Equiseto-Piceetum) were
vaguely grouped.
* E
3
= canopy (trees), E
2
= understorey (shrubs), E
1
= field layer (herbs, grasses, dwarf shrubs etc.), E
0
= ground layer
(bryophytes, lichens) (Klika, 1948, pp. 29–30; Rodwell et al., 1991).
e2
– Detailed examination of those relevés revealed that their species composition is biased towards the respective
author. The most species-rich relevés – irrespective of their final classification here – come from the studies of the
recognized Slovak geobotanist Prof L. Šomšák (1976, 1979) in collaboration with Dr J. Foltínová as the bryophyte
specialist. The reported species richness applies especially to bryophytes: no other author recorded similarly species-
abundant relevés within the particular unit.
– In contrast, Šomšák's students recorded in general (markedly) less abundant species composition in their
unpublished theses. Moreover, relevé data on bryophytes were frequently recorded as the presence data only. The
author bias is probably also displayed in marked alternation of species Agrostis canina/A. stolonifera and Glyceria
nemoralis/G. fluitans in the respective relevés (missidentification?). Determination of Sphagnum species and their
recognition in the field could also be a source of inaccuracy.
– Thus, the species-rich relevés of Leucobryum glaucum-dominated wet Picea woodland (author Šomšák; see
Kučera 2019) were usually classified outside of the corresponding group. Similarly, species less abundant relevés of
the group defined by set of species unique to community Sphagno palustris-Piceetum described by Šomšák (1979)
were partially mixed with the relevés classified here as Equiseto-Piceetum.
The differential attributes of the respective syntaxa (fidelity and frequency values, rounded to units) and resulting
tables were elaborated within JUICE (Tichý, 2016); the concept of fidelity was used (Chytrý et al., 2002; phi
coefficient – φ). The characteristics of associations are derived from final dataset without one relevé "group" of the
Carex rostrata-Picea abies community (see below). Fidelity calculation was based on the presence/absence data
with a standardization of relevé groups to an equal size. Performing the Fisher's exact test, zero fidelity was given to
species with significance P > 0.05 in a particular cluster (Tichý, Chytrý, 2006).
Fidelity values in the descriptions of the units of wet Picea woodland alliances of the order Sphagno palustris-
Piceetalia (units 2–7, see Tab. 4 – in electronical appendix B2) are computed from the final dataset reduced
by another 6 relevés of the supramontane community belonging to the alliance Piceion excelsae Pawłowski ex
Pawłowski et al. 1928 (see below).
e3
Figure 1. Dendrogram of hierarchical clustering of the relevés of wet Picea communities from Slovakia
1 – Sphagno acutifolii-Piceetum Zukrigl 1973
2 – Soldanello montanae-Piceetum Volk in Br.-Bl. et al. 1939
3 – Carex rostrata-Picea abies community (1 relevé)
4 – Leucobryo glauci-Piceetum abietis Šomšák ex P. Kučera 2019
5 – centre of relevés of Sphagno palustris-Piceetum Šomšák 1979
6 – centre of relevés of Equiseto sylvatici-Piceetum Šmarda 1950 (with admixed relevés of Sphagno palustris-Piceetum)
7 – Stellario nemorum-Abietetum albae P. Kučera 2019
8 – Valeriano dioicae-Abietetum P. Kučera 2019 ass. nov.
e4
1. Sphagno acutifolii-Piceetum Zukrigl 1973
Characteristic species combination within the whole dataset of wet woodlands with Picea abies in Slovakia (6
relevés, Tab. 3):
A) differential species (φ (× 100) ≥ 25):
E
3
: Sorbus aucuparia (41),
E
2
: Pinus mugo (55),
E
1
: Pinus mugo (55),
Juncus filiformis (87), Eriophorum vaginatum (81), Dryopteris dilatata (72), Athyrium distentifolium (72), Dryopteris
expansa (66), Nardus stricta (63), Homogyne alpina (57), Carex nigra (50), Carex canescens (38), Avenella flexuosa
(38),
E
0
: Sphagnum capillifolium (58), S. rubellum (55), Lophocolea heterophylla (43), Polytrichum formosum (41),
Calypogeia azurea (40), Dicranum fuscescens (38), Polytrichum alpinum (38), Barbilophozia floerkei (38), Sphagnum
fuscum (38), Barbilophozia attenuata (38), Pleuridium subulatum (38), Polytrichum commune (37);
B) frequent species (constancy ≥ 50%):
E
3
: Picea abies,
E
2
: Pinus mugo (50),
E
1
: Picea abies (67), Sorbus aucuparia (67),
Vaccinium myrtillus (100), Homogyne alpina (100), Avenella flexuosa (83), Vaccinium vitis-idaea (83), Eriophorum
vaginatum (83), Calamagrostis villosa (83), Dryopteris dilatata (83), Juncus filiformis (83), Athyrium distentifolium
(67), Carex canescens (67), Nardus stricta (67), Carex nigra (67), Dryopteris expansa 50),
E
0
: Polytrichum commune (100), Polytrichum formosum (83), Sphagnum capillifolium (83), Dicranum scoparium
(83), S. girgensohnii (67), Plagiothecium curvifolium (67).
2. Soldanello montanae-Piceetum Volk in Br.-Bl. et al. 1939
Characteristic species combination (24 relevés, Tab. 4 – in the electronical appedix B2):
A) differential species (φ (× 100) ≥ 25):
E
3
: –,
E
2
: Pinus sylvetris (27),
E
1
: Eriophorum vaginatum (38), Lycopodium annotinum (34), Listera cordata (28), Thelypteris palustris (27),
E
0
: Sphagnum girgensohnii (25), Polytrichum commune (25);
B) constant species (constancy ≥ 40%):
E
3
: Picea abies (100),
E
2
: Picea abies (96),
E
1
: Picea abies (92),
Vaccinium myrtillus (100), V. vitis-idaea (100), Calamagrostis villosa (63), Oxalis acetosella (50), Equisetum
sylvaticum (42), Homogyne alpina (42), Dryopteris carhusiana (42),
E
0
: Sphagnum girgensohnii (83), Polytrichum commune (75), Dicranum scoparium (67), Pleurozium schreberi (54),
S. palustre agg. (50).
4. Leucobryo glauci-Piceetum abietis Šomšák ex P. Kučera 2019
Characteristic species combination (15 relevés, Tab. 4 – in the electronical appedix B2):
A) differential species (φ (× 100) ≥ 25):
E
3
: –,
E
2
: –,
E
1
: Melampyrum sylvaticum (48), Calamagrostis arundinacea (32), Avenella flexuosa (30), Luzula luzuloides (30),
Calluna vulgaris (29), Luzula pilosa (28),
E
0
: Leucobryum glaucum (83), Orthodicranum undulatum (34), Hylocomium splendens (29), Dicranella heteromalla
(27);
B) frequent species (constancy ≥ 50%):
E
3
: Picea abies (100),
Supplement A2: Formal characteristics of the distinguished associations
e5
E
2
: Picea abies (93),
E
1
: Picea abies (93), Sorbus aucuparia (60),
Vaccinium myrtillus (100), Luzula pilosa (93), V. vitis-idaea (93), Avenella flexuosa (67), Athyrium filix-femina
(67), Calamagrostis arundinacea (67), Dryopteris carthusiana (67), Maianthemum bifolium (67), Calamagrostis villosa
(60), Equisetum sylvaticum (60),
E
0
: Leucobryum glaucum (100), Dicranum scoparium (93), [Sphagnum spp. (80)], Pleurozium schreberi (80),
Hylocomium splendens (53).
5. Sphagno palustris-Piceetum Šomšák 1979
Characteristic species combination (37 relevés, Tab. 4 – in the electronical appedix B2):
A) differential species (φ (× 100) ≥ 25):
E
3
: –,
E
2
: –,
E
1
: Salix aurita (39), Corylus avellana (26),
Agrostis canina (58), Viola palustris (53), Ranunculus flammula (49), Juncus effusus (43), Potentilla erecta (41),
Agrostis stolonifera (39), Carex rostrata (37), Valeriana simplicifolia (35), Carex canescens (33), Ajuga reptans (31),
Carex pallescens (30), Moneses uniflora (28), Senecio nemorensis agg. (ut S. nemorensis) (28), Carex echinata (27),
Lysimachia vulgaris (27), Peucedanum palustre (26), Galium uliginosum (26), Melampyrum pratense (26), Galium
palustre (26),
E
0
: Brachythecium starkei (30), Lepidozia reptans (28), Sphagnum quinquefarium (26), Cephalozia bicuspidata
(26), Calliergon cordifolium (26), Rhodobryum roseum (26), Chiloscyphus pallescens (26), Sphagnum palustre agg.
(25);
B) frequent species (constancy ≥ 50%):
E
3
: Picea abies (100),
E
2
: Picea abies (95),
E
1
: Picea abies (89), Sorbus aucuparia (54),
Vaccinium myrtillus (97), V. vitis-idaea (92), Equisetum sylvaticum (86), Luzula pilosa (86), Calamagrostis villosa
(81), Dryopteris carthusiana (76), Athyrium filix-femina (73), Maianthemum bifolium (68), Potentilla erecta (65),
Caltha palustris (54), Deschampsia cespitosa (54), Agrostis canina (54) [A. canina + A. stolonifera (86)], Crepis
paludosa (54), Lysimachia vulgaris (54), Orthilia secunda (51), Oxalis acetosella (51),
E
0
: Dicranum scoparium (81), Sphagnum palustre agg. (70), S. girgensohnii (65), Pleurozium schreberi (65),
Polytrichum commune (57).
6. Equiseto sylvatici-Piceetum Šmarda 1950
Characteristic species combination (13 relevés, Tab. 4 – in the electronical appedix B2):
A) differential species (φ (× 100) ≥ 25):
E
3
: –,
E
2
: Alnus incana (39),
E
1
: Lonicera nigra (26),
Trientalis europaea (44), Lysimachia vulgaris (35), Calamagrostis villosa (28), Veratrum album subsp. lobelianum
(25),
E
0
: Lophocolea bidentata (40), Bazzania trilobata (25);
B) frequent species (constancy ≥ 50%):
E
3
: Picea abies (100),
E
2
: Picea abies (92),
E
1
: Picea abies (100), Sorbus aucuparia (62),
Calamagrostis villosa (100), Equisetum sylvaticum (100), Vaccinium myrtillus (100), V. vitis-idaea (92), Luzula
pilosa (85), Oxalis acetosella (85), Dryopteris carthusiana (77), Maianthemum bifolium (77), Lysimachia vulgaris
(62), Athyrium filix-femina (62), Crepis paludosa (54), Caltha palustris (54), Rubus idaeus (54),
E
0
: Dicranum scoparium 92), Pleurozium schreberi (69), Plagiothecium curvifolium (62).
e6
7. Stellario nemorum-Abietetum albae P. Kučera 2019
Characteristic species combination (36 relevés, Tab. 4 – in the electronical appedix B2):
A) differential species (φ (× 100) ≥ 25):
E
3
: Abies alba (63), Fagus sylvatica (47),
E
2
: Fagus sylvatica (61), Lonicera xylosteum (38), Sorbus aucuparia (35), Acer pseudoplatanus (34), Lonicera nigra
(26), Salix caprea (25),
E
1
: Lonicera xylosteum (34), Fagus sylvatica (29), Abies alba (29),
Stellaria nemorum (84), Chrysosplenium alternifolium (72), Petasites albus (71), Lysimachia nemorum (65),
Geranium robertianum (59), Cardamine trifolia (59), Adenostyles alliariae (59), Luzula luzulina (57), Impatiens noli-
tangere (57), Prenanthes purpurea (56), Gentiana asclepiadea (55), Rubus hirtus (54), Galium odoratum (52), Dryopteris
dilatata (47), Milium effusum (47), Chaerophyllum hirsutum (45), Urtica dioica (44), Ranunculus lanuginosus (44),
R. platanifolius (44), Phyteuma spicatum (41), Senecio ovatus (40), Geum rivale (39), Carex sylvatica (39), Cardamine
flexuosa (38), Poa remota (38), Calamagrostis epigejos (38), Veronica anagallis-aquatica (38), Homogyne alpina (37),
Phegopteris connectilis (37), Cicerbita alpina (34), Rubus idaeus (34), Dryopteris filix-mas (32), Luzula sylvatica (31),
Oxalis acetosella (31), Epilobium montanum (29), Symphytum tuberosum (27), Sanicula europaea (27), Calamagrostis
arundinacea (27), Cardamine amara (26), Poa palustris (26),
E
0
: Cirriphyllum piliferum (54), Plagiomnium affine (47), Plagiothecium undulatum (38), Conocephalum conicum
(34), Plagiomnium rostratum (34), Plagiomnium undulatum (31), Thuidium tamariscinum (31);
B) frequent species (constancy ≥ 50%):
E
3
: Picea abies (100), Abies alba (78),
E
2
: Picea abies (64), Sorbus aucuparia (56),
E
1
: Picea abies (78), Abies alba (72), Sorbus aucuparia (56),
Equisetum sylvaticum (100), Oxalis acetosella (100), Senecio ovatus (83), Rubus idaeus (83), Stellaria nemorum
(78), Chaerophyllum hirsutum (78), Vaccinium myrtillus (75), Athyrium filix-femina (75), Chrysosplenium alternifolium
(69), Gentiana asclepiadea (67), Petasites albus (67), Prenanthes purpurea (61), Calamagrostis arundinacea (61),
C. villosa (61), Dryopteris carthusiana (58), Homogyne alpina (58), Hieracium murorum (53), Dryopteris filix-mas
(53), Deschampsia cespitosa (53), Lysimachia nemorum (50),
E
0
: Plagiomnium affine (72), Dicranum scoparium (61), Sphagnum girgensohnii (53).
8. Valeriano dioicae-Abietetum P. Kučera 2019 ass. nov.
Characteristic species combination (13 relevés from Slovakia, Tab. 4 – in the electronical appedix B2):
A) differential species (φ (× 100) ≥ 25):
E
3
: Pinus sylvestris (28), Alnus incana (26), A. glutinosa (26),
E
2
: Viburnum opulus (45), Lonicera nigra (45), Frangula alnus (36),
E
1
: Daphne mezereum (55), Viburnum opulus (45), Lonicera nigra (34), Ribes petraeum (32), Sorbus aucuparia
(32), Abies alba (26),
Rubus saxatilis (94), Valeriana dioica (74), Polygonatum verticillatum (71), Clematis alpina (59), Cirsium
oleraceum (59), Filipendula ulmaria (55), Thalictrum aquilegiifolium (52), Carex alba (52), Crepis paludosa (51),
Caltha palustris (50), Galium schultesii (49), Fragaria vesca (48), Astrantia major (45), Melica nutans (45), Solidago
virgaurea (39), Chaerophyllum hirsutum (37), Dactylorhiza maculata (agg.) (36), Carex remota (36), Actaea spicata
(36), Carex digitata (36), Maianthemum bifolium (36), Equisetum palustre (35), Luzula pilosa (34), Epipactis palustris
(32), Polygonatum multiflorum (32), Valeriana tripteris (32), Paris quadrifolia (31), Angelica sylvestris (29), Senecio
ovatus (27), Bistorta major (25),
E
0
: Trichocolea tomentella (39), Eurhynchium angustirete (28), Tetraphis pellucida (26);
B) frequent species (constancy ≥ 50%):
E
3
: Picea abies (100), Pinus sylvestris (62),
E
2
: Picea abies (69), Lonicera nigra (62),
E
1
: Picea abies (100), Sorbus aucuparia (92), Abies alba (69), Lonicera nigra (54),
Crepis paludosa (100), Caltha palustris (100), Vaccinium myrtillus (100), Luzula pilosa (100), Maianthemum
bifolium (100), Rubus saxatilis (92), Equisetum sylvaticum (77), Valeriana dioica (77), Oxalis acetosella (77), V. vitis-
idaea (69), Chaerophyllum hirsutum (69), Senecio ovatus (69), Calamagrostis villosa (69), Athyrium filix-femina (69),
Polygonatum verticillatum (69), Solidago virgaurea (62), Filipendula ulmaria (54), Rubus idaeus (54), Fragaria vesca
(54),
E
0
: Dicranum scoparium (85), Pleurozium schreberi (69), Sphagnum girgensohnii (62).
e7
Section I) Nomenclatural and syntaxonomical notes to the association Sphagno acutifolii-Piceetum Zukrigl 1973
The association was described by Zukrigl (1973) under the name "Sphagno-Piceetum", thus creating a name
similar to earlier and also later descriptions of Sphagnum-Picea abies woodland units (cf. Exner, 2007; Kučera, 2012;
Chytrý et al., 2013). The relevé table 6 of Zukrigl (1973) comprises in a matter of fact four different phytocoenoses
with varying peat moss species participation:
– rel. 1 is a non-forest community on a montane bog with height-reduced trees (canopy cover 10 %, understorey
cover 20 %) of the class Scheuchzerio-Caricetea nigrae Tüxen 1937;
– rel. 2 shows an example of phytocoenosis with elements of both climax supramontane Picea forest and bog
communities;
– rel. 3 presents a different (semiforest?) phytocoenosis of the bog rand with Molinia;
– rels 4–6 represent a phytocoenosis similar to Soldanello montanae-Piceetum Volk in Br.-Bl. et al. 1939 (probably
subass. caricetosum fuscae Kasprowicz ex P. Kučera 2019).
Typification of the association by Willner, Zukrigl (1999) considerably consolidated the syntaxonomical content
of the syntaxon as a good interpretable and useful phytocoenological unit. As only one peat mos species was
mentioned in the type relevé, the association name could be specified as "Sphagno acutifolii-Piceetum" (Recommend.
10C; Weber et al., 2000, p. 749).
Remarks on the name type "Sphagno-Piceetum"
Sphagno-Piceetum (Tüxen 1937) Hartmann 1953
There is a striking difference between the characteristics of the association by Hartmann (1953) (= "Moor-
Fichtenwald") and the supplied original diagnosis which consists of the subassociation Piceetum excelsae
sphagnetosum Tüxen 1937 (= humid slope forest phytocoenoses with numerous Sphagnum-pillows) (Willner,
2007; Kučera 2007, 2012). The name Sphagno-Piceetum (Tüxen 1937) Hartmann 1953 (often with various author
citations) was repeatedly used in the Hartmann's incorrect concept until recently (e.g. Jirásek, 2002); however,
Hartmann later changed his syntaxonomical evaluation and did not use the label Sphagno-Piceetum (cf. Hartmann,
Jahn, 1967).
Tüxen (1937) presented three peat moss species in his original diagnose in the form of a synoptic table consisting
of total 8 relevés, all of them were labelled as differential species: Sphagnum capillifolium (ut S. acutifolium Russ. et
Warnst.) in 4 relevés, S. quinqefarium in 2 relevés and S. girgensohnii in 2 relevés. According to the differences in
ecology of these peat moss species (Pilous 1971) and at the same time in compliance with Tüxen (1937), I propose
the following completion of Tüxen's name following the Code (Weber et al., 2000, Rec. 10C):
Piceetum excelsae sphagnetosum quinquefarii Tüxen 1937; see Tüxen (1937), p. 123.
Consequently, the association name published by Hartmann (1953) is to be completed as follows:
Sphagno quinquefarii-Piceetum (Tüxen 1937) Hartmann 1953; see Hartmann (1953), Anhang, p. XIII.
Sphagno-Piceetum Kuoch 1954 (with one subassociation thuidietosum)
The name is a nomen superfluum (Willner 2007, p. 240; Art. 29c) because Kuoch (1954, p. 227) proposed the
name as a unit including also communities Soldanello-Piceetum Volk in Br.-Bl. et al. 1939 as well as Bazzanio-
Piceetum (Schmid et Gaisberg 1936) Br.-Bl. et Sissingh in Br.-Bl. et al. 1939 (cf. Kučera, 2007). Provided relevés
(Kuoch, 1954, tab. 13) actually display affinities to Soldanello-Piceetum Volk in Br.-Bl. et al. 1939 and part of them
also towards Equiseto-Piceetum Šmarda 1950.
Kuoch (1954, p. 227, also Tab. 13) in express words stated that S. girgensohnii is a provisional character species
of the association. Also other peat moss species are recorded in Kuoch's relevés. As the name is nomen superfluum,
completion of the name according the Art. 10 (Rec. 10C) is redundant.
Supplement A3: Nomenclatural and syntaxonomical notes to selected syntaxa
e8
Sphagno-Piceetum Richard 1961
Sphagno-Piceetum Richard 1961 is cited by Šomšák (1979) and Chytrý et al. (2013) as a separate association
name. However, Richard (1961, p. 110) stated that he labelled the unit "according to Moor and Kuoch" and
described two new subassociations. Thus, the author actually used the name Sphagno-Piceetum Kuoch 1954 and did
not describe a new association, therefore also completion of this name to "Sphagno girgensohnii-Piceetum (Moor
1942) Richard 1961" (cf. Art. 10C) made by Boeuf et al. (2014, p. 111) was redundant.
Species composition of Richard's relevés are in accordance with concept of Kuoch (1954) and the relevés of his
two subassociations belong to the association Soldanello-Piceetum Volk in Br.-Bl. et al. 1939 (see above) and partly
to Equiseto-Piceetum Šmarda 1950.
Sphagno girgensohnii-Piceetum Polakowski 1962 nom. cons. propos.
The association was described with the original form of the name Piceo-Sphagnetum Girgensohnii by Polakowski
(1962) from the north-east Poland where outskirts of boreal Picea abies distribution range extend. The correct
inverted form of the name (Art. 10b) was proposed already by Czerwiński (1966) and was used since then
(Medwecka-Kornaś, 1972; J. Matuszkiewicz, 1977; Sokołowski, 1980) and at the same time is recognized as a
distinct syntaxon in the national surveys till the recent time (cf. W. Matuszkiewicz, 1964; J. Matuszkiewicz, 1977,
2002; W. Matuszkiewicz 1981 till 2014; W. Matuszkiewicz, J. Matuszkiewicz, 1996).
The long and well established use of this unit is the reason for preservation of the name and for preferential
conservation against a potential older homonymous name. A competing name could be Sphagno-Piceetum (Tüxen
1937) Hartmann 1953; however, completion of the latter name (see above) to Sphagno quinquefarii-Piceetum (Tüxen
1937) Hartmann 1953 (see above) solve that question.
For nomenclatural purposes, lectotypes are here chosen for the association Sphagno girgensohnii-Piceetum
Polakowski 1962 and its subunits:
Sphagno girgensohnii-Piceetum Polakowski 1962 nom. cons. propos.
Nomenclatural type: Polakowski (1962), tab. 4, rel. 29; lectotypus hoc loco.
Sphagno girgensohnii-Piceetum lycopodietosum annotini Polakowski 1962
Nomenclatural type: Polakowski (1962), tab. 4, rel. 29; lectotypus hoc loco.
Sphagno girgensohnii-Piceetum vaccinietosum myrtilli Polakowski 1962 (Art. 14)
Nomenclatural type: Polakowski (1962), tab. 4, rel. 54; lectotypus hoc loco.
Sphagno-Piceetum Zukrigl 1973
Zukrigl (1973) described a new association. The name could be completed to Sphagno acutifolii-Piceetum Zukrigl
1973 (see above), thus it is questionable to label it as a later homonymum (cf. Chytrý et al., 2013, p. 432).
Sphagno-Piceetum Ellenberg et Klötzli 1974
Ellenberg, Klötzli (1972) (on the correct publication date see Willner 2007, p. 239) actually differentiated
two separate communities: Sphagno-Piceetum typicum Ellenberg et Klötzli 1972 and Sphagno-Piceetum
calamagrostietosum villosae Ellenberg et Klötzli 1972. However, from the nomenclatural point of view the first
subassociation is a nomen superfluum (Art. 29c) to Sphagno-Piceetum betuletosum Richard 1961 in the first
succession. Similarly, Sphagno-Piceetum calamagrostietosum villosae should be nomenclaturally treated as a nomen
superfluum to Sphagno-Piceetum thuidietosum Kuoch 1954 even though the supposed syntaxonomical content was
different.
In both cases, Sphagno-Piceetum Ellenberg et Klötzli 1974 leads nomenclaturally directly to Soldanello-Piceetum
Volk in Br.-Bl. et al. 1939 (see above).
Sphagno palustris-Piceetum Šomšák 1979
Along with the description of a new association Šomšák (1979) specified the name-giving Sphagnum species,
thus the unit was clearly differentiated from older Sphagnum-Picea names. It represents distinct wet woodland with
Picea abies.
e9
Sphagno-Piceetum sensu Sofron 1981 non (Tüxen 1937) Hartmann 1953
This is the most common example of the approach when an author used the concept of bog woodland intended by
Hartmann (1953), but the nomenclatural implications were not followed (see above). Application of the incorrect
use of the association name continued more decades (cf. Jirásek, 1996, 2002 vs. Chytrý et al., 2013).
Sphagno acutifolii-Piceetum (Březina et Hadač in Hadač et al. 1969) Hadač 1987
Hadač (1987) published the nomen novum for Sphagno-Piceetum excelsae tatricum (Art. 34) and at the same time
specified the peat moss species used in the association name. Nevertheless, Hadač’s name is a younger homonym to
Sphagno acutifolii-Piceetum Zukrigl 1973 (Art. 31).
Only few relevés of this community was published from Slovakia. According to my field knowledge, it represents
strongly human-influenced Picea-Abies forests on habitat where Fagus sylvatica was originally naturally present
(Kučera, 2009a, 2012, p. 250).
Sphagno magellanici-Piceetum Bick ex Boeuf in Boeuf et al. 2014
Boeuf (Boeuf et al. 2014, p. 109; cf. also p. 306, 316) proposed this new name since the name of Bick (1952;
Pino rotundatae-Sphagnetum piceetosum Bick 1985) was considered invalidly published due to the absence of the
required typification (Art. 3o etc.). However, Bick (1985, p. 166) published his subassociation name validly (Art. 2b,
5) therefore Boeuf's proposal should be assessed as raising of the rank (Art. 27d).
Proposal to complete the association name with the epiteth "magellanici" (cf. Recom. 10C) is also problematic
because the original diagnosis cited by Bick (1985), i.e. facies "Pinetum uncinatae piceosum" by Kästner, Flößner
(1933, tab. XXII), does not contain Sphagnum magellanicum (table 18 of Bick contains this species).
Moreover, the name Sphagno magellanici-Piceetum (proposed by Boeuf) would be a later homonym to the name
Piceo abietis-Sphagnetum magellanici Krisai 1986 (Art. 31d) (cf. Wallnöfer 1993, p. 309). While the subassociation
Pino rotundatae-Sphagnetum piceetosum abietis Bick 1985 (cf. Art. 10b) represents a plant community of the class
Vaccinio uliginosi-Pinetea Passarge 1968, the name Piceo abietis-Sphagnetum magellanici Krisai 1986 belongs to
a plant community of non-forest vegetation of the class Oxycocco-Sphagnetea Br.-Bl. et Tx. ex Westhoff et al. 1946.
Section II) Chosen syntaxonomical notes to the association Soldanello montanae-Piceetum Volk in Br.-Bl. et al.
1939
Although the original of the association Soldanello montanae-Piceetum Volk in Br.-Bl. et al. 1939 published by
Braun-Blanquet et al. (1939, p. 32) was very distinctive and explicit, the unit was repeatedly confused with other
syntaxa in the past (cf. also Sofron, 1981, p. 56–60). In brief:
(1) Trautmann (1952) united this association with Mastigobryo-Piceetum (= Bazzanio-Piceetum) (Schmid et
Gaisberg 1936) Br.-Bl. et Sissingh in Br.-Bl. et al. 1939 from the Black Forest [Schwarzwald], probably following
the occurrence pattern of Bazzania trilobata (cf. Braun-Blanquet et al. 1939). Although Oberdorfer (1957) formally
retained the label Bazzanio-Piceetum, it seems that the determining factor of the later application of the name
(cf. Wallnöfer, 1993) was simultaneous presentation of the species Bazzania trilobata along with the name
Bazzanio-Piceetum.
Most of the authors in the former Czechoslovakia and in Poland followed the concept of assignment of stands
with Bazzania trilobata to the association Bazzanio-Piceetum (Schmid et Gaisberg 1936) Br.-Bl. et Sissingh in
Br.-Bl. et al. 1939 (auctorum (excl. Zlatník, p. 63 in Ružička, 1961); Samek, 1961; Magic, 1966, 1986;
Neuhäuslová-Novotná, 1968; Šomšák, 1979; Bujakiewicz, 1981; and other later authors in the respective country).
However, the association Bazzanio-Piceetum (Schmid et Gaisberg 1936) Br.-Bl. et Sissingh in Br.-Bl. et al. 1939
is defined by the subassociations Piceetum myrtilletosum and Piceetum vaccinietosum both described by Schmid, von
Gaisberg (1936) from which only the second one marginally contains wet Picea woodlands (cf. Braun-Blanquet
et al., 1939; Bartsch, Bartsch, 1940; Kučera, 2007, p. 62, 2010, p. 834, 2012, p. 242). In contrast to these units,
e10
association Soldanello montanae-Piceetum Volk in Br.-Bl. et al. 1939 represents true wet woodland (Braun-Blanquet
et al. 1939, Trautmann 1952, tab. 2).
(2) The second (and in the eastern Central Europe only sporadical) source of confusion is Oberdorfer's (1957)
integration of the supramontane Lophozio-Piceetum Volk in Br.-Bl. et al. 1939 into Soldanello montanae-Piceetum
Volk in Br.-Bl. et al. 1939 (cf. also Willner, 2007, p. 240) expressed by Soldanella montana Willd. as proposed
character species of the latter association. These two communities co-occurre in the Bohemian Forest but they are
floristically and ecologically distinct units.
Similarly Samek (1961) in the former Czechoslovakia misidentified Soldanello-Piceetum with a supramontane
Picea woodland of the (Czech part of) Bohemian Forest labelled by him Homogyno-Piceetum Samek 1961 (= syntax.
syn. of Lophozio-Piceetum Volk in Br.-Bl. et al. 1939). Šoltés (1969) applied Volk's name Soldanello-Piceetum to
Western Carpathians’ supramontane Picea woodlands; however, later he used only the name Vaccinio myrtilli-
Piceetum Šoltés 1976 (Šoltés 1976).
Concept of fusion of Soldanello montanae-Piceetum and Lophozio-Piceetum was preserved by German and
Austrian phytocoenologists (see Seibert, 1992; Wallnöfer, 1993); however, Exner (2007, p. 205–207) correctly
separated corresponding wet woodland (subass. Soldanello-Piceetum equisetetosum Oberdorfer 1957 nom. inval.,
Art. 3m = Mastigobryo-Piceetum homogynetosum alpinae Trautmann 1952, Art. 14) from the climax supramontane
woodland.
(3) Lately Exner (2007) unified "Soldanello montanae-Piceetum s. str." (= homogynetosum Trautmann 1952; ut
S.-P. equisetetosum Oberdorfer 1957) with Equiseto-Piceetum Šmarda 1950. Again, these two units represent two
separate associations which was demonstrated also by Chytrý et al. (2013); proposal of conservation of the name
Equiseto-Piceetum against Soldanello montanae-Piceetum (Willner, 2007, p. 240) is therefore controversial.
Section III) Nomenclatural and syntaxonomical note to the association Equiseto sylvatici-Abietetum Moor 1952
Although Willner (2007, p. 239) lectotypified the association with the name "Equiseto-Abietetum Moor ex Kuoch
1954" (see the notes below), the lectotype was chosen from Kuoch's (1954) relevés and not from the original data
for the association already provided by Moor (1952). Such manner of lectotype choice should be rejected and
a corresponding rule should be implemented into International Code of Phytosociological Nomenclature (cf. latest
edition: Weber et al., 2000).
In the recent vegetation surveys of the surrounding countries there are no mentions of the association Equiseto
sylvatici-Abietetum Moor 1952 except for Exner (2007). However, the used association concept might not
be corresponding to the original description of the syntaxon by Moor (1952) as Exner (2007) included in this
association several units including Petasito-Piceetum Samek 1961 which is floristically and ecologically non-related
unit (see this paper and Samek, 1961).
Moreover, Exner (2007) used the name "Equiseto-Abietetum Moor ex Kuoch 1954" indicating the suggestion that
Kuoch (1954) validated (Art. 3b) the name proposed by Moor (1952). It should be emphasized that Moor (1952)
only in the p. 68 gave provisional statement on the association name, though, he clearly adopted the name Equiseto-
Abietetum throughout his whole study and in the relevé table as well as he did not use the expressions "nom. prov."
or "ass. prov." (cf. Art 3b, Weber et al. 2000, p. 745). Nor was the name treated by Kuoch (1954) as provisionally
described by Moor as the original author.
I propose that such clear adoption of a name by the original author should be accepted, which was explained
already in the case of the name Aceri pseudoplatani-Fagetum J. Bartsch et M. Bartsch 1940 (Kučera, 2007, 2013,
p. 23).
It is possible that a part of Kuoch's (1954) relevés classified to the association Equiseto sylvatici-Abietetum Moor
1952 really belong to this unit. Data on species as Calamagrostis varia and Bellidiastrum michelii indicate different
phytocoenological quality which is here classified as a separate syntaxon in the rank of an alliance – Valeriano
dioicae-Abietion albae P. Kučera 2019. More extensive examination of phytocoenoses resembling Moor's (1952)
description of Equiseto sylvatici-Abietetum Moor 1952, relevés of Kuoch (1954, tab. 8) and the presented data
from Slovakia included, would facilitate establishment of floristic patterns for division of phytocoenoses between
the base-rich alliance Valeriano dioicae-Abietion albae P. Kučera 2019 and the less nutrient-rich alliance Stellario
nemorum-Abietion albae P. Kučera 2019.
e11
Supplement B1: Table 1. Differential tables of orders of the class Piceetea excelsae Klika 1948 with fidelity
(φ (× 100) ≥ 25) and constancy (%) in the exponent
A – Sphagno palustris-Piceetalia abietis P. Kučera 2019
B – Piceetalia excelsae Pawłowski ex Pawłowski et al. 1928
C – Cortuso matthioli-Piceetalia abietis P. Kučera nom. prov. (= Athyrio-Piceetalia sensu auct. non Hadač 1962)
Species with constancy less than 10 % in a single column are omitted.
Group A B C
No. of relevés 145 234 97
Differential tree and shrub species
E
3
Pinus sylvestris 4225 –. –.
Abies alba 3825 –. –3
Alnus incana 3618 –. –.
Betula pendula 3114 –. –.
Alnus glutinosa 2912 –. –.
Betula pubescens 2718 –1 –4
Pinus cembra –. 4642 –10
Sorbus aucuparia –4 731 2844
Acer pseudoplatanus –. –. 2811
E
2
Picea abies 4883 –40 –24
Alnus incana 2912 –. –.
Frangula alnus 2811 –. –.
Ribes petraeum –. –1 3721
Salix silesiaca –. –3 2916
E
1
Abies alba 4739 –2 –4
Picea abies 4388 –51 –33
Alnus incana 2912 –. –.
Frangula alnus 2710 –. –.
Salix aurita 2610 –. –.
Sorbus aucuparia –54 2988 –64
Daphne mezereum –3 –. 6049
Ribes petraeum –2 –4 4031
Salix silesiaca –1 –2 2614
Other tree and shrub species
E
3
Picea abies –100 –99 –98
E
2
Sorbus aucuparia –22 –21 1233
Pinus mugo –1 1922 –15
Lonicera nigra 2219 –1 –8
Fagus sylvatica 2011 –1 –3
E
1
Lonicera nigra –17 –12 2435
e12
Group A B C
No. of relevés 145 234 97
Pinus cembra –. 2117 –8
Acer pseudoplatanus –8 –1 2218
Betula pubescens 2012 –. –4
Rosa pendulina –4 –1 1710
Differential field layer species (E
1
)
Equisetum sylvaticum 8277 –1 –.
Luzula pilosa 6352 –2 –.
Caltha palustris 5237 –. –1
Deschampsia cespitosa 4536 –2 –4
Dryopteris carthusiana 4359 –15 –18
Potentilla erecta 4326 –. –.
Lysimachia vulgaris 4224 –. –.
Maianthemum bifolium 3952 –4 –27
Myosotis palustris agg. 3825 –1 –2
Carex echinata 3822 –1 –.
Carex canescens 3723 –3 –.
Ranunculus repens 3421 –. –3
Agrostis canina 3316 –. –.
Orthilia secunda 3225 –. –7
Vaccinium vitis-idaea 3274 –49 –30
Lysimachia nemorum 3013 –. –.
Athyrium filix-femina 3057 –22 –31
Carex nigra 3015 –1 –.
Glyceria nemoralis 2912 –. –.
Agrostis stolonifera 2912 –. –.
Veronica officinalis 2817 –1 –3
Viola palustris 2811 –. –.
Filipendula ulmaria 2710 –. –.
Impatiens noli-tangere 2612 –. –1
Juncus effusus 2610 –. –.
Cardamine trifolia 2610 –. –.
Ranunculus flammula 2610 –. –.
Valeriana dioica 2610 –. –.
Petasites albus 2518 –1 –6
Dryopteris dilatata –12 5379 –36
Avenella flexuosa –38 4488 –44
Homogyne alpina –32 3488 –74
Vaccinium myrtillus –93 2599 –71
Valeriana tripteris –2 –3 7571
Primula elatior –2 –1 6657
Cortusa matthioli –. –. 6046
Phyteuma spicatum –5 –2 5954
Myosotis sylvatica –3 –. 5948
Polygonatum verticillatum –9 –6 5962
e13
Group A B C
No. of relevés 145 234 97
Cirsium erisithales –. –. 5640
Calamagrostis varia –. –. 5438
Astrantia major –2 –1 5341
Asplenium viride –. –. 5337
Galeobdolon luteum agg. –3 –2 5243
Polystichum lonchitis –. –. 5236
Geranium sylvaticum –. –1 5237
Heracleum sphondylium –. –. 5135
Viola biflora –1 –2 5039
Leucanthemum rotundifolium –1 –1 5036
Bellidiastrum michelii –. –. 4528
Cardaminopsis arenosa agg. –. –. 4528
Clematis alpina –3 –. 4432
Galium schultesii –3 –1 4433
Mercurialis perennis –. –. 4326
Cicerbita alpina –3 –11 4343
Mycelis muralis –12 –1 4341
Sesleria albicans –. –. 4225
Tanacetum clusii –1 –. 4024
Adenostyles alliariae –10 442 4067
Hieracium murorum –30 –7 3957
Soldanella hungarica –4 –17 3944
Luzula sylvatica –8 2367 3877
Moneses uniflora –5 –1 3728
Ranunculus platanifolius –6 –3 3731
Lilium martagon –. –1 3720
Prenanthes purpurea –19 –29 3761
Pimpinella major –. –. 3619
Fragaria vesca –13 –. 3634
Senecio subalpinus –1 –1 3420
Senecio nemorensis agg. –42 –36 3475
Aconitum variegatum –. –. 3416
Dentaria enneaphyllos –. –. 3416
Epilobium montanum –13 –2 3434
Soldanella carpatica –. –8 3427
Thalictrum aquilegiifolium –3 –. 3421
Cystopteris fragilis –. –2 3420
Campanula rotundifolia agg. –. –2 3319
Phyteuma orbiculare –. –. 3315
Alchemilla spp. –2 –1 3219
Carduus glaucinus –. –. 3214
Cystopteris montana –. –. 3214
Crepis jacquini –. –. 3113
Sesleria tatrae –. –. 3113
e14
Group A B C
No. of relevés 145 234 97
Carex digitata –1 –. 3015
Ranunculus oreophilus –. –. 2912
Poa alpina –. –. 2912
Carex sempervirens subsp. laxiflora (Schur) Jáv. –. –. 2912
Doronicum austriacum –3 –9 2827
Campanula cochleariifolia –. –. 2811
Veratrum album subsp. lobelianum –16 –17 2741
Tofieldia calyculata –. –. 2710
Dryopteris filix-mas –23 –9 2640
Campanula serrata –. –1 2610
Ranunculus lanuginosus –6 –1 2618
Poa stiriaca –. –. 259
Carex ornithopoda –. –. 259
Corallorrhiza trifida –1 –. 2510
Swertia perennis –1 –. 2510
Euphorbia amygdaloides –1 –. 2510
Paris quadrifolia –6 –3 2521
Calamagrostis villosa 2472 2572 –20
Other field layer species (E
1
)
Oxalis acetosella –65 1285 –85
Rubus idaeus –47 –49 –39
Gentiana asclepiadea –21 –39 2455
Luzula luzuloides –21 1942 –27
Calamagrostis arundinacea –32 –22 1138
Solidago virgaurea –19 –19 1230
Stellaria nemorum –20 –14 1429
Crepis paludosa 2237 –. 1835
Athyrium distentifolium –6 1223 –21
Gymnocarpium dryopteris –17 –15 –22
Chaerophyllum hirsutum 2032 –2 1429
Lycopodium annotinum –12 1820 –3
Huperzia selago –7 –13 –15
Melampyrum sylvaticum –10 –6 2123
Rumex alpestris –1 713 1416
Chrysosplenium alternifolium 1319 –. 1420
Milium effusum –6 –8 1719
Geum rivale 913 –. 1415
Urtica dioica 1916 –1 –8
Rubus saxatilis –9 –1 2118
Phegopteris connectilis –10 –5 –6
Hypericum maculatum –2 –2 1910
Geranium robertianum 1910 –. –3
Aconitum firmum s. l. –1 –2 2110
e15
Group A B C
No. of relevés 145 234 97
Differential ground layer species (E
0
)
Polytrichum commune 5552 –9 –.
Sphagnum palustre agg. 5440 –1 –1
Sphagnum girgensohnii 4859 –22 –4
Sphagnum squarrosum 3317 –1 –.
Leucobryum glaucum 3115 –. –1
Plagiomnium affine 3028 –10 –2
Pohlia nutans 3016 –2 –.
Pleurozium schreberi 2850 –27 –19
Polytrichum formosum –28 3964 –20
Lophozia ventricosa –. 2811 –.
Bazzania tricrenata –1 2611 –.
Calypogeia integristipula –12 2523 –.
Mnium spinosum –1 –1 5642
Ctenidium molluscum –. –. 4023
Tortella tortuosa –. –3 3926
Other ground layer species (E
0
)
Dicranum scoparium –76 1179 –61
Plagiothecium curvifolium –29 1440 –25
Hylocomium splendens –26 –31 1240
Rhytidiadelphus triquetrus –23 –14 –24
Lepidozia reptans 1921 –11 –4
Rhizomnium punctatum 1721 –3 –15
Plagiochila asplenioides 2121 –2 –12
Plagiothecium undulatum –8 1314 –4
Sphagnum capillifolium 1514 –9 –2
Tetraphis pellucida –10 –9 –5
Dicranella heteromalla –8 –10 –3
Blepharostoma trichophyllum –1 1712 –6
Bazzania trilobata 1712 –5 –2
Mylia taylorii –1 2110 –2
Eurhynchium angustirete 58 –1 1110
Plagiomnium undulatum 2011 –2 –2
Cirriphyllum piliferum 1210 –. –7
Brachythecium velutinum –1 –2 2212
Data:
Col. A: – Relevé dataset of wet woodlands with Picea abies used in this study.
Col. B: – 135 relevés of Piceion excelsae Pawłowski ex Pawłowski et al. 1928 prepared for the survey Plant communities of
Slovakia, Forest and shrub vegetation (Kučera in Valachovič et al., in prep.),
– 2 relevés of Piceion excelsae Pawłowski ex Pawłowski et al. 1928 excluded from that survey (relevé plot sizes 100 m2),
– 93 relevés of Homogyno alpinae-Pinion cembrae P. Kučera 2017 from Slovakia (see Kučera 2017),
– 4 lately published relevés of Homogyno alpinae-Pinion cembrae P. Kučera 2017, included in the survey Plant communities of
Slovakia, Forest and shrub vegetation (Kučera in Valachovič et al., in prep.).
Col. C: – 76 relevés of Cortuso-Piceion P. Kučera nom. prov. (Oxalido-Piceion sensu Hadač et al. 1969) prepared for the survey
Plant communities of Slovakia, Forest and shrub vegetation (Kučera in Valachovič et al., in prep.),
– 11 relevés of Cortuso-Piceion P. Kučera nom. prov. excluded from that survey (relevé plot sizes 150 m2 and 100 m2),
– 10 relevés of Calamagrostio variae-Pinion cembrae P. Kučera 2017 (see Kučera 2017).
e16
Supplement B2: Table 4. Differential table of associations of the order Sphagno palustris-Piceetalia abietis P. Kučera
2019 with fidelity (φ (× 100) ≥ 25)
B – alliance Sphagno palustris-Piceion abietis P. Kučera 2019
2 – Soldanello montanae-Piceetum Volk in Br.-Bl. et al. 1939
4 – Leucobryo glauci-Piceetum abietis Šomšák ex P. Kučera 2019
5 – Sphagno palustris-Piceetum Šomšák 1979
6 – Equiseto sylvatici-Piceetum Šmarda 1950
C – alliance Stellario nemorum-Abietion albae P. Kučera 2019
7 – Stellario nemorum-Abietetum albae P. Kučera 2019
D – alliance Valeriano dioicae-Abietion albae P. Kučera 2019
8 – Valeriano dioicae-Abietetum P. Kučera 2019
Alliance B C D
Group No. 2 4 5 6 7 8
No. of relevés 24 15 37 13 36 13
Trees and shrubs
E
3
Abies alba – – – – 63 –
Fagus sylvatica – – – – 47 –
Pinus sylvestris – – – – – 28
Alnus incana – – 12 – – 26
Alnus glutinosa – – – – – 26
E
2
Pinus sylvestris 27 – – – – –
Alnus incana – – 8 39 – –
Fagus sylvatica – – – – 61 –
Lonicera xylosteum – – – – 38 –
Sorbus aucuparia – – – – 35 –
Acer pseudoplatanus – – – – 34 –
Salix caprea – – – – 25 –
Sambucus racemosa – – – – 24 –
Viburnum opulus – – – – – 45
Frangula alnus – – – – – 36
Lonicera nigra – – – – 26 45
E
2
Salix aurita – – 39 – – –
Corylus avellana – – 26 – – –
Lonicera xylosteum – – – – 34 –
Fagus sylvatica – – – – 29 –
Daphne mezereum – – – – – 52
Viburnum opulus – – – – – 45
Ribes petraeum – – – – – 32
Sorbus aucuparia – – – – – 32
Frangula alnus – – – – – 25
Lonicera nigra – – – 26 – 34
Abies alba – – – – 29 26
Differential field layer species (E
1
)
Eriophorum vaginatum 38 – – – – –
Lycopodium annotinum 34 – – – – –
Listera cordata 28 – – – – –
Thelypteris palustris 27 – – – – –
Melampyrum sylvaticum – 48 – – – –
Avenella flexuosa – 30 – – – –
e17
Alliance B C D
Group No. 2 4 5 6 7 8
No. of relevés 24 15 37 13 36 13
Luzula luzuloides – 30 – – – –
Calluna vulgaris – 29 – – – –
Agrostis canina – – 58 – – –
Viola palustris – – 53 – – –
Ranunculus flammula – – 49 – – –
Juncus effusus – – 43 – – –
Potentilla erecta – – 41 – – –
Agrostis stolonifera – – 39 – – –
Carex rostrata – – 37 – – –
Valeriana simplicifolia – – 35 – – –
Carex canescens – – 33 – – –
Ajuga reptans – – 31 – – –
Carex pallescens – – 30 – – –
Moneses uniflora – – 28 – – –
Senecio "nemorensis" – – 28 – – –
Carex echinata – – 27 – – –
Galium uliginosum – – 26 – – –
Peucedanum palustre – – 26 – – –
Melampyrum pratense – – 26 – – –
Galium palustre – – 26 – – –
Trientalis europaea – – – 44 – –
Calamagrostis villosa – – – 28 – –
Veratrum album subsp. lobelianum – – – 25 – –
Stellaria nemorum – – – – 84 –
Chrysosplenium alternifolium – – – – 72 –
Petasites albus – – – – 71 –
Lysimachia nemorum – – – – 65 –
Geranium robertianum – – – – 59 –
Cardamine trifolia – – – – 59 –
Adenostyles alliariae – – – – 59 –
Impatiens noli-tangere – – – – 57 –
Luzula luzulina – – – – 57 –
Prenanthes purpurea – – – – 56 –
Gentiana asclepiadea – – – – 55 –
Rubus hirtus – – – – 54 –
Galium odoratum – – – – 52 –
Dryopteris dilatata – – – – 47 –
Milium effusum – – – – 47 –
Urtica dioica – – – – 44 –
Ranunculus lanuginosus – – – – 44 –
Ranunculus platanifolius – – – – 44 –
Phyteuma spicatum – – – – 41 –
Geum rivale – – – – 39 –
Carex sylvatica – – – – 39 –
Cardamine flexuosa – – – – 38 –
Poa remota – – – – 38 –
Calamagrostis epigejos – – – – 38 –
Veronica anagallis-aquatica – – – – 38 –
Homogyne alpina – – – – 37 –
Phegopteris connectilis – – – – 37 –
e18
Alliance B C D
Group No. 2 4 5 6 7 8
No. of relevés 24 15 37 13 36 13
Cicerbita alpina – – – – 34 –
Rubus idaeus – – – – 34 –
Dryopteris filix-mas – – – – 32 –
Luzula sylvatica – – – – 31 –
Oxalis acetosella – – – – 31 –
Epilobium montanum – – – – 29 –
Symphytum tuberosum – – – – 27 –
Sanicula europaea – – – – 27 –
Cardamine amara – – – 26
Poa palustris – – – – 26 –
Rubus saxatilis – – – – – 94
Valeriana dioica – – – – – 74
Polygonatum verticillatum – – – – – 71
Clematis alpina – – – – – 59
Cirsium oleraceum – – – – – 59
Filipendula ulmaria – – – – – 55
Thalictrum aquilegiifolium – – – – – 52
Carex alba – – – – – 52
Crepis paludosa – – 10 – – 51
Caltha palustris – – 8 – – 50
Galium schultesii – – – – – 49
Fragaria vesca – – – – 12 48
Astrantia major – – – – – 45
Melica nutans – – – – – 45
Solidago virgaurea – – – – – 39
Dactylorhiza maculata – – – – – 36
Carex remota – – – – – 36
Actaea spicata – – – – – 36
Carex digitata – – – – – 36
Maianthemum bifolium – – – – – 36
Equisetum palustre – – – – – 35
Epipactis palustris – – – – – 32
Polygonatum multiflorum – – – – – 32
Valeriana tripteris – – – – – 32
Paris quadrifolia – – – 20 – 31
Angelica sylvestris – – – – – 29
Persicaria bistorta – – – – – 25
Calamagrostis arundinacea – 32 – – 27 –
Luzula pilosa – 28 22 20 – 34
Lysimachia vulgaris – – 27 35 – –
Equisetum sylvaticum – – – 24 24 –
Chaerophyllum hirsutum – – – – 45 37
Senecio ovatus – – – – 40 27
Differential ground layer species (E
0
)
Sphagnum girgensohnii 25 – – – – –
Polytrichum commune 25 – – – – –
Leucobryum glaucum – 83 – – – –
Orthodicranum undulatum – 34 – – – –
Hylocomium splendens – 29 22 – – –
Dicranella heteromalla – 27 – – – –
e19
Alliance B C D
Group No. 2 4 5 6 7 8
No. of relevés 24 15 37 13 36 13
Brachythecium starkei – – 30 – – –
Lepidozia reptans – – 28 – – –
Sphagnum quinquefarium – – 26 – – –
Cephalozia bicuspidata – – 26 – – –
Calliergon cordifolium – – 26 – – –
Rhodobryum roseum – – 26 – – –
Chiloscyphus pallescens – – 26 – – –
Sphagnum palustre agg. – – 25 – – –
Lophocolea bidentata – – 12 40 – –
Bazzania trilobata 14 – – 25 – –
Cirriphyllum piliferum – – – – 54 –
Plagiomnium affine – – – – 47 –
Plagiothecium undulatum – – – – 38 –
Conocephalum conicum – – – – 34 –
Plagiomnium rostratum – – – – 34 –
Plagiomnium undulatum – – – – 31 –
Thuidium tamariscinum – – – – 31 –
Trichocolea tomentella – – – – – 36
Eurhynchium angustirete – – – – – 28
Tetraphis pellucida – – 10 – – 26
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