71 Diagnostic-value-of-the-Polypodium-vulgare-L.-complex-in-rocky-communities 71

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1 71 Diagnostic-value-of-the-Polypodium-vulgare-L.-complex-in-rocky-communities 71 Occurrence and diagnostic value of the Polypodium vulgare L. complex in the communities of Asplenietea trichomanis (Br. Bl. in Meier & Br. Bl. 1934) Oberd class in Polish Sudety Mts and their foreland KRZYSZTOF ŒWIERKOSZ Museum of Natural History, University of Wroc³aw, Sienkiewicza 21, PL Wroc³aw, ABSTRACT: In , 298 relevés of plant rocky communities of Asplenietea trichomanis class were collected in the Sudety Mts and their foreland. Polypodium vulgare occurred in 93 relevés, i.e. in about one third of collected material, however the distribution and abundance of the species were not equal in all syntaxonomical units. The species was found in 56% of relevés representing Hypno-Polypodion vulgaris alliance, in 26% of relevés of Asplenion septentrionalis alliance, in the same percentage in Asplenion cuneifolii phytocoenoses, and in less than 10% of all the relevés made on calcareous substrata. Thus, P. vulgare should be considered as a character species of the Androsacetalia vandelii order rather than the Asplenietea trichomanis class as a whole. It is especially abundant in phytocoenoses belonging to the Hypno-Polypodion vulgaris alliance and dominating in Hypno-Polypodietum associations, in which it reaches the highest coverage (10% up to 80%). The results indicate that the most important factors which determine the occurrence of P. vulgare in rocky communities are the type of the geological substrate (non-calcareous rocks) and shade, generally exceeding 60%. Occurrence of Polypodium mantoniae has been confirmed so far in the phytocoenoses belonging to the association Asplenio- Polypodietum and Asplenietum cuneifolii. It may occur in ass. Woodsio-Asplenietum septentrionalis as well. ABSTRAKT: W latach wykonano 298 zdjêæ fitosocjologicznych w zbiorowiskach naskalnych na terenie Sudetów i ich Przedgórza. Wystêpowanie Polypodium vulgare stwierdzono w 93 zdjêciach, czyli w 31% zebranego materia³u, jednak udzia³ gatunku w zdjêciach z poszczególnych jednostek syntaksonomicznych nie by³ równomierny. Paprotkê pospolit¹ stwierdzono w 56% zdjêæ reprezentuj¹cych fitocenozy ze zwi¹zku Hypno-Polypodion vulgaris, w 26% zdjêæ ze zwi¹zku Asplenion septentrionalis oraz zwi¹zku Asplenion cuneifolii, jednoczeœnie nie wiêcej ni w 10% Œwierkosz K Occurrence and diagnostic value of the Polypodium vulgare L. complex in the communities of Asplenietea trichomanis (Br. Bl. in Meier & Br. Bl. 1934) Oberd class in Polish Sudety Mts and their foreland. In: E. Szczêœniak, E. Gola (eds), Genus Polypodium L. in Poland. Polish Botanical Society, Wroc³aw, p

2 72 Krzysztof-Œwierkosz 72 zdjêæ wykonanych na pod³o ach z udzia³em wêglanu wapnia. Oznacza to, e gatunek nale y uznaæ za charakterystyczny dla rzêdu Androsacetalia vandelii, nie zaœ dla klasy Asplenietea trichomanis. Szczególnie czêsto wystêpuje w fitocenozach zwi¹zku Hypno-Polypodion vulgaris i dominuje w p³atach zespo³u Hypno-Polypodietum, w których osi¹ga tak e najwy sze stopnie pokrycia (od 10 do 80%). Uzyskane wyniki wskazuj¹, e najwa niejszymi czynnikami, które warunkuj¹ wystêpowanie P. vulgare w zbiorowiskach naskalnych w Sudetach i na ich przedgórzu s¹ rodzaj pod³o a (ska³y bezwapienne) oraz zacienienie (z regu³y przekraczaj¹ce 60%). Wystêpowanie Polypodium mantoniae zosta³o jak dot¹d potwierdzone w fitocenozach zaliczanych do zespo³ów Asplenio-Polypodietum i Asplenietum cuneifolii; mo e wystêpowaæ tak e w Woodsio-Asplenietum septentrionalis. Key words: Polypodium vulgare, Polypodium mantoniae, Asplenietea trichomanis, rocky plant communities, Sudety Mts, Central Europe. Introduction Rocky communities of Asplenietea trichomanis (Br. Bl. in Meier & Br. Bl. 1934) Oberd class are not a very popular subject of the studies. The communities with domination of Polypodium vulgare L. were described in Poland only in some papers (Anio³-Kwiatkowska, Œwierkosz 1992; Œwierkosz 1994; Kwiatkowski 1994; Berdowski, Panek 1999; Wilczek 2006, Nowak et al. 2008). Many theorems on the rocky communities variability and their ecological requirements come from the days when the phytosociological investigations have just begun. Even today many of the analyses are conducted based on the insufficient data what may result in erroneous interpretations. This causes many problems with the classification of the rocky communities during the field work, and later, during phytosociological data analysis. The detailed analysis of P. vulgare occurrence in the rocky communities belongs to the many problems that require solution. Is it really a species character for the Asplenietea trichomanis class? Is there a differentiation between the P. vulgare forms dominating in rocky communities? What are the preferred species habitat conditions? What are the most frequently co-occurring plant species? This article attempts to answer these questions. Materials and methods Research concerning the rocky and wall communities of the Sudety Mts and their foreland were conducted in relevés were collected

3 73 Diagnostic-value-of-the-Polypodium-vulgare-L.-complex-in-rocky-communities 73 according to Braun-Blanquet approach (Westhoff, van der Maarel 1978; Mueller- Dombois, Ellenberg 2003) and stored in a TURBOVEG database (Hennekens, Schaminée 2001). The area of the relevés was adapted to that proposed by Chytrý and Otypková (2003) for rocky communities. Shading, within the relevé, was determined as the sum of percentage cover generated by trees and shrubs in vertical projection. Relevés were classified using the TWINSPAN method (Roleèek et al. 2009) embedded within the JUICE program (Tichý 2002). Pseudospecies cut levels were set to 5, and values of cut levels to 0.1%, 0.5%, 2%, 16% and 32%. Three relevés were selected as a minimum group size for division. A Total Inertia method was used as a measure of cluster heterogeneity (Tichý et al. 2007). Diagnostic species for particular vegetation units were determined using the phi coefficient as a measure of fidelity (Chytrý et al. 2002) in a synoptic table. The phi coefficient was used for clusters of equalized size (Tichý, Chytrý 2006). Only species with both a significant concentration in particular vegetation units (using the Fisher s exact test and the significance level P < 0.05) and phi coefficient > 0.30 were considered to be diagnostic species. In this paper only relevés with presence of Polypodium sp. are analyzed and presented. The identification of the phytocoenoses was based both on the publications (Oberdorfer 1957, 1977; Jurko, Peciar 1963; Valachoviè 1995, 1996; Œwierkosz 2004; Œwierkosz et al. 2004; Matuszkiewicz 2008, Chytrý 2009), and on the analysis of the total phytosociological data. The nomenclature of the vascular plants follows Mirek et al. (2002), and the mosses nomenclature follows Ochyra et al. (2003). Identification of Polypodium mantoniae Rothm. during the field investigation is very difficult, and needs collection of samples from each relevé, and the laboratory analyses of each sample ploidy. All the data on this hybrid, presented in phytosociological tables, come from the research carried out by Szczêœniak et al. (mscr.). Results Polypodium vulgare was found in all documented wall and rocky communities in the Sudety Mts and their foreland, however its participation in different syntaxonomical units varied within wide limits. The species was found in 93 relevés which were 31% of all collected materials. It occurred in 56% of relevés representing Hypno-Polypodion vulgaris Mucina 1993 alliance, in 26% of relevés of Asplenion septentrionalis Oberd alliance, in 26% of Asplenion cuneifolii Br.-Bl. ex Eggler 1955 phytocoenoses, and in 6% of relevés from Potentilletalia caulescentis Br.-Bl. in Br.-Bl. et Jenny 1926 order. Single occurrences of Polypodium mantoniae have been confirmed so far in the phytocoenoses belonging to the association Asplenio-Polypodietum

4 74 Krzysztof-Œwierkosz 74 Firbas 1924 and Asplenietum cuneifolii Gauckler It may also occur in Woodsio-Asplenietum septentrionalis R. Tx. 1937, but it has not been confirmed yet (see also Szczêœniak et al. mscr.). Ass. Asplenietum rutae-murariae-trichomanis Kuhn 1937 Occurrence of P. vulgare was confirmed at two localities: in the Pe³cznica Gorge, on the wall near the Stary Ksi¹ castle in Wa³brzysko-Bolkowskie Foothills (Œwierkosz, Reczyñska 2009), and in one relevé made near Romanowo (the range of Krowiarki Hills) on marbles (Tab.1, relevé 1). The species cover did not exceed 5% in all relevés, and the habitat varied from exposed to shaded. Table 1. Occurrence of Polypodium vulgare L. in rocky communities on calcareous substrata. Tabela 1. Wystêpowanie Polypodium vulgare L. w zbiorowiskach naskalnych na pod³o ach wêglanowych. Relevé no. (Numer zdjêcia) Date: y/m/d/ (Data: r/m/d) Relevé area (Powierzchnia zdjêcia) m Altitude m. a.s.l. (Wysokoœæ n.p.m.) Aspect in degrees (Wystawa w stopniach) Slope (Nachylenie) Cover herb layer c (Pokrycie warstwy runa) % Cover moss layer d (Pokrycie mszaków) % Shadow (Zacienienie) % Number of species (Liczba gatunków) Locality (Stanowisko) ROM JEZ RG CG CG Ch. et D*. Ass. Asplenietum rutae-murariae trichomonis Asplenium trichomanes Asplenium ruta-muraria Tortula muralis d Ch. et D*. Ass. Cystopteridetum fragilis Asplenium viride Cystopteris fragilis Polystichum aculeatum* Saxifraga rosacea* Cardaminopsis arenosa ssp. borbasi* Ch. Cl. Asplenietea trichomanis et. D*. O. Potentilletalia caulescentis Polypodium vulgare Ctenidium molluscum* d Neckera complanata* d

5 75 Diagnostic-value-of-the-Polypodium-vulgare-L.-complex-in-rocky-communities 75 Accompanying (Towarzysz¹ce): Mycelis muralis Oxalis acetosella 1 + r + + Geranium robertianum Urtica dioica Galeobdolon luteum. r. + + Picea abies c.. r r r Epilobium montanum. + + r. Fagus sylvatica c r. r.. Mercurialis perennis Asarum europaeum Athyrium filix-femina Poa nemoralis Senecio ovalis Sporadic (Sporadycznie): Chelidonium majus 1 (1); Impatiens noli-tangere 2 (1); Conocephalum conicum d 2 (1); Campanula rapunculoides 3 (1); Hieracium murorum 3 (+); Actaea spicata 3 (r.); Dryopteris filix-mas 3 (+). The localities and their geological substrata: ROM - Romanowo, the Krowiarki Hills, marbles; JEZ - Je owice, RG - Rogowa Kopa and CG - the Czernica Gorge, all localities in the Sto³owe Mts, marls. Lokalizacja i pod³o e geologiczne: ROM - Romanowo, Krowiarki, marmur; JEZ - Je owice, RG - Rogowa Kopa i CG - w¹wóz Czernicy, wszystkie w Górach Sto³owych, margle. Ass. Cystopteridetum fragilis Oberd In this association in the Sudety Mts, P. vulgare has been found so far at three localities only. They are in the Sto³owe Mts, in deep gorges cut in marl, on the edges of summit plane (Tab.1, relevés 2-5). The P. vulgare cover varied between 0.1 and 5%. The most common co-occurring species (in more than 50% of relevés) were: Cystopteris fragilis, Asplenium viride, Mycelis muralis, Oxalis acetosella, Geranium robertianum, Epilobium montanum, Galeobdolon luteum, mosses: Ctenidium molluscum, Neckera complanata, and seedlings of Picea abies. The habitat was shaded (average 67.5% of shade) and moist. Most of the collected relevés of Cystopteridetum fragilis are devoid of P. vulgare, even if the habitat conditions are convenient for this species. Ass. Asplenietum cuneifolii Gauckler 1954 Polypodium vulgare was found in 9 relevés (i.e. 26%) representing Asplenion cuneifolii phytocoenoses (Tab. 2). There, the P. vulgare cover varied between 0.1 and 20%. The most common co-occurring species (in more than 50% of relevés) were Asplenium cuneifolium, A. adulterinum, Silene vulgaris, and only one moss, Hypnum cupressiforme. The habitat was partly shadowed (average 32.2% of shade), and moderately moist, developed on the serpentine rocks (Fig.1). Occurrence of P. mantoniae in the phytocoenoses of Asplenietum

6 Table 2. Occurrence of Polypodium vulgare L. and P. mantoniae Rothm. in ass. Asplenietum cuneifolii Gauckler Tabela 2. Wystêpowanie Polypodium vulgare L. i P. mantoniae Rothm. w fitocenozach zespo³u Asplenietum cuneifolii Gauckler Relevé no. (Numer zdjêcia) Date: y/m/d/ (Data: r/m/d) Relevé area (Powierzchnia zdjêcia) m C Altitude m. a.s.l. (Wysokoœæ n.p.m.) o Aspect in degrees (Wystawa w stopniach) n Slope (Nachylenie) s Cover shrub layer b (Zwarcie krzewów) % t Cover herb layer c (Pokrycie warstwy runa) a Cover moss layer d (Pokrycie mszaków) % n Shadow (Zacienienie) % c Number of species (Liczba gatunków) y Locality (Stanowisko) WK WK WK PRZ PRZ RAD RAD RAD JOR Ch. et. D.* Ass. Asplenietum cuneifolii Asplenium cuneifolium r + + r r Silene vulgaris* r +. r + + r Asplenium adulterinum Ch. et. D.* Cl. Asplenietea trichomanis Polypodium vulgare et P. mantoniae Hypnum cupressiforme* d Asplenium ruta-muraria Asplenium septentrionale Asplenium trichomanes Cystopteris fragilis Krzysztof-Œwierkosz 76

7 D. O. Androsacetalia vandelii Calamagrostis arundinacea Campanula rotundifolia r r.. 33 Dryopteris carthusiana Deschampsia flexuosa Sporadic (Sporadycznie): Dicranum scoparium d 8 (1); Luzula pilosa 7 (r.); Polytrichastrum formosum d 7 (+); Sedum maximum 6 (+); Sorbus aucuparia c 6 (+); Vaccinium myrtillus 7 (r.). Accompanying (Towarzysz¹ce): Rubus idaeus b Rubus idaeus c... r Poa nemoralis Achillea millefolium r Arenaria serpyllifolia Galium verum Phleum phleoides Sedum acre Festuca ovina..... r Dryopteris filix-mas Pohlia nutans d Sporadic (Sporadycznie): Acer platanoides c 1 (r.); Arrhenatherum elatius 9 (+); Brachythecium albicans 9 (2); Campanula persicifolia 8 (r.); Centaurea rhenana 2 (+); Dianthus carthusianorum 9 (+); Epilobium angustifolium 7 (+); Euphorbia cyparissias 3 (+); Festuca pallens 6 (2); Fragaria vesca 3 (+); Fraxinus excelsior c 9 (r.); Galium pumilum agg. 8 (1); Viscaria vulgaris 2 (+); Potentilla arenaria 2 (+); Cerasus avium 1 b (+), c (+); Senecio sp. 3 (r.); Solanum dulcamara 1 (r.); Stellaria media 9 (+); Taraxacum officinale agg. 2 (+); Thymus pulegioides 2 (+); Veronica chamaedrys 1 (+). Explanations: 1 - occurrence of Polypodium mantoniae was confirmed in the relevés from this locality (Szczêœniak et al. mscr.), but it was not possible to determine its abundance when it occurred together with P. vulgare. Objaœnienia: 1 - wystêpowanie Polypodium mantoniae w tym zdjêciu by³o potwierdzone (Szczêœniak et al. mscr.), lecz nie by³o mo liwe okreœlenie pokrycia, gdy wystêpowa³ razem z P. vulgare. Localities: WK - the Wzgórza Kie³czyñskie Hills; PRZ - Przemi³ów near Sulistrowice (the Wzgórza Oleszeñskie Hills); RAD - "Radunia" Nature Reserve (the Wzgórza Oleszeñskie Hills); JOR - a small quarry between Glinica and Jordanów (the Wzgórza Oleszeñskie Hills). Lokalizacje: WK - Wzgórza Kie³czyñskie; PRZ - Przemi³ów k. Sulistrowic (Wzgórza Oleszeñskie); RAD - rezerwat przyrody Radunia (Wzgórza Oleszeñskie); JOR - ma³y kamienio³om miêdzy Glinic¹ a Jordanowem (Wzgórza Oleszeñskie). 77 Diagnostic-value-of-the-Polypodium-vulgare-L.-complex-in-rocky-communities 77

8 78 Krzysztof- wierkosz 78 Fig. 1. Polypodium vulgare L. in ass. Asplenietum cuneifolii Gauckler 1954; the Wzgórza Kie³czyñskie Hills, Sudety Foreland (photo K. wierkosz). Ryc. 1. Polypodium vulgare L. w zespole Asplenietum cuneifolii Gauckler 1954; Wzgórza Kie³czyñskie, Przedgórze Sudeckie (fot. K. wierkosz). cuneifolii was confirmed at least for one locality in Radunia (Szczê niak et al. mscr.), but its detailed distribution needs further research. Ass. Asplenio-Polypodietum Firbas 1924 Polypodium vulgare was found in most of the patches of this association, so in Tab. 3 only 10 representative relevés are shown. The species cover in this association varied between 0.1% and 5%; higher coverage, up to 80%,was achieved only exceptionally. The most common species co-occurring with P. vulgare in more than 40% of relevés were Asplenium trichomanes, Mycelis muralis, Cystopteris fragilis, Acer pseudoplatanus, Galeobdolon luteum, Geranium robertianum, Poa nemoralis, Oxalis acetosella and the moss Hypnum cupressiforme. The habitat was shaded (average 64% of shade), and moderately moist. The geological substrate of plots included neutral or weakly acidic rocks: basalt, greenstone, trachyte, and some kinds of granodiorite, greywacke and mudrocks.

9 Table 3. Occurrence of Polypodium vulgare L. and P. mantoniae Rothm. in ass. Asplenio-Polypodietum Firbas 1924 (representative relevés). Tabela 3. Wystêpowanie Polypodium vulgare L. i P. mantoniae Rothm. w fitocenozach zespo³u Asplenio-Polypodietum Firbas 1924 (przyk³adowe zdjêcia). Relevé no. (Numer zdjêcia) Date: y/m/d/ (Data: r/m/d) Relevé area (Powierzchnia zdjêcia) m C Altitude m. a.s.l. (Wysokoœæ n.p.m.) o Aspect in degrees (Wystawa w stopniach) n Slope (Nachylenie) s Cover shrub layer b (Zwarcie krzewów) % t Cover herb layer c (Pokrycie warstwy runa) a Cover moss layer d (Pokrycie mszaków) % n Shadow (Zacienienie) % c Number of species (Liczba gatunków) y Locality (Stanowisko) WMY WMY WMY MIE LUT PST PST OST MOR WLI Ch. et D*. Ass. Asplenio-Polypodietum Asplenium trichomanes* Cystopteris fragilis* r r Mycelis muralis* r + + r. + + r Galeobdolon luteum* Phyllitis scolopendrium Urtica dioica* Acer pseudoplatanus* b + r Acer pseudoplatanus* c Ch. et D.* Cl. Asplenietea trichomanis Polypodium vulgare et P. mantoniae 1 r Hypnum cupressiforme* d Asplenium ruta-muraria r Diagnostic-value-of-the-Polypodium-vulgare-L.-complex-in-rocky-communities 79

10 D. O. Androsacetalia vandellii Sedum maximum Calamagrostis arundinacea r 20 Dryopteris carthusiana Sporadic (Sporadycznie): Campanula rotundifolia 10 (r.); Dicranum scoparium d 8 (2); Luzula luzuloides 5 (+); Polytrichastrum formosum 8 (2). Accompanying (Towarzysz¹ce): Dryopteris filix-mas +. + r Geranium robertianum Poa nemoralis r 50 Epilobium montanum r Impatiens parviflora r. r Stellaria media r r Campanula rapunculoides. r r. 20 Galium schultesii. r Ribes grossularia b Ribes grossularia c... r Adoxa moschatellina r Chrysosplenium alternifolium Oxalis acetosella Moehringia trinervia r Conocephalum conicum d Cardamine impatiens. r Sporadic (Sporadycznie): Acer platanoides b 6 (1); Athyrium filix femina 7 (1); Bartramia sp. d 8 (1); Bilderdykia dumetorum 8 (r.); Festuca pallens 9 (+); Fraxinus excelsior c 7 (+); Galeopsis pubescens 8 (r.); Hieracium murorum 7 (+); Lonicera xylosteum b 1 (1); Mercurialis perennis 7 (+); Mnium hornum d 3 (3); Mnium sp. d 8 (+); Plagiothecium sp. d 8 (1); Rubus idaeus 4 (r.); Senecio ovatus 7 (1); Taraxacum officinale agg. 4 (r.); Ulmus glabra b 3 (3). Explanations: 1 - occurrence of Polypodium mantoniae was confirmed in the relevés from this locality (Szczêœniak et al. mscr.), 2 - occurrence of Polypodium mantoniae was excluded. A laboratory test showed no hybrids within the collected Polypodium samples. Objaœnienia: 1 - obecnoœæ Polypodium mantoniae w zdjêciu z tego stanowiska zosta³a potwierdzona (Szczêœniak et al. mscr., 2 - obecnoœæ Polypodium mantoniae zosta³a wykluczona. Badania laboratoryjne nie wykaza³y obecnoœci mieszañca w pobranych próbkach. Localities: WMY - W¹wóz Myœliborski (the Pogórze Kaczawskie Foothill, greenstone); MIE - Mieroszów (the Góry Kamienne Mts, trachyte); LUT - Lutynia (the Góry Z³ote Mts, basalt); PST - Pstr¹ na (the Wzgórza Lewiñskie Hills, granodiorite); OST - Ostrzyca Proboszczowicka Hill (the Pogórze Kaczawskie Foothill, basalt); MOR - Morzyszów (the Góry Bardzkie Mts, mudrocks and greywacke); WLI - W¹wóz Lipa Gorge (the Pogórze Kaczawskie Foothill, greenstone). Lokalizacje: WMY - W¹wóz Myœliborski (Pogórze Kaczawskie, zieleñce); MIE - Mieroszów (Góry Kamienne, trachity); LUT - Lutynia (Góry Z³ote, bazalt); PST - Pstr¹ na (Wzgórza Lewiñskie, granodioryt); OST - Ostrzyca Proboszczowicka (Pogórze Kaczawskie, bazalt); MOR - Morzyszów (Góry Bardzkie, mu³owce i szarog³azy); WLI - W¹wóz Lipy (Pogórze Kaczawskie, zieleñce). 80 Krzysztof-Œwierkosz 80

11 81 Diagnostic-value-of-the-Polypodium-vulgare-L.-complex-in-rocky-communities 81 Occurrence of P. mantoniae in the phytocoenoses of Asplenio-Polypodietum was confirmed at least in two localities: the hill of Ostrzyca Proboszczowicka and the Lipa Gorge (Szczê niak et al. mscr.), but it needs further research, especially on basalt and greenstone rocks. Ass. Hypno-Polypodietum Jurko et Peciar 1963 Polypodium vulgare was found in all of the patches of this association, so in Tab. 4 only 10 representative relevés are shown. The species cover in this association varied between 5% and 80%. The most common species co-occurring with P. vulgare (in more than 40% of relevés) were Poa nemoralis, Dryopteris filix-mas, and the moss Hypnum cupressiforme.the habitat was shaded (average 68% of shade), and moist (Fig. 2). The geological substrate included rocks very poor in nutrients mainly acidic (granite, rhyolite) and sedimentary (quartz sandstone, conglomerate), rarely metamorphic (amphibolite, migmatite) or alkaline plutonic rocks (gabbro). Fig. 2. Polypodium vulgare L. in ass. Hypno-Polypodietum Jurko et Peciar 1963; Panieñskie Ska³y, Pogórze Izerskie Foothill (photo K. wierkosz). Ryc. 2. Polypodium vulgare L. w zespole Hypno-Polypodietum Jurko et Peciar 1963; Panieñskie Ska³y, Pogórze Izerskie (fot. K. wierkosz).

12 Table 4. Occurrence of Polypodium vulgare L. in ass. Hypno-Polypodietum Jurko et Peciar 1963 (exemplary relevés). Tabela 4. Wystêpowanie Polypodium vulgare L. w fitocenozach zespo³u Hypno-Polypodietum Jurko et Peciar 1963 (przyk³adowe zdjêcia). Relevé no. (Numer zdjêcia) Date: y/m/d/ (Data: r/m/d) Relevé area (Powierzchnia zdjêcia) m C Altitude m. a.s.l. (Wysokoœæ n.p.m.) o Aspect in degrees (Wystawa w stopniach) n Slope (Nachylenie) s Cover shrub layer b (Zwarcie krzewów) % t Cover herb layer c (Pokrycie warstwy runa) a Cover moss layer d (Pokrycie mszaków) % n Shadow (Zacienienie) % c Number of species (Liczba gatunków) y Locality (Stanowisko) BAR WPE MIC NIE DKA DKA BOL OLS OLS DEM D. (dom.) Ass. Hypno-Polypodietum Polypodium vulgare Hypnum cupressiforme d D. O. Androsacetalia vandelli et Ch.* Cl. Asplenietea trichomanis Calamagrostis arundinacea Dicranum scoparium d Polytrichastrum formosum d Dryopteris dilatata Dryopteris carthusiana Krzysztof-Œwierkosz 82

13 Sporadic (Sporadycznie): Cystopteris fragilis* 6 (+); Deschampsia flexuosa 7 (+); Luzula luzuloides 9 (+); Vaccinium myrtillus 1 (1). Accompanying (Towarzysz¹ce): Dryopteris filix-mas Poa nemoralis. r Oxalis acetosella + r Impatiens parviflora. r Plagiothecium cf. denticulatum d Fagus sylvatica c +. r Pohlia nutans d Sambucus racemosa b Sambucus racemosa c Sporadic (Sporadycznie): Acer platanoides c 2 (r.); Dicranella heteromalla d 3 (1); Epilobium angustifolium 5 (+); Fraxinus excelsior b 8 (1); Gymnocarpium dryopteris 5 (2); Mnium sp. d 2 (3); Picea abies c 3 (r.); Populus tremula c 9 (1); Ribes uva-crispa 2 b (2), c (+); Rubus hirtus agg. 10 (+); Rubus idaeus c 1 (+); Rubus sect. Corylifolii 9 (+); Sambucus nigra b 2 (1); Senecio ovatus 3 (r.). Explanations: 2 - occurrence of Polypodium mantoniae was excluded. A laboratory test showed no hybrids within the collected Polypodium samples. Objaœnienia: 2 - wystêpowanie Polypodium mantoniae zosta³o wykluczone. Badania laboratoryjne nie wykaza³y obecnoœci mieszañca w pobranych próbkach Polypodium. Localities: BAR - Barcinek (the Pogórze Izerskie Foreland, granite); WPE - W¹wóz Pe³cznicy Gorge (the Pogórze Wa³brzyskie Foothills, conglomerate); MIC - Micha³kowa (the Góry Sowie Mts, gneiss); NIE - Nielestno (the Pogórze Kaczawskie Foreland, quartz sandstone); DKA - Dolina Kamienicy Gorge (the Pogórze Izerskie Foreland, granite); BOL - north of Bolków (the Pogórze Kaczawskie Foothills, rhyolite); OLS - Olszak (the Góry Opawskie Mts, conglomerate); DEM - Dêbówka by Nowa Ruda (the Góry Sowie Mts, gabbro). Lokalizacje: BAR - Barcinek (Pogórze Izerskie, granit); WPE - W¹wóz Pe³cznicy (Pogórze Wa³brzyskie, zlepieniec); MIC - Micha³kowa (Góry Sowie, gneis); NIE - Nielestno (Pogórze Kaczawskie, piaskowce kwarcytowe); DKA - Dolina Kamienicy (Pogórze Izerskie, granit); BOL - na pó³noc od Bolkowa (Pogórze Kaczawskie, ryolit); OLS - Olszak (Góry Opawskie, zlepieniec); DEM - Dêbówka k. Nowej Rudy (Góry Sowie, gabro). 83 Diagnostic-value-of-the-Polypodium-vulgare-L.-complex-in-rocky-communities 83

14 84 Krzysztof-Œwierkosz 84 Occurrence of P. mantoniae in the phytocoenoses of Hypno-Polypodietum has not been confirmed so far, and it is doubtful due to adverse habitat conditions, namely rocks poor in nutrients and strong shading of the sites. Ass. Woodsio-Asplenietum septentrionalis R. Tx Polypodium vulgare was found in 26% of relevés (24) belonging to this association, so the Tab. 5 shows only 10 exemplary ones. The species abundance in this association varied between 0.1% and 30%. The most common species co-occurring with P. vulgare in more than 40% of relevés were Poa nemoralis, Sedum maximum, Asplenium septentrionale, Hieracium sabaudum and mosses: Ceratodon purpureus and Hypnum cupressiforme.the habitat was exposed (average 19% of shade), and moderately dry. The geological substrate included all kinds of rock, both poor in nutrients, such as granite, gabbro, gneiss, quartz sandstone, and moderately rich, e.g. basalt, greywacke, greenstone, granodiorite, trachyte. Occurrence of P. mantoniae in the phytocoenoses of Woodsio-Asplenietum septentrionalis has not been confirmed so far, but is very probable because at least in two localities, near the Zbiornik Bystrzycki Reservoir and the Ostrzyca Proboszczowicka Hill, the hybrid has been found in the close vicinity of the relevés presented in the table (Szczêœniak et al. mscr.). Discussion and conclusion The syntaxonomical role of Polypodium vulgare in the rocky communities of Central Europe is still unclear. Most of the authors place it between the character species of the class Asplenietea trichomanis, except for Jurko and Peciar (1963), who proposed a new class Polypodietea, and other subordinate units as orders, alliances and associations. The results presented in this paper suggest that P. vulgare should not be longer treated as a character species of the Asplenietea trichomanis class, because it occurs with variable constancy in its subunits. The last syntaxonomical revision of this class (Chytrý 2009) also noted this differentiation and considered P. vulgare as a character only for the alliances Asplenion septentrionalis, Asplenion cuneifolli, and especially for association Asplenio-Polypodietum. It means that the species should be treated as a character of the Androsacetalia vandelii order rather than the Asplenietea trichomanis class as a whole. Another problem to solve is a differentiation of communities with domination of P. vulgare in Central Europe. The concept presented by Jurko and Peciar (1963), was disputed by Valachoviè (1996), who compared the material of P. vulgare-dominated communities from Central Europe and showed the lack of methodological bases to distinguish specific units, not only at the class and

15 Table 5. Occurrence of Polypodium vulgare L. and P. mantoniae Rothm. in ass. Woodsio-Asplenietum septentrionalis R. Tx (representative relevés). Tabela 5. Wystêpowanie Polypodium vulgare L. i P. mantoniae Rothm. w fitocenozach zespo³u Woodsio-Asplenietum septentrionalis R. Tx (przyk³adowe zdjêcia). Relevé no. (Numer zdjêcia) Date: y/m/d/ (Data: r/m/d) Relevé area (Powierzchnia zdjêcia) m C Altitude m. a.s.l. (Wysokoœæ n.p.m.) o Aspect in degrees (Wystawa w stopniach) n Slope (Nachylenie) s Cover shrub layer b (Zwarcie krzewów) % t Cover herb layer c (Pokrycie warstwy runa) a Cover moss layer d (Pokrycie mszaków) % n Shadow (Zacienienie) % c Number of species (Liczba gatunków) y Locality (Stanowisko) ZB PP ZB PP BAR JAR JAR ZPI ZPI OST Ch. et D*. Ass. Woodsio-Asplenietum septentrionalis Asplenium septentrionale c Ceratodon purpureus* d Hieracium sabaudum* c + r r Sedum maximum* c Polytrichum piliferum* d D. O. Androsacetalia vandelli Campanula rotundifolia c r Calamagrostis arundinacea c Deschampsia flexuosa c r Diagnostic-value-of-the-Polypodium-vulgare-L.-complex-in-rocky-communities 85

16 Sporadic (Sporadycznie): Dicranum scoparium d 5 (1); Luzula luzuloides 1 (+); Sorbus aucuparia c 4 (r.). Ch. et D*. Cl. Asplenietea trichomanis Polypodium vulgare et P. mantoniae 1 c r 2 r 1 + r Hypnum cupressiforme* d Asplenium trichomanes c Asplenium ruta-muraria c r.. 10 Accompanying (Towarzysz¹ce): Poa nemoralis c. r Silene nutans c Dryopteris filix-mas c r Hieracium lachenalii c. r Hieracium murorum c r Thymus pulegioides c Betula pendula c Rhacomitrium cf. canescens d Fagus sylvatica c r Quercus petraea b r Quercus petraea c Sporadic (Sporadycznie): Acer platanoides c 4 (r.); Acinos arvensis 10 (+); Bartramia sp. 5 (1); Bryum sp. 1 (+), 6 (1); Calluna vulgaris 5 (+); Festuca duriuscula 8 (1); Festuca pallens 10 (2); Fraxinus excelsior 4 (r.); Genista tinctoria 4 (+); Impatiens parviflora 3 (1); Potentilla argentea 10 (r.); Potentilla collina 8 (r.); Cerasus avium b 7 (1); Rosa canina b 4 (+); Rubus idaeus b 3 (1); Silene vulgaris 10 (1); Taraxacum officinale agg. 9 (r.); Tortella sp. d 10 (+); Urtica dioica 6 (r.). Explanations: 1 - occurrence of Polypodium mantoniae is probable, the hybrid was confirmed in the close vicinity of the plot (Szczêœniak et al. mscr.). 2 - occurrence of Polypodium mantoniae was excluded. A laboratory test showed no hybrids within the collected Polypodium samples. Objaœnienia: 1 - wystêpowanie P. mantoniae jest prawdopodobne, mieszaniec zosta³ potwierdzony w s¹siedztwie p³atu (Szczêœniak et al. mscr.). 2 - wystêpowanie P. mantoniae wykluczono. Badania laboratoryjne nie wykaza³y obecnoœci mieszañca w zebranych próbach Polypodium. Localities: ZB - Zbiornik Bystrzycki Reservoir (the Góry Sowie Mts, gneiss); PP - "Piaskowcowe Porwaki" Nature Monument (the Pogórze Izerskie Foothills, basalt); BAR - Janowiec (the Góry Bardzkie Mts, greywacke); JAR - Jarno³tówek (the Góry Opawskie Mts, greywacke); ZPI - Zbiornik Pilichowicki Reservoir (the Pogórze Izerskie Foothills, basalt); OST - Ostrzyca Proboszczowicka Hill (the Pogórze Kaczawskie Foothills, basalt). Lokalizacje: ZB - Zbiornik Bystrzycki (Góry Sowie, gnejs); PP - pomnik przyrody Piaskowcowe Porwaki (Pogórze Izerskie, bazalt); BAR - Janowiec (Góry Bardzkie, szarog³az); JAR - Jarno³tówek (Góry Opawskie, szarog³az); ZPI - Zbiornik Pilichowicki (Pogórze Izerskie, bazalt); OST - Ostrzyca Proboszczowicka (Pogórze Kaczawskie, bazalt). 86 Krzysztof-Œwierkosz 86

17 87 Diagnostic-value-of-the-Polypodium-vulgare-L.-complex-in-rocky-communities 87 order levels, but even at the level of association. From that moment, most of the authors (comp. Chytrý 2009) found no grounds to distinguish the Hypno- Polypodietum association, and treated it as a synonym of earlier described Asplenio-Polypodietum. However it is worth mentioning that all the data, which had been collected to that time and then analyzed by Valachoviè (1996) contained only 46 relevés from Europe, and none of them came from Poland. The preliminary results presented in this paper suggest the presence of significant differences between the phytocoenoses of Asplenio-Polypodietum and Hypno-Polypodietum, as it is showed in Tab. 2, Tab. 4 and Tab. 6. These differences are related to both floristic compositions (see Tab. 6), and main ecological factors, particularly the type of geological substrata. This relationship was also noticed by Chytrý (2009), but it became the basis for distinguishing Table 6. Differentiation of Androsacetalia vandelli communities based on 84 relevés with the occurrence of Polypodium vulgare L. and P. mantoniae Rothm. (shortened table). Tabela 6. Zró nicowanie zbiorowisk z rzêdu Androsacetalia vandelli w oparciu o klasyfikacjê 84 zdjêæ fitosocjologicznych z udzia³em Polypodium vulgare L. i P. mantoniae Rothm. (tabela skrócona). Association (Zespó³) As-P W-As As. c Hy-P Number of relevés (Liczba zdjêæ) Ch. et D*. Ass. Mycelis muralis* c Cystopteris fragilis c Acer pseudoplatanus juv.* c Galeobdolon luteum* c Epilobium montanum* c Urtica dioica* c Senecio ovatus* c Phyllitis scolopendrium c Asplenium trichomanes c Ceradoton purpureus* d Hieracium sabaudum* c Asplenium septentrionale c Sedum maximum* c Asplenium cuneifolium c Silene vulgaris* c Asplenium adulterinum c.. 56.

18 88 Krzysztof-Œwierkosz 88 Ch. et D.* Cl. Asplenietea trichomanis Polypodium vulgare and P. mantoniae c 100 r r Hypnum cupressiforme* d Asplenium ruta-muraria c Epilobium collinum c D. O. Androsacetalia vandellii Calamagrostis arundinacea c Polytrichastrum formosum d Campanula rotundifolia c Sorbus aucuparia b/c Dicranum scoparium d Luzula luzuloides c Dryopteris carthusiana c Deschampsia flexuosa c Solidago virgaaurea c Accompanying (Towarzysz¹ce): Poa nemoralis c Dryopteris filix-mas c Rubus idaeus c Campanula persicifolia c Acer platanoides b/c Impatiens parviflora c Fraxinus excelsior juv. c Hieracium murorum c Fagus sylvatica juv. c Stellaria media c Sambucus racemosa b/c Taraxacum officinale agg. c Campanula rapunculoides c Epilobium angustifolium c Festuca pallens c Oxalis acetosella c Geranium robertianum c Abbreviations of the association names (Skróty nazw zespo³ów): As-P Asplenio-Polypodietum Firbas 1924, W-As Woodsio-Asplenietum septentrionalis R. Tx.1937, As.c Asplenietum cuneifolii Gauckler 1954, Hy-P Hypno-Polypodietum Jurko et Peciar 1963.

19 89 Diagnostic-value-of-the-Polypodium-vulgare-L.-complex-in-rocky-communities 89 variants only. Also, it should be stated here that the analyses of Androsacetalia vandelli communities, prepared for the whole Czech Republic, were based on 96 relevés. In comparison to material analyzed here, it makes the number of data a small part of all the material collected in the Sudety Mts. Status of both associations require further study and analyses based on representative material of other Central European mountain ranges. Also the syntaxonomical position of P. mantoniae should be solved in the future. Although it seems to have similar preferences to its parent, P. vulgare, probably it does not occur in the Hypno-Polypodietum phytocoenoses at all. Acknowledgements Special thanks to Dr. Stephan Hennekens for permission to use the free Turboweg software, to Dr. Ewa Szczêœniak for inspiring discussions and to Kamila Reczyñska, M.Sc. for her irreplaceable assistance in field studies and in the preparation of this paper. I would also thank to two anonymous Reviewers for their valuable comments and advice. References ANIO -KWIATKOWSKA J., ŒWIERKOSZ K Flora i roœlinnoœæ rezerwatu Ostrzyca Proboszczowicka oraz jego otoczenia. Acta Univ. Wrat Pr. Bot. 48: BERDOWSKI W., PANEK E Roœlinnoœæ rezerwatu Góra Radunia w województwie wroc³awskim. Parki Narodowe i Rezerwaty Przyrody 18(2): CHYTRÝ M. (ed.) Vegetace Èeské republiky 2. Ruderální, plevelová, skalní a su ová vegetace. Academia Praha, 488 pp. CHYTRÝ M., OTYPKOVÁ Z Plot sizes used for phytosociological sampling of European vegetation. J. Veg. Sci. 14: CHYTRÝ M., TICHÝ L., HOLT J., BOTTA-DUKÁT Z Determination of diagnostic species with statistical fidelity measures. J. Veg. Sci. 13: HENNEKENS S. M., SCHAMINÉE, J. H. J TURBOVEG, a comprehensive data base management system for vegetation data. J. Veg. Sci. 12: JURKO A., PECIAR V Pflanzengesellschaften am schattigen Felsen in den Westkarpaten. Vegetatio 11: KWIATKOWSKI P Szata roœlinna projektowanego rezerwatu Góra Zamkowa ko³o Wlenia. Acta Univ. Wrat Prace Bot. 60: MATUSZKIEWICZ W Przewodnik do oznaczania zbiorowisk roœlinnych Polski. Wyd. Nauk. PWN, Warszawa, 537 pp. MIREK Z., PIÊKOŒ-MIRKOWA H., ZAJ C A., ZAJ C M Flowering plants and pteridophytes of Poland: a checklist. W. Szafer Institute of Botany, Polish Academy of Science, Kraków, 442 pp. MUELLER DOMBOIS D., ELLENBERG H Aims and methods of vegetation ecology. The Blackburn Press, 547 pp.

20 90 Krzysztof-Œwierkosz 90 NOWAK A., NOWAK S., STEBEL A The association of Hypno-Polypodietum Jurko et Peciar 1963 in the Opawskie Mts. (the Eastern Sudetes Mis). In: E. SZCZÊŒNIAK, E. GOLA (eds.), Club mosses, horsetails and ferns in Poland resources and protection. Polish Botanical Society & Institute of Plant Biology, University of Wroc³aw, pp OBERDORFER E Süddeutsche Pflanzengesellschaften. Pflanzensiozologie 10. Gustav Fischer Verlag, Stuttgart Jena. OBERDORFER E Klasse: Asplenietea rupestria Br.-Bl. 34 in Meier et Br.-Bl. 34. In: E. OBERDORFER (ed.), Süddeutsche Pflanzengesellschaften. 1: Gustav Fischer Verlag, Stuttgart New York. OCHYRA R., BEDNAREK-OCHYRA H., ARNOWIEC J Census catalogue of Polish mosses. W. Szafer Institute of Botany, Polish Academy of Science, Kraków, 372 pp. ROLEÈEK J., TICHÝ L., ZELENÝ D., CHYTRÝ M Modified TWINSPAN classification in which the hierarchy respects cluster heterogeneity. J. Veg. Sci. 20: SZCZÊŒNIAK E., PIELECH R., JÊDRZEJCZYK I., RECZYÑSKA K., ŒWIERKOSZ K. Occurrence of Polypodium interjectum Shivas and P. mantoniae Rothm. in Polish Sudety Mts. (2012) mscr. ŒWIERKOSZ K Zbiorowiska roœlinne Góry Chojnik eksklawy Karkonoskiego Parku Narodowego. Czêœæ 2. Zbiorowiska nieleœne. Parki Narodowe i Rezerwaty Przyrody 13: ŒWIERKOSZ K Notes on the syntaxonomy of the Asplenietea trichomanis class in Poland. Polish Botanical Journal 49(2): ŒWIERKOSZ K., PERZANOWSKA J., MRÓZ W Œciany skalne i urwiska krzemianowe ze zbiorowiskami z Androsacetalia vandellii. In: HERBICH J. (ed.) Poradniki ochrony siedlisk i gatunków Natura 2000, podrêcznik metodyczny, T. 4. Œciany, piargi i rumowiska skalne, jaskinie. Ministerstwo Œrodowiska, Warszawa, pp ŒWIERKOSZ K., RECZYÑSKA K Livid hawkweed (Hieracium bifidum Kit. ex Hornem), a new species for the Red List of Lower Silesia, in the Pe³cznica Gorge (Wa³brzych-Bolków Foothills, Sudetes). Opole Scientific Society Nature Journal 42: TICHÝ L., CHYTRÝ M Statistical determination of diagnostic species for site groups of unequal size. J. Veg. Sci. 17: TICHÝ L JUICE, software for vegetation classification. J. Veg. Sci. 13: TICHÝ L., ROLEÈEK J., ZELENÝ M., CHYTRÝ New stopping rules for TWINSPAN. European vegetation survey, 16 th Workshop, Rome, Italy. March VALACHOVIÈ M Asplenietea trichomanis (Br.-Bl. in Meier et Br.-Bl. 1934) Oberd In: M. VALACHOVIÈ (ed.), Rastlinné spoloèenstvá Slovenska. 1. Pionierska vegetácia, pp Veda, Bratislava. VALACHOVIÈ M The role of cryptogams in differentiation of plant communities on the shady rocks in Western Carpathians. Thaiszia 5: WESTHOFF V., VAN DER MAAREL E The Braun Blanquet approach. In: WHITTAKER R.H. (ed.). Classification of plant communities. W. Junk, The Hague, pp. WILCZEK Z Fitosocjologiczne uwarunkowania ochrony przyrody Beskidu Œl¹skiego (Karpaty Zachodnie). Wydawnictwo Uniwersytetu Œl¹skiego 2418, Katowice, 223 pp.

21 91 Diagnostic-value-of-the-Polypodium-vulgare-L.-complex-in-rocky-communities 91 Wystêpowanie i wartoœæ diagnostyczna taksonów z kompleksu Polypodium vulgare L. w zbiorowiskach z klasy Asplenietea trichomanis (Br. Bl. in Meier & Br. Bl. 1934) Oberd w Sudetach i na ich Przedgórzu. Streszczenie Zbiorowiska naskalne z klasy Asplenietea trichomanis s¹ zbadane w stopniu niewystarczaj¹cym, a wiele z za³o eñ dotycz¹cych ich zmiennoœci florystycznej oraz siedliskowej pochodzi z pocz¹tkowego okresu badañ fitosocjologicznych. Badania te, prowadzone na niewielkim i wyrywkowo zgromadzonym materiale, mog¹ prowadziæ do b³êdnych lub uproszczonych wniosków, co powoduje problemy podczas identyfikacji zbiorowisk naskalnych w trakcie badañ terenowych lub póÿniejszego opracowania zebranego materia³u fitosocjologicznego. W œwietle wyników uzyskanych w trakcie analizy 298 zdjêæ fitosocjologicznych wykonanych na terenie Sudetów oraz ich Przedgórza, a tak e syntezy ostatnich opracowañ syntaksonomicznych, Polypodium vulgare nale y uznaæ za gatunek charakterystyczny dla rzêdu Androsacetalia vandelii Br.-Bl. in Meier et Br.-Bl.1934, nie zaœ dla klasy Asplenietea trichomanis (Br. Bl. in Meier & Br. Bl. 1934) Oberd. 1977, jak do tej pory przyjmowano w literaturze przedmiotu. Szczególnie czêsto paprotka pospolita wystêpuje w fitocenozach rozwijaj¹cych siê na œródleœnych i zacienionych ska³ach, zaliczanych do zwi¹zku Hypno-Polypodion vulgaris Mucina 1993 i dominuje w p³atach zespo³u Hypno- Polypodietum Jurko et Peciar 1963, w którym osi¹ga tak e najwy sze stopnie pokrycia (do 80%). Czêsto spotykana jest tak e w fitocenozach zespo³u Asplenio-Polypodietum Firbas 1924, gdzie jednak wystêpuje ze znacznie ni szym pokryciem (0,5 do 10%, wyj¹tkowo do 80%). Uzyskane wyniki sugeruj¹ wystêpowanie pomiêdzy tymi dwoma zespo³ami znacz¹cych ró nic florystycznych (podsumowanych w Tab. 6), których istotnoœæ wymaga jednak potwierdzenia w trakcie analizy wiêkszej iloœci danych z terenu ca³ej Europy Œrodkowej. Najwa niejszymi czynnikami, które warunkuj¹ wystêpowanie Polypodium vulgare w zbiorowiskach naskalnych w Sudetach i na ich Przedgórzu s¹ rodzaj pod³o a (ska³y bezwapienne) oraz zacienienie (z regu³y przekraczaj¹ce 60%). Na pod³o ach wapiennych oraz na stanowiskach nas³onecznionych paprotka pospolita wystêpuje znacznie rzadziej, a jej udzia³ w p³atach rzadko przekracza 1% pokrycia. Wystêpowanie P. mantoniae Rothm. zosta³o jak dot¹d potwierdzone w fitocenozach zaliczanych do zespo³ów Asplenio-Polypodietum i Asplenietum cuneifolii Gauckler 1954; potencjalnie mo e wystêpowaæ tak e w Woodsio-Asplenietum septentrionalis R. Tx Okreœlenie preferencji siedliskowych oraz pozycji syntaksonomicznej P. mantoniae wymaga dalszych badañ.