The association Saxifrago-Poetum compressae in Lower Silesia, south-western Poland

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1 Fragm. Flor. Geobot. 39(2): , 1994 The association Saxifrago-Poetum compressae in Lower Silesia, south-western Poland KRZYSZTOF ŚWIERKOSZ ŚWIERKOSZ, K The association Saxifrago-Poetum compressae in Lower Silesia, southwestern Poland. Fragmenta Floristica et Geobotánica 39(2): Kraków. PL ISSN x. ABSTRACT. Saxifrago-Poetum compressae (Kreh 1945) Genu et Leric 1957 is reported from eleven localities in Lower Silesia of south-western Poland and this is the first record of this association in our country. It belongs within the alliance Alysso-Sedion and the order Sedo-Scleranthetalia. Variability of the association is assessed using available comparative material from Central and Western Europe. This made it possible to distinguish four subassociations on the basis of different species composition and habitat conditions. Two of these, S.-P. arrhenatheretosum and S.-P. chenopodietosum, have been recognized in Lower Silesia. KEY WORDS: phytosociology, plant communities, grasslands, Lower Silesia, Poland K. Świerkosz, Museum of Natural History, Wroclaw University, ul. Sienkiewicza 21, PL , Wrocław, Poland INTRODUCTION Epilithic grassland associations of the class Sedo-Scleranthetea are poorly known in Poland. Matuszkiewicz (1982) showed the potential possibility of their occurrence in our country, but without mentioning any associations. In spite of the distinct impoverishment of the group of characteristic and differential species of the alliance Alysso-Sedion, it seems that at least part of grassland associations known from Europe occur also in western Poland. In particular this is true of the association Saxifrago-Poetum compressae (Kreh 1945) Gehu et Leric 1957, widespread in Lower Silesia, which I first observed during my studies in Wrocław (Świerkosz 1993), where it occurred in extremely impoverished form, making its correct identification difficult. In I found additional localities of this association in Lower Silesia making it possible to describe it in detail. I compared also the material accumulated during the present studies with data available from Central and Western European literature, which enabled preliminary conclusions to be drawn concerning the presence of lower syntaxa within the association. When describing the association I used the papers of Korneck (1977), Gehu (1961), Bornkamm (1961), Hilbig and Reichhoff (1977), Klimes (1986) and others. Taxa charac-

2 640 Fragm. Flor. Geobot. Ann. 39, Pars 2, 1994 teristic for particular units were distinguished following Korneck (1977), but papers by Hilbig and Reichhoff (1977) and Głowacki (1981) have also been consulted. METHODS Thirty-eight phytosociological releves were made in at eleven localities throughout Lower Silesia using the method of Braun-Blanquet (1964). Their areas ranged from 0.5 to 10 m 2. The resulting material is arranged in Tables 2 and 3. It represents the two subassociations distinguished in Lower Silesia. Table 1 contains basic climatic data for localities of the phytocoenoses of Saxifrago-Poetum compressae in Lower Silesia. Depending on the locality the annual precipitation ranges from 527 to 855 mm, mean annual temperature from 6.3 to 8.5 C, and the vegetation period varies from 193 to 227 days. Table 1. Climatic data for localities of Saxifrago-Poetum compressae in Lower Silesia. Station (altitude in m) Mean temp, of I( C) Mean temp, of VII ( C) Mean annual temperature ( Q Amplitude < C) Annual precipitation (mm/year) Vegetation period (days) Number of localities Wrocław (127) (621)* 227 2, 3, 4, 5, 6 Legnica (140) , 10 Głuszyca (469) ** Boguszów (595) ,8 * Świdnica; ** Walim; (Schmuck 1948, 1960; Walczak 1968,1970) RESULTS Association Saxifrago-Poetum compressae and its differentiation The characteristic species often dominating in patches of the association Saxifrago- Poetum compressae is Poa compressa. P angustifolia, a differential species of the association, occurs infrequently in Lower Silesia and, additionally, it is found in other phytocoenoses. In the study area I have not found another characteristic species of the association, Saxífraga tridactylites. The patches of Saxifrago-Poetum compressae form in most cases low and loose grassland of the two above mentioned grass species, sometimes with an admixture of Bromus tectorum, Arrhenatherum elatius and/or Dactylis glomerata. These are always accompanied by species of the class Sedo-Scleranthetea such as Potentilla argéntea, Sedum acre and Ceratodon purpureus which constitute the ground layer. The two following subassociations of Saxifrago-Poetum compressae are recognized in Lower Silesia. 1. Saxifrago-Poetum compressae (Kreh 1945) Gehu et Leric 1957 chenopodietosum albi (Klimes 1986) Swierkosz, comb, nova (Table 2) It occurs throughout Lower Silesia (Fig. 1) in ruderal habitats on the top and sloping

3 K. Świerkosz: Saxifrago-Poetum compressae in Poland 641 Fig. 1. Distribution of Saxifrago-Poetum compressae (Kreh 1954) Genu et Leric 1957 in Lower Silesia. 1: Saxifrago- Poetum compressae chenopodietosum albii (Klimes 1985) Swierkosz; 2: Saxifrago-Poetum compressae arrhenatheretosum elatioris (Genu 1961) Świerkosz. List of localities: 1 - Złotoryja (Legnica Prov.); 2 - Środa Śląska (Wroclaw Prov.); 3 - Wroclaw. 4 - Sokolniki near the massif of Ślęża (Wrocław Prov.); 5 - Stoszów near the massif of Ślęża (Wrocław Prov.); 6 - Radzików near the massif of Ślęża (Wrocław Prov.); 7 - Walim in Góry Sowie (Wałbrzych Prov.); 8 - Wałbrzych. 9 - Zamkowa Góra near Wałbrzych in Góry Wałbrzyskie; 10 - Wilcza Góra near Złotoryja (Legnica Prov.); 11 - Bardo Śląskie near Kłodzko (Wałbrzych Prov.). parts of walls, roofs of buildings, and river bank protections. Poa compressa is dominant, often with P. angustifolia. Of species of the alliance Alysso-Sedion only the differential species Asplenium ruta-muraria and Tortula muralis, are present. Of the order Sedo- Scleranthetalia two ergasio-epecophytes, Sempervivum tectorum and Sedum album, are found which have been known in Lower Silesia since the beginning of the century (Schube 1904). They may grow locally with a high constancy and abundance, though they occur at only limited number of localities. The subassociation is distinguished by a group of synanthropic species. In Lower Silesia these are Chenopodium album, Artemisia vulgaris, Plantago major, Stellaria media and Trifolium repens. Though much impoverished, the patches of the subassociation described from Lower Silesia are, with respect to their species composition, nearly identical to those described from similar ruderal habitats in Germany and the Czech Republic (Table 4). The subassociation is widespread throughout the range of Saxifrago-Poetum compressae. It was described under various names by Korneck (1977), Bornkamm (1961), Klimes (1986) and Hilbig and Reichhoff (1977). 2. Saxifrago-Poetum compressae (Kreh 45) Gehu et Leric 57 arrhenatheretosum elatioris (Gehu 61), subass. nova prov. (Table 3) This subassociation is much less frequent than the preceding one. It was found on

4 Table 2. Saxifrago-Poetum compressae (Kreh 1945) Gehu et Leric 1957 chenopodietosum albii (Klimes 1986) Swierkosz.

5 Localities - Zł - Złotoryja, Sr - Sroda Śląska, Wr - Wrocław, So - Sokolniki, St - Stoszów, R - Radzików, W - Walim, Wa - Wałbrzych, ZG - Zamkowa Góra near Wałbrzych. Table 2. Continued. Sporadic - D. Saxifrago-Poetum compressele: Chenopodium album 14 (1.2), Stellaria media 16 (r). Ch. and Sedo-Scleranthetalia and Sedo-Scleranthetea: Erodium D.00 cicutarium 1 (2.1), 13 (2.1), Hieracium pilosella 5 (+), Homalothecium sericeum00 19 (3.2), 20 (2.2), Cerastium arvense 20 (3.2), Sedum album 0 20 (2.1). Accompanying species: Achillea millefolium 14 (1.2), Ballota nigra 4 (1.1), Bryum caespiticium 17 (2.1), Campanula rotundifolia 15 (+), Cerastium fontanum subsp. triviale 16 (+), Chelidonium majus 3 (+), 19 (+), Cladonia chlorophaea 15 (1.1), 16 (1.1), Conyza canadensis 2 (+), 19 (1.1), Crepis tectorum 1 (1.1), 3 (1.1), Epilobium angustifolium 19 (+), Festuca rubra 5(1.1), Hieracium murorum 15 (+), Lamium album 3 (+), Lappula squarrosa 2 (+), Lepidium ruderale 3(1.1), Lolium perenne 16(1.1), Moehringia trinervia 19 (+), Poa annua 16 (r), Sisymbrium loeselii 18 (+), Urtica dioica 16 (r).

6 Table 3. Saxifrago-Poetum compressae (Kreh 1945) Géhu et Leric 1957 arrhenatheretosum elatioris (Géhu 1961) Swierkosz.

7 Table 3. Continued. Sporadic- Saxifrago-Poetum compressae arrenatheretosum: Daucus carota 16 (+). Accompanying species: Artemisia vulgaris 17 (+), 18 (+), Achillea millefolium 16 (1.1), 17 (1.2), Asplenium trichomanes 8 (+), Bilderdykia convolvulus 2 (1.1), Bryum caespiticium 16 (2.1), 18 (1.1), Campanula rotundifolia 11 (+), Cerasus avium c 6 (+), Festuca rubra 18 (1.2), Galium album 11 (+), 16 (1.2), G. verum 1(+), Hieracium murorum 3 (+), H. sabaudum 12 (+), H. umbellatum 4 (1.2), 5 (1.2), Leontodón hispidus 18 (1.2), Medicago lupulina 16 (1.2), 17(1.1), Melilotus officinalis 16 (+), 17 (1.1), Rubus caesius coll. 4 (+), Silene vulgaris 17 (1.2).

8 646 Fragm. Flor. Geobot. Ann. 39, Pars 2, 1994 basalt rocks of the "Wilcza Góra" Nature Reserve near Złotoryja in Legnica Province, and on the bridge head in Bardo Śląskie near Kłodzko in Wałbrzych Province. Differential species of the subassociation are those of the class Festuco-Brometea {Potentilla tabernaemontani, Sanguisorba minor, Scabiosa columbaria and Pimpinella saxífraga) and of the class Molinio-Arrhenatheretea {Arrhenatherum elatius, Daucus carota, Dactylis glomerata and Rumex acetosa). The core of the subassociation is formed by Poa compressa which, however, occurs here with lower coverage than in the preceding subassociation. Of species of the order Sedo-Scleranthetalia there occur Acinos arvensis and Scleranthus annuus subsp. polycarpos. Also species of the class Sedo-Scleranthetea are abundant. Variability of the association in Europe On the basis of material in Table 4, four subassociations of Saxifrago-Poetum compressae can be distinguished in Europe, which differ in floristic composition and habitat conditions. 1. Saxifrago-Poetum compressae subass. brometosum (Hilbig et Reichhoff 1977) Swierkosz, comb. nova. The differential species include Bromus sterilis, B. tectorum, Poa nemoralis and Geranium robertianum. This subassociation has two variants. The typical variant described by Korneck (1977) from West Germany and Hilbig and Reichhoff (1977) from East Germany, and the variant with Chenopodium album. Its differential species, such as Senecio vulgaris, Chenopodium album and Artemisia vulgaris, are associated with the subassociation chenopodietosum albi. This variant is known from France (Genu 1961) and the Czech Republic (Klimes 1986). The subassociation occurs mainly in ruderal habitats. 2. Saxifrago-Poetum compressae subass. arrhenatheretosum elatioris (Genu 1961) Swierkosz, subass. nova. prov. Recognition of this subassociation is based on the striking floristic similarity of the "community with Potentilla argéntea and P. verna" from the Ardennes (Genu 1961) and patches found in Lower Silesia (Table 3). Its differential species include Potentilla tabernaemontani, Pimpinella saxífraga, Scabiosa columbaria, Arrhenatherum elatius, Dactylis glomerata, Rumex acetosa, Ranunculus bulbosus, Daucus carota and Cytisus scoparius. The subassociation was found on basalt, limestone and stone walls. 3. Saxifrago-Poetum compressae subass. chenopodietosum albii (Klimes 1989) Swierkosz This is a form of the association tending towards synanthropic communities, to such an extent that it is sometimes included within the alliance Chenopodion albi (Bornkamm 1961; Korneck 1977). The differential species include Artemisia vulgaris, Chenopodium

9 K. Swierkosz: Saxifrago-Poetum compressae in Poland 647 album, Senecio vulgaris, Trifolium repens, Plantago major, Galinsoga parviflora, Stellaria media and Viola arvensis. 4. Saxifrago-Poetum compressae typicum Hilbig et Reichhoff 1977 This syntaxon is characterized by the absence of differential species of the remaining subassociations. There are only five records of its occurrence, all made on sandstone and porphyr in East Germany in the vicinity of Gera and Querfurt (Hilbig & Reichhoff 1977). The subassociations distinguished above often form intermediate forms which make their correct identification difficult. For this reason phytocoenoses of the association Saxifrago-Poetum compressae are known in the literature under different names (Table 4) with variation of the lower syntaxonomic units. DISCUSSION The association Saxifrago-Poetum compressae was described from ruderal habitats in France by Kreh in 1945 (Borkmann 1961). In subsequent years plant communities with similar species composition were described under various names from both synanthropic and natural habitats in various places on the continent. In France Genu (1961) described two communities (Table 4, columns 2 and 6) which he included in the order Festuco-Sedetalia of the class Festuco-Brometea. Local differences between them justify such an approach, but comparison with material from other regions of Europe makes them less distinct. Bornkamm (1961) analysed the roof vegetation in Gottingen and presented his rich relevé material in several tables recognizing five communities placed in four different classes. At least two of these associations, Poetum anceptis-compressae (Table 4, columns 9 and 10) and Sedo-Sempervivetum tectorum (Table 4, columns 11 and 12) display close similarity to the association Saxifrago-Poetum compressae. Hilbig and Reichhoff (1977) presented two synthetic tables of the association Saxifrago-Poetum from East Germany (Table 4, columns 4 and 13) - the typical variant and the variant with Promus tectorum. They regarded Promus tectorum, B. sterilis and Chelidonium majus as differential species of the latter. They also reported that species of the alliance Sisymbrion R. Tx., Lohm. et Preis 1950 and of the class Galio-Parietarion Riv.- Mart (i.e. alliance Tortulo-Cymbalarion in a more modern sense) were associated with that variant. They included the association Saxifrago-Poetum in the alliance Alysso- Sedion and order Sedo-Scleranthetalia. The same opinion was expressed by Korneck (1977) who presented a synthetic table of the association from south-western Germany (Table 4, column 3). The approach adopted in this paper makes it possible to identify the form of association described by him with the subassociation brometosum. Klimes (1986) published abundant relevé material from Moravia in the northern Czech Republic (Table 4, column 5). Considering the phytosociological affinity of the

10 648 Fragm. Flor. Geobot. Ann. 39, Pars 2, 1994 Table 4. Variability of Saxífrago-Poetum compressae (Kreh 1945) Géhu et Leric 1957 in Europe.

11 K. Swierkosz: Saxifrago-Poetum compressae in Poland 649 Table 4. Continued. Sporadic - Alysso-Sedion and Ch & D 0 0 Sedo-Scleranthetalia: Alyssum alyssoides 3 (11), Cerastium brachypetalum 1(20), Encalypta streptocarpa 00 2 (I), Festuca glauca 0 2 (I), Grimmia pulvinata 00 1 (40), Melica ciliata 0 1 (7), Minuartia hybrida 1 (13), Pleurochaete squarrosa 00 1 (7), Poa badensis 0 1 (7), P bulbosa 3 (4), Scleranthus annuus subsp. polycarpos 1 (17), Thlaspi perfoliatum 2 (I), Tortella tortuosa 13 (20), Veronica praecox 1 (13). In one table: 1: Holosteum umbellatum, 33; Cerastium semidecandrum, 20; Medicago minima, 13; Taraxacum laevigatum, 7; Valerianella locusta, 7. 5: Thymus serpyllum, 86; Festuca ovina, 29; Trifolium arvense, 14. 6: Polytrichum piliferum, 56; Rumex tenuifolius, 22.12: Euphorbia exigua, 66. Special characters used in the table -?: mosses and lichens not investigated; v: moss recorded in the locality, but only as present (cfr. Klimes 1986) Explanations I. Saxifrago-Poetum compressae arrhenatheretosum elatioris 1. Saxifrago-Poetum compressae, arrhenatheretosum elatioris, subass. nova prov. - Table 3 2. ass. a Potentilla argentea et Potentilla verna, Gehu (1961), France (intermediate variant between I and II) II. Saxifrago-Poetum compressae brometosum a) variant typicum 3. Saxifrago-Poetum compressae, Korneck (1977), West Germany

12 650 Fragm. Flor. Geobot. Ann. 39, Pars 2, 1994 Table 4. Continued. 4. Saxifrago-Poetum compressae var. with Bromus sterilis, Hilbig & Reichhoff (1977), East Germany b) variant with Chenopodium album 5. Sedo acri-poetosum compressae ass. nova, Klimes (1986), Czech Republic 6. ass. á Poa compressa et Saxífraga tridactylites, Genu (1961), France III. Saxifrago-Poetum compressae chenopodietosum albi 1. Saxifrago-Poetum compressae chenopodietosum albi - Table 2 8. Saxifrago-Poetum compressae chenopodietosum albi, Swierkosz (1993), Wroclaw 9. Poetum anceptis-compressae ass. nova subass. chenopodietosum albi, Borkamm (1961), Góttingen 10. Poetum anceptis-compressae ass. nova subass. typicum, Borkamm (1961), Góttingen 11. Sedo-Sempervivetum tectorum ass. nova, Borkamm 1961, Góttingen 12. Sedo-Sempervivetum tectorum ass. nova var. Sedum boloniense Borkamm (1961), Góttingen IV. Saxifrago-Poetum compressae typicum. 13. Saxifrago-Poetum compressae typicum, Hilbig & Reichhoff (1977), East Germany new association, he rejected the possibility of its inclusion in Saxifrago-Poetum compressae (an argument was its "more thermophilous character") and established a new association, "Sedo acri-poetum compressae". Interestingly, he regarded Poa compressa and P. angustifolia along with Sedum acre and Taraxacum officinale agg. as its characteristic species. He divided this association into two subassociations: subass. brometosum and subass. chenopodietosum albi. The association was included by Klimes (1986) in the alliance Convolvulo-Agropyrion of the class Agropyretea repentis, but the author provided no evidence to support his decision. The division proposed in the present paper resembles that of Klimes (1986), but is considerably modified, since only a synthetic approach to the material reveals that the striking differences in the frequency of occurrence of some species groups are actually less significant than they would appear at first glance. In its typical form described by Korneck (1977), Saxifrago-Poetum compressae (Kreh 1945) Genu et Leric 1957 undoubtedly belongs within the alliance Allysso-Sedion, the order Sedo-Scleranthetalia and the class Sedo-Scleranthetea as is shown by the species composition of the association in which species of the above syntaxa dominate (cf. Table 4, column 3). However, Poa compressa has a wide phytosociological tolerance, and on walls it also occurs as an accompanying species in the associations Asplenietum trichomano-rutae-murariae R. Tx and Cymbalarietum muralis Gors 1966 (Oberdorfer 1977; Swierkosz 1993). This species is also considered as a characteristic species of the class Festuco-Brometea (Matuszkiewicz 1982). Thus, the more abundant presence of Poa compressa is not a sufficient argument in favour of the distinct character of the association and its occurrence in Lower Silesia. The syntaxonomic position of Saxifrago-Poetum compressae described from Lower Silesia is not as clear as that of its forms known from areas situated to the west and south of Poland. However, its affinity to the class Sedo-Scleranthetea, whose characteristic species occur in nearly all stands of the association, is evident. The order Sedo-Scleranthetalia and the alliance Alysso-Sedion in Lower Silesia lacks many characteristic species. This testifies to the marginal character of the associations.

13 K. Świerkosz: Saxifrago-Poetum compressae in Poland 651 In order to characterize the association in detail, it is also necessary to consider specific habitat factors (no other grassland association of the class Sedo-Scleranthetea has been found on wall tops until now) and the characteristic combination of species of the association which includes, besides Poa compressa and P. angustifolia, numerous species of the alliance, order, and, in Poland, most of the class Sedo-Sclereanthetea. Their high systematic group values "D" and collective participation "G" (Pawłowski 1972) indicate the affinity of the association studied to the alliance Alysso-Sedion of the order Sedo- Scleranthetalia (Table 5). Table 5. Systematic group values "D" in Saxifrago-Poetum compressae (Kreh 1945) Genu et Leric 1957 in Lower Silesia. Group of species chenopodietosum arrhenatheretosum S(%) G(%) D S(%) G(%) D Ch. & D. of association, alliance and order Ch. Sedo-Scleranthetea Ch. Fetuco-Brometea Ch. Molinio-Arrhenatheretea Ch. Artemisietea Ch. Chenopodietea Accompanying species It appears that the most favourable conditions for the development of the association are in Przedgórze Sudeckie with its higher precipitation in comparison to that in Nizina Śląska, and a simultaneous long vegetation period (Walczak 1968). With decreasing precipitation or mean annual temperature phytocoenoses of Saxifrago-Poetum compressase become impoverished in species of the class Sedo-Scleranthetea which is distinctly visible in Wroclaw (climate with a high mean annual temperature but with relatively low precipitation), Walim and Wałbrzych (precipitation higher than in Przedgórze Sudeckie but low mean annual temperature) (Table 1). This makes it possible to formulate a conjecture on the subatlantic character of the association which is supported by its geographical distribution documented in the literature. Local habitat conditions of the occurrence of the phytocoenoses of the association were analysed by Klimeś (1986) in the Czech Republic and Bornkamm (1961) in Germany. In Lower Silesia phytocoenoses of Saxifrago-Poetum compressae occur on walls within a wide range of ph ( in H2O), with the optimum at ph ; on soils rich in all important nutrients, with a high content of CaC03 and skeletal parts (15-39%). Thus, these habitat conditions are close to those found on natural soils of the para-limestone soil type, which was pointed out by the authors cited above. Likewise, soils of the lithosol type formed on basalt eluvium should be regarded as rich. Their ph ranges from 4.5 to 5.0, and only phosphorus is deficient there (Wożakowska-Natkaniec 1986). According to Klimes (1986) a high deficiency of humidity, a shallow and quickly drying soil layer and high insolation also have a significant effect on the phytocoenoses of the association described. All these factors determine the habitat specificity of wall tops, roofs and crevices, similar to that of epilithic habitats. 22*

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