THE GENUS SCHISTIDIUM (GRIMMIACEAE, MUSCI) IN HUNGARY ...

Studia bot. hung. 39, pp. 27–88, 2008 

THE GENUS SCHISTIDIUM (GRIMMIACEAE, MUSCI) 

IN HUNGARY 

P. ERZBERGER 1 and W. SCHRÖDER 2 

1 Belziger Str. 37, D-10823 Berlin, Germany; erzberger@erzfisch.de 

2 Ludwigsstädter Str. 51, D-96337 Ludwigsstadt, Germany 

All available specimens of Schistidium (collected in Hungary) in Hungarian herbaria (BP, 

EGR and private herbaria) and collections of the first author (B) were revised. The 

following 15 taxa were found to occur in Hungary: Schistidium apocarpum, S. brunnescens 

subsp. brunnescens, S. brunnescens subsp. griseum, S. confertum, S. confusum, S. crassipilum, 

S. dupretii, S. elegantulum, S. flaccidum, S. helveticum, S. lancifolium, S. papillosum, S. 

platyphyllum (new to South-Eastern Central Europe), S. pruinosum, and S. robustum. All 

except S. apocarpum, S. brunnescens subsp. brunnescens, S. crassipilum, S. flaccidum and S. 

helveticum are new to Hungary. Descriptions, illustrations, distribution maps and a key are 

provided. Some taxonomic, bryogeographical and conservation issues are briefly discussed. 

Key words: bryophytes, distribution maps, habitat requirements, illustrated key, Schistidium 

platyphyllum 

INTRODUCTION 

Since the revision of the Schistidium apocarpum complex by BLOM 

(1996, 1998), his concepts – developed for the taxa found in Scandinavia – 

have been widely and rather successfully applied in other parts of Europe 

(BLOCKEEL and LONG 1998, GRIMS 1999, HOLZ 2000, CASAS et al. 2001, 

CORTINI-PEDROTTI 2001, SOTIAUX and VANDERPOORTEN 2001, SIEBEL 

2003, MÜLLER 2004, SMITH 2004, CASAS et al. 2006, BLOM et al. 2006, 

IGNATOV et al. 2006, IGNATOVA et al. 2006, MEINUNGER and SCHRÖDER 

2007). A recent molecular study seems to support the narrow species concept 

of Blom (GORYUNOV et al. 2007). As information on the distribution 

of taxa in a wider geographical area is accumulating, their ecological profile 

and additional knowledge about their variability is becoming more precise 

as well. At the same time, some new species have been described (BLOM 

1998, BLOM and LÜTH 2002). 

Studia Botanica Hungarica, 39, 2008 

Hungarian Natural History Museum, Budapest

28 ERZBERGER, P. and SCHRÖDER, W. 

After the preliminary treatment of Schistidium in the checklist of Hungarian 

bryophytes (ERZBERGER and PAPP 2004), the present work aims at 

closing this gap by the revision of a large number of specimens of this genus. 

Apart from answering the question of which taxa occur in Hungary, their 

distribution in the floristical regions of the country as defined by BOROS 

(1968) is assessed and mapped. For each taxon, selected morphological characters 

are described, based on observations in Hungarian material and the 

literature, and illustrated. The authors propose a determination key for 

Schistidium in Hungary. Habitat preference and distribution in Hungary 

are discussed and compared with the results from other countries. 

MATERIAL AND METHODS 

All specimens collected in Hungary and labelled Schistidium (or synonyms) in the 

bryophyte herbarium of the Hungarian Natural History Museum (BP), the herbarium of 

the Eszterházy College in Eger (EGR), and of some private herbaria (P. Ódor, Budapest, P. 

Szûcs, Almásfüzitõ) as well as the recent collections of the first author were examined by 

light microscopy and revised, apart from incomplete or non-fruiting material. Drawings of 

some morphological features were achieved with a Leitz drawing apparatus. In order to 

observe the pattern of the exothecial cells and stomata in the basal part of the capsule, 

sporophytes were emptied and well soaked in 2% KOH, sometimes by boiling for some 

seconds with a cigarette lighter to remove air bubbles, and then cut in half using a razor 

blade. In order to completely flatten the capsule wall, an additional longitudinal cut of 

about half of the capsule length was applied to the basal part of each half. 

In randomly selected specimens measurements of urn length and width were taken 

from emptied capsules soaked in 2% KOH. From these data, the ratio length/width was 

computed, and means and standard errors were also evaluated for each taxon. 

Distribution maps were prepared on the basis of the Central European mapping 

scheme (NIKLFELD 1971). Open circles represent collections before 1977, closed circles 

after that year. 

Frequency of taxa is estimated by the number of different growth sites as noted on 

the specimen labels. 

Nomenclature of bryophytes follows ERZBERGER and PAPP (2004) with the exception 

of Schistidium, Bryum moravicum Podb. (= B. laevifilum Syed) and B. kunzei Hornsch., 

where HILL et al. (2006) is followed. 

Studia bot. hung. 39, 2008

THE GENUS SCHISTIDIUM IN HUNGARY 29 

Taxonomic characters 

– Visible in stem cross sections of sporophyte bearing plants. In 

sterile shoots, the central strand may not fully be developed. 

– Two types of hair-point can be distinguished (BLOM 1996): 

(1) (Figs 3C, 9C, E, 11B 15C, 23C, 25B): cell walls clearly visible; transverse section 

canaliculate, flattened or lens-shaped; hair-point irregularly bent or flexuose in the dry 

state (but straight in S. dupretii and S. confusum). (2) (Figs 5E, 7B, E, 13C, 17E, 21C, 

30B, C, 32B, D): cell walls usually not clearly visible; transverse section lens-shaped or 

terete; hair-point usually straight. 

– Sinuose (e.g. in S. robustum, Fig. 32E) or esinuose, size in different 

parts of leaf (basal, central and apical) is also important. 

– Hemispherical or conical cell wall structures on leaf lamina (Figs 11E, 

25F, 30G, H), costa and margins (Figs 3B, 13B–D, 15C, 23B, C, E, 28B, 30B, C). Since in 

papillose species only a fraction of lamina cells bear papillae, papillosity is a property of the 

lamina, not of cells. Young leaves should be examined for papillosity. 

K+ – Upper leaves torn off the stem and placed in a drop of 2% KOH solution develop 

either yellow (in Confertum group) or red (in all other taxa) colours. 

– These important characters 

are best assessed in empty capsules soaked in 2% KOH solution, because after this procedure 

even old, fragile or damaged capsules assume their original shape, the capsule wall 

becomes less fragile and can thus be cut more easily, and the pattern of exothecial cells and 

stomata will also become much clearer. If only green, unripe capsules are available, these 

should be boiled for a short time in 2% KOH solution and then cut and emptied for examination 

of exothecial cells and stomata; however their shape and size will not correspond to 

that of ripe, emptied urns. 

Urn form may be the widest at the mouth (e.g. S. platyphyllum, Fig. 27D) or about the 

middle (e.g. S. elegantulum, Fig. 17F); measurements of length and width are important in 

absolute terms (e.g. for the differentiation between S. apocapum and S. lancifolium, compare 

Figs 3E and 23F) as well as their ratio. Measurements should be taken under a stereomicroscope 

without flattening the urn. 

The pattern of exothecial cells must be carefully observed in different parts of the 

capsule (see Fig. 1C–E), most relevant is the pattern in the lower half, especially at the base 

immediately above the seta, where the stomata may be observed. The pattern of seta surface 

cells is also sometimes useful (e.g. in the Rivulare group, Fig. 27G, 28F; BREMER 1980). 

In some taxa, ripe emptied urns may show a very fine striolation due to the arrangement 

of exothecial cells in vertical rows (Fig. 1). Transverse sections of the capsule wall 

(Fig. 1A) show an exothecium of even thickness, as in the capsule walls of Grimmia species 

(MAIER 2004). Sometimes (e.g. in S. dupretii) striolae develop due to the fact that long narrow 

exothecial cells, alternating with shorter cells, collapse (Fig. 15I, J). These striolae are 

not comparable to the plicae in the urns of other genera, e.g. Orthotrichum (MAIER 2004). 

– Orientation of peristome teeth should be observed in freshly 

deoperculate capsules. Peristome teeth may be patent to spreading with ascending tips 



(e.g. S. apocarpum, Fig. 3E) to squarrose-recurved (e.g. S. confusum, Fig. 11G). Apart from 

colour and papillosity, the size and shape of peristome teeth is important, as is the amount 

of perforation by pits and holes in cribrose peristome teeth. 

B 

C 

D 

F 

A 

E 

G 

Capsule walls without ( ) and with striolae ( ). = transverse section of capsule 

wall (exothecium stippled); = exothecial cells in Schistidium apocarpum; = exothecial 

cells in S. crassipilum at top of urn / in mid urn / at base of urn; , = exothecial cells in S. 

lancifolium in upper/lower part of urn. Scale bar: 200 μm. [Del. Schröder] 

RESULTS AND DISCUSSION 

According to the revision of all available herbarium material presented 

here, in Hungary the genus Schistidium comprises fifteen taxa, which can 

be grouped in the following informal infrageneric categories (BLOM 1998): 

Rivulare group: S. platyphyllum (Mitt.) H. Perss. subsp. platyphyllum; 

Apocarpum group, Apocarpum subgroup: S. apocarpum (Hedw.) 

Bruch et Schimp., S. lancifolium (Kindb.) H. H. Blom; 

Apocarpum group, Strictum subgroup: S. confusum H. H. Blom, S. 

papillosum Culm., S. pruinosum (Wilson ex Schimp.) G. Roth; 



Robustum group: S. dupretii (Thér.) W. A. Weber, S. robustum (Nees 

et Hornsch.) H. H. Blom; 

Confertum group: S. confertum (Funck) Bruch et Schimp., S. flaccidum 

(De Not.) Ochyra; 

Atrofuscum group: S. brunnescens Limpr. subsp. brunnescens, S. brunnescens 

subsp. griseum (Nees et Hornsch.) H. H. Blom, S. crassipilum H. H. 

Blom, S. elegantulum H. H. Blom, S. helveticum (Schkuhr) Deguchi (= S. singarense 

(Schiffn.) Laz.). 

Historical aspects 

The oldest specimens of Schistidium in the bryophyte collections of 

the Hungarian Natural History Museum (BP) date from the second half of 

the 19th century. Two are without date (S. apocarpum s. str.: “Magyarhon 

virányából Hazslinszky Frigyes, Grimmia apocarpa L. Ágasvár Mátra l. 

Borbás” BP 5758; S. crassipilum:“Grimmia apocarpa Kalkfelsen b. Tapolcza; 

ex herbario Frid. Hazslinskyi” BP 5699). The oldest specimens with a date 

are S. crassipilum from 1870 (BP 5667 Herbar. Musei Nat. Hungar. Budapest 

Specimen in herb. Simonkai L. inter Grimmia apocarpa insertum erat 

“In rupestribus montis ‘Kis Eged’ prope Agriam. Comitat. Heves in Hungaria. 

6. Apr. 1870 Vrabélyi”), and S. elegantulum from 1886 (BP 5624 “Ex 

herbario C. Schilberszky, Farkasvölgy, mészkõ [18]86.V.2”). Among the 

older collections there are some by Á. Degen from 1901 and L. Simonkai 

from 1904. Table 1 lists the collectors, the period of collection (of Schistidium 

specimens), and the number of specimens (corrected for duplicates). 

In his bryophyte flora of the surroundings of Budapest and the Pilis Mts 

SZEPESFALVI (1941) treats Schistidium under Grimmia and mentions 2 species: 

S. flaccidum (as G. flaccida) and S. apocarpum s. l. (as G. apocarpa) with 

three varieties: var. brunnescens,var.conferta, and var. gracilis. While his specimens 

of S. flaccidum and var. brunnescens could be confirmed in the present 

revision, the plants labelled var. conferta proved to be S. apocarpum s. str., S. 

brunnescens subsp. brunnescens, orS. cf. crassipilum, but not S. confertum. 

We did not see any specimens collected by Szepesfalvi labelled var. gracilis. 

In an earlier paper SZEPESFALVI (1935) stated that S. brunnescens was 

rare in the surroundings of Budapest and in Hungary, thus contradicting 



PÉTERFI (1916), who held the opposite opinion. It is interesting that Péterfi 

considered the possibility that plants named S. confertum growing on limestone 

rocks might prove to be S. brunnescens – and that in our revision this 

turned out to be true. 

BOROS’s (1968) taxonomic treatment is similar to that of SZEPESFALVI 

(1941), he lists some more forms (f. irrigata,f.epilosa,f.laxum = f. gracile), 

varieties (var. nigrescens) and subspecies (subsp. conferta,subsp.brunnescens) 

of S. apocarpum s. l. His f. irrigata was revised to S. apocarpum s. str.; under f. 

epilosa many different taxa were found, some belonging to different genera 

(S. apocarpum s. str., S. brunnescens subsp. brunnescens, S. crassipilum, S. helveticum, 

S. lancifolium, Didymodon acutus, D. insulanus, D. luridus, D. rigidulus); 

similarly “f. gracile” was found on specimen labels of S. crassipilum, 

S. apocarpum and S. lancifolium. Some specimens of S. crassipilum and S. apocarpum 

were labelled “var. nigrescens” by Boros. Specimens labelled “ssp. 

conferta” by Boros proved to be S. crassipilum or S. brunnescens subsp. brunnescens. 

However, plants that he named subsp. brunnescens consistently 

could be referred to S. brunnescens subsp. brunnescens. 

ORBÁN and VAJDA (1983) treat all taxa of Schistidium as subspecies, 

varieties or forms of S. apocarpum, even S. flaccidum, in this respect adopting 

the concept of LOESKE (1930). As for specimens collected by Vajda and 

bearing various subspecific labels, roughly the same applies as for Boros’s 

collections: his f. irrigata and S. rivulare proved to be S. apocarpum, subsp. 

gracile was revised to S. crassipilum and S. apocarpum, but one specimen (BP 

67976) labelled Grimmia apocarpa var. conferta could be confirmed as S. 

confertum, and S. brunnescens was also consistently confirmed. 

In Figure 2 the number of specimens of Schistidium collected in the 

years between 1870 and 2007 (including the collections of the first author) 

is depicted, clearly showing two periods of collecting activity with respect 

to Schistidium in Hungary: the first between 1905 and 1965, with major contributions 

by Boros, Degen, Szepesfalvi, Szurák and Vajda (compare Table 1), 

and the second between 1985 and 2007, with major contributions of Rajczy, 

Papp and the first author. In the distribution maps, collections from these 

two periods are represented by different symbols (open circles for the older 

collections, closed circles for the recent ones). 



Bryological activity in Hungary – number of specimens of Schistidium (without 

duplicates) collected between 1870 and 2007 

Collectors of Schistidium from Hungary in BP, EGR, and B 

Collector Period or year of collection No. of specimens 

Balanyi, L. 1959 1 

Bán, E. 1934 1 

Borbás, V s.t. 1 

Boros, Á. 1917–1968 259 

Degen, Á. 1901–1924 15 

Erzberger, P. 1991–2008 179 

Felföldy, L. 1952–1956 6 

Gelencsér, I. 1954 1 

Glatz, W. 1936 1 

Gyõrffy, I. 1921–1928 2 

Héder, I. 1953 1 

Igmándy, J. 1924–1941 4 

Károlyi, Á. 1947–1956 10 

Kenyeres, J. [= “uxor mea”, Boros’s wife] 1956 1 

Kis, G. 1984 1 

Kovács, M. 1957 1 

Lengyel, G. 1909–1921 2 

Ódor, P. 1996–2001 4 

Ötvös, L. 1933 1 

Papp, B. 1989–2008 39 

Pénzes, A. 1947 1 

Péter, B. 1936 1 



(continued) 

Collector Period or year of collection No. of specimens 

Pócs, T. 1950–2007 7 

Polgár, S. 1932–1935 3 

Prágai, Z. 1958 1 

Priszter, Sz. 1953 1 

Rajczy, M. 1976–1994 21 

Redinger, K. 1931 3 

Sándor s.t. 1 

Sass-Gyarmati, A. 2001 1 

Simonkai, L. 1904 2 

Schilberszky, K. 1885 1 

Szemes, G. 1947 1 

Szepesfalvi (Szurák), J. 1911–1937 61 

Szûcs, P. 2004–2006 14 

Timár, L. 1914–1952 12 

Timkó, Gy. 1924 1 

Vajda, L. 1942–1971 93 

Vidéki, R. 2006 1 

Visnya, A. 1931–1953 9 

Vrabélyi, M. 1870 1 

Wagner, J. 1927 1 

Descriptions, selected diagnostic features 

The descriptions are based on BLOM (1996, 1998), HOLZ (2000), 

WEIBULL (2006), and our own observations. The data on the urn length/ 

width ratio of Hungarian material include (minimum) mean ± 2s.e. (maximum), 

sample size. Vertical distribution: elevation of lowest and highest location 

according to specimen labels. For convenience, in this paragraph, 

the taxa will be treated in alphabetical order. 

(Hedw.) Bruch et Schimp. 

(Figs 3, 4) 

Plants medium-sized to large, forming lax or dense tufts or mats. Central strand absent 

or narrow and indistinct. Hair-point 0–0.8 mm, fine, thin and ± flexuose, from not to 

longly decurrent. Abaxial side of costa and leaf margin papillose, distantly denticulate in 

upper part of leaf, rarely smooth (longipilose plants). 



A B C 

E 

D 

F 

H 

G 

Schistidium apocarpum. =leaves; , =leafapices; = perichaetial leaf; =capsule; 

, =exothecialcellsinmidurn/atbaseofurnandsetasurfacecells;H =peristome 

tooth. Scale bar: A, D, E – 2 mm; B – 800 μm; C, H – 400 μm; F, G – 200 μm. [Erzberger 

12084, del. Erzberger] 



Lamina without papillae, smooth, predominantly unistratose, but often with bistratose 

patches or striae in upper half. Lamina cells from slightly to strongly sinuose, in upper part 

8–10 μm wide. K+ red. 

Sporophytes common, deeply immersed. Urn dark red, widest at mouth, length/width 

ratio (1.2–)1.3–1.6(–2.0) (BLOM 1996), in Hungarian material (1.0–)1.51 ±0.06(–2.0), 

n = 65. Exothecial cells predominantly isodiametric and short transversely rectangular, but 

often with patches of elongate and rectangular cells. Stomata (4–)8–12(–18) per urn. Peristome 

teeth (350–)400–710 μm, red, patent to spreading with ascending tips, longly tapering 

to a fine point, in lower part ± semi-perforated, in central and upper parts from almost 

entire to strongly perforated with small oval cracks in submarginal rows. 

Normally easily identified by the combination of denticulate leaf tips and isodiametric 

exothecial cells, but longipilose forms occur with smooth leaf margin and costa. 

: On often shaded or moist, but sometimes also exposed or dry siliceous (andesite, 

porphyric rock, aleurolitic slate, sandstone, basalt) or more rarely calcareous rocks, 

often near streams or springs, but also on walls, even on loess. 

: Amphidium mougeotii, Brachythecium populeum, Ceratodon 

purpureus, Didymodon rigidulus, D. sinuosus, D. vinealis, Grimmia hartmanii, Hedwigia 

ciliata var. ciliata, Hypnum cupressiforme, Lejeunea cavifolia, Orthotrichum urnigerum, 

Orthotrichum sp., Schistidium crassipilum, S. dupretii, S. elegantulum, S. lancifolium, S. pruinosum, 

Taxiphyllum densifolium, Tortella tortuosa, Tortula ruralis. 

: 83–900 m a.s.l. 

: : Comit. Abaúj-Torna. In rupibus 

irrigatis rivi Ósva-patak (pars superior) prope pag. Telkibánya, montes Sátor-hegység, 

17.09.1959, leg. L. Vajda, BP 63350; 

Distribution of Schistidium apocarpum 



: Comit. Abaúj-Torna. In rupibus calcareis umbrosis silvat. ad 

fontem vallis Kopolya prope Szinpetri, 200 m, 19.06.1953, leg. Á. Boros, BP 110955; 

: Comit. Abaúj-Torna et Borsod. In rupibus calcar. alvei rivi vallis 

Telekes-völgy prope Perkupa, 180 m, 24.06.1953, leg. Á. Boros, BP 110966; : 

Felsõtárkány, Lök-völgy, aleurolite slate rocks at road, N 48° 00’ 25.2”, E 20° 27’ 

57.8”, 337 m, 06.04.2007, leg. P. Erzberger and T. Pócs, herb. Erzberger, (B) 12108; 

: Sorkõ/Sombokor, 800–860 m, 02.06.2001, leg. B. Papp, BP 170576; 

: Radiolarit-Felsen an der Straße ca 1 km S Bátor, 230 m, 26.03.2008, leg. P. 

Erzberger and T. Pócs, herb. Erzberger, (B) 12842a; : Cserhátszentiván, 

Cserkuti patak, 05.08.2002, leg. P. Erzberger, herb. Erzberger, (B) 7312; 

: Comit. Pest. In saxis andesit. vallis “Bükkös” ad “Sikáros” in 

Fageto, 350 m, 05.07.1933, leg. J. Szepesfalvi, BP 5859; : Comit. 

Hont. In rupibus andes. silvat. alvei rivi Kemence-patak infra Barsi-bükk pr. Kemence, 

300 m, 12.10.1958, leg. Á. Boros, BP 110874; Kemence-patak, on andesite boulder near the 

water, N 47° 59’ 07.8”, E 18° 58’ 20.7”, 384 m, 05.04.2007, leg. P. Erzberger, herb. Erzberger, 

(B) 12084; : Comit. Pest. In petrosis andesiticis in ripa rivi vallis 

“Apátkuti völgy” prope Visegrád, 200–300 m, 24.04.1924, leg. Á. Boros, BP 110905; 

: Szendehely-Katalinpuszta, Lósi-patak, sandstone near stream, N 47° 51’ 

01.1”, E 19° 06’ 24.4”, 180 m, 09.04.2007, leg. P. Erzberger and P. Szûcs, herb. Erzberger, (B) 

12163; : Budapest: János-hegy, mészkövön, 21.04.1924, leg. J. Szurák, 

BP 5628; : Comit. Esztergom. In cava rupe in decliv. sept. silvat. 

montis Nagyteke-hegy prope pag. Süttõ, 300 m, 17.05.1941, leg. Á. Boros, BP 110850; 

: County Veszprém. On rock in forest on the southern slope of 

Szent György-hegy at Hegymagas, 300 m, 13.08.1999, leg. B. Papp, BP 166930; 

: Tátika, 30.07.2000, leg. P. Erzberger, herb. Erzberger, (B) 6258; : 

: Querceto-Luzuletum sub font. Szénégetõ-forrás, supra pag. Velem, 13.07.1954, 

leg. T. Pócs et I. Gelencsér, BP 58482; : Comit. Baranya. In rupibus 

irrigatis vallis Csatornavölgy prope pag. Vasas, 26.07.1952, leg. L. Vajda, BP 5619; 

: Comit. Baranya. In rupibus calcareis umbrosis in sylva Tenkes-erdõ in 

monte Tenkes prope pag. Bisse, 300 m, 07.08.1999, leg. B. Papp, BP 166507; 

: Comit. Csongrád. Sándorfalva. Tegmen lapidarium (andesiticum) 

aggeri ad Sajtos, 83 m, 09.08.1957, leg. L. Timár, EGR; : In cippo in sepultura 

Felsõ-temetõ prope Hajdunánás, com.: Hajdu, 02.04.1941, leg. J. Igmándy, BP 

111365; : Comit. Ung. In muris pontis viae ferreae ad Záhony, 

100 m, 07.09.1925, leg. Á. Boros, BP 111357. 

Limpr. subsp. 

(Figs 5, 6) 

Plants small, forming dense tufts, brownish. Central strand mostly broad and distinct. 

Hair-point absent or short in leaves below perigonia, in subperichaetial and 

perichaetial leaves 0.15–1.0 mm, coarse and terete, but flattened towards insertion, not or 

shortly decurrent, but embracing parts of the upper lamina. 



A B E 

F C D 

I 

H 

G 

Schistidium brunnescens subsp. brunnescens. = leaves; = subperichaetial leaves; 

, = perichaetial leaves; = perichaetial leaf apex; = capsules; , = exothecial cells 

in lower third / at base of urn; I = part of peristome. Scale bar: A–D, F – 2 mm; E – 800 μm; 

I – 400 μm; G, H – 200 μm. [A–C, E–F, I: Erzberger 12198; D: EGR Vajda 1946 (Pomáz); 

H: BP 37650, A–F, H: del. Erzberger, G: del. Schröder] 



Costa and leaf margins smooth. Lamina smooth, in lower part unistratose, in upper part 

with bistratose spots. Upper lamina cells rounded, 6–9 μm wide, lower cells wider and 

slightly sinuose, basal marginal cells sometimes subhyaline, with slightly thickened 

cross-walls. K+ red. 

Sporophytes frequent, mostly shallowly immersed. Urn orange to reddish-brown, often 

finely striolate, widest below middle, length/width ratio (1.3–)1.4–1.6–1.8 (BLOM 

1996), in Hungarian material (0.94–)1.34 ±0.04(–1.93), n = 91. Exothecial cells elongate, 

especially in basal part, at least some up to 50 μm (often up to 80 μm) long, arranged in a 

regular pattern. Stomata absent. Peristome teeth 220–340 μm, orange, in central and 

upper parts mostly strongly perforated to cribrose. 

Variability: The exothecial cells in the lower part of the urn sometimes only reach 

(50–)55 μm, but in other samples they attain 75–80 μm. 

The brown, dense cushions with hair-points only around the sporophytes give this 

plant a very characteristic appearance. However, some forms of S. crassipilum can look 

quite similar, and in that case the strongly elongate exothecial cells (Fig. 5G, H, compare 

with Fig. 13I, J), and the shorter and wider urn of S. brunnescens are diagnostic. Confusion 

could also occur with forms of S. helveticum lacking the typical black colour, but again the 

pattern of exothecial cells is quite different. For the difference between subsp. brunnescens 

and subsp. griseum see the note under the latter. 

: S. brunnescens subsp. brunnescens grows nearly exclusively on exposed calcareous 

rocks (limestone, dolomite), and is a very typical member of the bryophyte vegetation in 

calcareous grasslands. It was found associated with the lichen Fulgensia fulgens (det. V. Otte). 

: Bryum argenteum, Didymodon rigidulus, D. vinealis, Grimmia 

dissimulata, G. orbicularis, G. pulvinata, Grimmia tergestina, Pseudocrossidium revolutum, 

Pseudoleskeella catenulata, Schistidium crassipilum, S. helveticum, Tortella inclinata, Tortula 

crinita, Trichostomum sp. 

: 150–780 m a.s.l. 

: : Comit. Gömör. In rupibus 

calc. montis “Baradla-tetõ” prope pag. Aggtelek, 350 m, 03.06.1928, leg. Á. Boros, BP 

111757; : Comit. Borsod. In rupibus calcar. montis Bélkõ prope Bélapátfalva, 

500–780 m, 03.09.1959, leg. Á. Boros, BP 110944; Lehmschieferfelsen an der 

Straßenböschung zwischen Bükkzsérc und Felsõtárkány, ca 400 m, mit S. crassipilum, 27.03. 

2008, leg. P. Erzberger and T. Pócs, herb. Erzberger, (B) 12886; : 

Glaukonit-Sandsteinfelsen “Noé szõllõje” im Ort Istenmezeje, 230 m, 26.03.2008, leg. P. 

Erzberger and T. Pócs, herb. Erzberger, (B) 12861; : Comit. Pest. In 

rupibus dolomit. apricis montis Vár-hegy prope Csõvár, 350 m, 06.05.1953, leg. Á. Boros, 

BP 111748; : Comit. Pest. In monte Somlyó prope Fóth, 21.05.1901, 

leg. Á. Degen, BP 37650; : Comit. Pest. In rupibus dolomiticis merid. 

montis Nagykevély prope pag. Pilisborosjenõ, 500 m, 25.03.1947, leg. Á. Boros, BP 111750; 

Hung. centr., com. Pest. In rupibus calcareis supra pag. Pomáz, 31.03.1946, leg. L. Vajda, 

EGR; : Budakalász, comit. Pest, in monte “Monalovác” ad saxa calcarea, 

250 m, III.1911, leg. J. Szurák, revid. M. Péterfi, BP 5645; : Gyermely, 



Kecske-kõ, N 47° 35’ 52.9”, E 18° 36’ 57.9”, 250 m, 11.04.2007, leg. P. Erzberger, herb. 

Erzberger, (B) 12198; : Comit. Fejér. In rupibus dolomit. montis adv. 

“Pap irtás” prope Csákberény, 350–400 m, 26.04.1936, leg. Á. Boros, BP 111730; 

Comit. Fejér. In rupibus dolomiticis apricis montis Szóló-kõ prope Csákvár, 200–240 m, 

21.03.1937, leg. Á. Boros, BP 111726, EGR; 

Distribution of Schistidium brunnescens subsp. brunnescens 

: Comit. Veszprém. In dolomiticis montis Móroc-tetõ prope Várpalota, 

500 m, 08.04.1951, leg. Á. Boros, BP 111693; : Comit. 

Zala. In rupibus basalt. tophac. merid. montis Csucs-hegy prope Tihany, 200 m, 

14.11.1962, leg. Á. Boros, BP 111010; : Comit. Baranya. In calcareis 

montis Harsányi-hegy ad Nagyharsány, 200–400 m, 22.03.1925, leg. Á. Boros, BP 111385. 

subsp. (Nees et Hornsch.) H. H. Blom 

(Figs 7, 8) 

Differs from subsp. brunnescens in the following characters. Plants larger, mediumsized. 

Central strand of few cells or absent. Hair-point not distinctly longer in subperichaetial 

leaves, more finely spinulose, 0.25–0.9 mm. Leaves of different shape (Figs 5A 

and 7A), often with ridge-like plicae (Fig. 7F, G), costa stronger (58–90 μm wide versus 

45–60 μm wide in central part of subsp. brunnescens), 3–4 stratose in upper and 5–6 stratose 

in lower part (Fig. 7F, G). 

Sporophytes sparse. Urn length/width ratio 1.4–1.6–1.9 (BLOM 1996), in Hungarian 

material (1.3–)1.48 ±0.08(–1.6), n = 7. Peristome teeth 190–320(–370) μm, orange, less 

perforated than in subsp. brunnescens (Figs 5I, 7I). 

: Exposed dry calcareous rocks. 



A B C 

F 

E 

G 

D 

H 

I 

Schistidium brunnescens subsp. griseum. = leaves; = leaf apex; = subperichaetial 

leaf; = perichaetial leaves; = perichaetial leaf apex; , = transverse sections 

in upper/lower part of leaf (showing bistratose ridges near margin: ); =capsule; = peristome 

tooth. Scale bar: A, C, D, H – 2 mm; B, E – 800 μm; I – 400 μm; F, G 200 μm. 

[Erzberger 12147, A–E, H, I: del. Erzberger; F, G: del. Schröder] 



: Grimmia orbicularis, G. tergestina, Orthotrichum sp. 

: 300–826 m a.s.l. 

Distribution of Schistidium brunnescens subsp. griseum 

Selected specimens examined: : Répáshuta, Három-kõ, auf exponiertem 

Kalkstein, N 48° 03’ 41.0”, E 20° 28’ 54.1”, 826 m, 08.04.2007, leg. P. Erzberger, herb. 

Erzberger, (B) 12147; : Gipfel des Pilis-Berges, offener Felsrasen, auf 

exponiertem Kalkstein, N 47° 41’ 16.5”, E 18° 51’ 56.6”, 750 m, 12.04.2007, leg. P. Erzberger, 

herb. Erzberger, (B) 12204; : Com. Fejér. In rupibus calcareis siccis 

montis Csókahegy prope pag. Mór, montes Vértes, 01.07.1951, leg. L. Vajda, EGR, BP 5668. 

(Funck) Bruch et Schimp. 

(Figs. 9, 10) 

Plants small, glossy, usually forming dense cushions. Central strand narrow or sometimes 

absent in sterile shoots. Hair-point short (0–0.45 mm), weak and flattened, straight or 

irregularly bent, not decurrent, sharply and strongly spinulose-denticulate with erecto-patent 

to squarrose spinulae. Costa and leaf margins smooth. Lamina smooth, partly bistratose in 

upper and central parts. Upper lamina cells rounded, 5–8(–9) μm wide, lower cells wider 

and slightly sinuose, basal marginal cells often subhyaline, forming a rectangular alar group 

composed of several rows of square to shortly rectangular cells with thickened cross walls. 

K+yellow. 

Sporophytes common, shallowly or deeply immersed. Urn orange-yellow, length/ 

width ratio 0.9–1.4–1.7 (BLOM 1996), in Hungarian material 1.29, n = 1. Exothecial cells 

predominantly elongate. Stomata 3–8 per urn. Peristome teeth 230–320 μm, orange, 

strongly perforated with long and ± wide cracks along the median line, sometimes with 

prongs split off at margins, coarsely papillose with rather distant, short papillae with broad 

basal part (appearing pebble-like). 



A 

D E F G 

D 

B 

C 

H 

K 

I 

J 

M 

L 

Schistidium confertum. , = leaves; , , = leaf apices; F = perichaetial leaves; 

, = perichaetial leaf apices; , = capsules; , = exothecial cells in lower third / at 

base of urn; = peristome tooth (showing papillae, in part). Scale bar: A, D, F, I, J – 2 mm; 

B, G – 800 μm; E, H, K – 400 μm; C, L, M – 200 μm. [A–C, K, L: EGR Vajda 1956 

(Börzsöny, Hollókõ); D–I: BP 67976 Vajda 1962 (Börzsöny, Hollókõ); J, M: BP 111013 

Vajda 1952 (Mátra, Disznókõ), del. Erzberger] 



The spinulose, flattened hair-point with visible lumina is quite unique to this species, 

it is not likely to be confused with any other Schistidium species in Hungary. The rather 

short urn with strongly perforated peristome, and the preference of siliceous substrata are 

further characteristics. 

: In Hungary, S. confertum grows exclusively on dry, andesitic rock. In Scandinavia, 

it grows on dry, ± exposed siliceous or base-rich rocks (BLOM 1996, 1998). It prefers 

acid rocks in Catalonia (CASAS et al. 2001). 

: 600–750 m a.s.l. 

Distribution of Schistidium confertum 

Selected specimens examined: Mátra Mts 8186/1 Com. Heves. In rupibus andesiticis 

siccis montis Disznókõ prope pag. Parádfürdõ, 750 m, 24.04.1952, leg. L. Vajda, BP 

5608 and 111013 (sub Grimmia plagiopodia); : Comit. Nógrád. In 

rupibus andesiticis siccis montis Hollókõ, prope pag. Perõcsény, montes Börzsöny, leg. L. 

Vajda, 30.05.1962, BP 67976 (sub Grimmia apocarpa var. conferta); Comit. Nógrád. In 

rupibus andesiticis siccis cacuminis montis Hollókõ prope pag. Kemence, montes Börzsöny, 

21.04.1956, leg. L. Vajda, EGR. 

H. H. Blom 

(Figs. 11, 12) 

Plants medium-sized, forming rather loose tufts of variable colour (olivaceous, brownish 

or blackish). Central strand wide and distinct. Hair-point 0–0.65 mm, narrow throughout 

or sometimes ± abruptly widened at insertion, not coarse but stiff and ± straight, 

from not to longly decurrent, irregularly spinulose-denticulate with short, blunt spinulae. 



A B C G 

D 

E 

F 

J 

J 

I 

Schistidium confusum. = leaves; = leaf apex; C = perichaetial leaf; =lamina 

cells in lower third of leaf; E = papillae and lamina cells at abaxial side of leaf near 

costa; = capsule (moist); perichaetium with capsule (dry) showing reflexed peristome 

teeth; = exothecial cells at lower third / at base of urn; = part of peristome. 

Scale bar: A, C, F, G – 2 mm; B, H, J – 400 μm; D, E, I – 200 μm. [Erzberger 12896, del. 

Erzberger] 

H 



Abaxial side of costa with scattered low papillae in upper part; leaf margins papillosedenticulate 

in upper part, more rarely smooth, broadly and strongly recurved throughout 

or in upper 3/4–4/5 of leaf length, ± bistratose. Lamina with scattered dorsal papillae, 

particularly near costa, and mostly with few ventral papillae, unistratose with bistratose 

spots and striae in upper part. Lamina cells in upper part rounded, slightly sinuose, 8–11 μm 

wide, in lower part wider and sinuose to nodulose, basal marginal cells with slightly 

thickened crosswalls. K+ red. 

Sporophytes abundant, shallowly immersed. Urn red-orange-brown, glossy, length/ 

width ratio (1.3–)1.5–1.9–2.3(–2.5) (BLOM 1996), in Hungarian material 1.78–1.83, n = 2. 

Exothecial cells predominantly isodiametric and transversely rectangular, but also rectangular, 

forming an irregular pattern. Stomata 4–9 per urn. Peristome teeth 310–420 μm, 

squarrose-recurved, red, gradually tapering to a fine point, from entire to strongly and irregularly 

perforated in central part, densely papillose. 

In Hungary, S. confusum is the only member of the Strictum subgroup that grows on 

calcareous substrates, whereas S. papillosum and S. pruinosum grow on siliceous rocks. From 

those species, S. confusum differs by its less papillose lamina (Fig. 11E), with more papillae 

on the abaxial, fewer on the adaxial side of the leaf (S. pruinosum is strongly papillose on 

both sides: Fig. 30G, H). S. papillosum is similar in this respect (usually more papillae on 

the abaxial side than on the adaxial side), but differs by the lack of a distinct central strand 

and the peristome teeth erecto-patent (Fig. 25G, but squarrose when old). Other differences 

between S. confusum and S. pruinosum include the wider and more sinuose lamina cells 

(Figs 11D, 30F) and the narrower perichaetial leaves of the former species (Figs 11C, 30D). 

:InHungary,S. confusum was found on exposed concrete adjacent to dolomite. 

In Scandinavia, it grows in similar situations, on exposed calcareous substrates (BLOM 1996). 

: None. 

: 250 m a.s.l. 

Distribution of Schistidium confusum 



Specimen examined: Buda Mts: 8580/1 Buda: Sas-hegy, on exposed concrete adjacent 

to dolomite, N 47° 28’ 59.5”, E 19° 01’ 09.3”, ca 250 m a.s.l., 28.03.2008, leg. P. 

Erzberger, herb. Erzberger, (B) 12896. 

H. H. Blom 

(Figs 13, 14) 

Plants medium-sized or small, forming dense or lax tufts of variable colour (olivaceous, 

brownish, black). Central strand distinct. Hair-point 0–1.1 mm, coarse and stiff, 

mostly somewhat flattened in lower part, but sometimes terete throughout, straight or 

more rarely irregularly curved in upper part, from shortly to longly and broadly decurrent or 

sometimes not decurrent, finely to coarsely spinulose-denticulate. Abaxial side of costa 

with scattered low papillae in upper part, more rarely smooth; leaf margins distantly and 

finely denticulate to papillose-denticulate in upper part, more rarely smooth. Lamina smooth, 

unistratose in lower part to irregularly bistratose in upper part. Lamina cells in upper part 

rounded, not or slightly sinuose, 8–9 μm wide, in lower part wider and sinuose, basal cells often 

noticeably wider than cells above, basal marginal cells with thickened crosswalls. K+ red. 

Sporophytes common, mostly deeply immersed. Urn orange-brown (yellowish when 

empty and becoming finely striolate), length/width ratio (1.3–)1.6–1.8–2.2 (BLOM 1996), in 

Hungarian material (1.2–)1.78 ±0.03(–2.6), n = 190. Exothecial cells predominantly elongate. 

Stomata 0–4(–6) per urn, sometimes ± rudimentary. Peristome teeth 300–450 μm, red, 

smooth or finely papillose-striated in lower part, papillae often in oblique rows. 

Very variable. The most common morph with denticulate leaf apices (Fig. 13C, D) and 

shortly elongate exothecial cells (Fig. 13I) is easily identified, but more deviant forms can 

be hard to name. They may closely resemble S. elegantulum or S. helveticum. According to 

our observations, the number of stomata can be greater than 6, the number given by BLOM 

(1996). This makes the differentiation between S. crassipilum and S. elegantulum subsp. 

wilsonii (8–16 stomata per urn, BLOM 1996) more problematic. S. elegantulum differs from 

S. crassipilum in the absence of a central strand and the structure of the hair-point, which is 

terete throughout in the former (Fig. 17), and usually distinctly flattened and somewhat 

broadened towards insertion in the latter species (Fig. 13B). See also the note under S. 

elegantulum. S. helveticum has a shorter hair-point of different structure (Fig. 21B, C), no 

stomata and a characteristically different exothecial cell pattern. (Fig. 21G, H). 

:InHungary(asinScandinavia)S. crassipilum is by far the most common 

Schistidium species on calcareous rocks (limestone, dolomite) or concrete, but it is also 

found more rarely on siliceous rock types (andesite, trachyte, breccia, sandstone, granite, 

basalt, muskovite) and basic or even acidic types of soil, in a great variety of conditions 

(dry or moist, exposed to the sun or shaded, even in caves, in northern or southern exposition, 

in forests or in open habitats, e.g. calcareous grasslands). A rather weedy species. 



A B E F D 

C 

G 

I 

K 

H 

Schistidium crassipilum. = leaves; = leaf apices (B, C in lateral view, 

costa); E = perichaetial leaf; = perichaetial leaf apex; = capsule; , , = exothecial 

cells at mid urn / at lower third of urn / at base of urn; = part of peristome. Scale bar: A, 

E, G – 2 mm; F – 800 μm; B, C, D, I, K – 400 μm; H, J – 200 μm. [A–C, E–H, J–K: Erzberger 

12090; D, I: EGR Vajda 1941 (Pilishegy), del. Erzberger] 

J 



: Amblystegium serpens, Anomodon attenuatus, Barbula unguiculata, 

Bryoerythrophyllum recurvirostrum, Bryum argenteum, B. cf. kunzei, B. moravicum, 

Ceratodon purpureus, Cirriphyllum tommasinii, Didymodon acutus, D. fallax, D. rigidulus, 

Ditrichum flexicaule, Eurhynchium hians, E. pulchellum, Grimmia laevigata, G. pulvinata, 

Hedwigia ciliata var. leucophaea, Homalothecium lutescens, Homomallium incurvatum, 

Leskea polycarpa, Orthotrichum anomalum, O. cupulatum, O. sp., Oxystegus tenuirostris, 

Pottia lanceolata, Pseudoleskeella catenulata, P. nervosa, Pylaisia polyantha, Schistidium 

apocarpum, S. brunnescens subsp. brunnescens, S. dupretii, S. elegantulum, S. helveticum, S. 

lancifolium, S. robustum, Seligeria pusilla, Thuidium abietinum, Tortella tortuosa, Tortula 

crinita, T. muralis, T. ruralis. 

: 83–910 m a.s.l. 

Distribution of Schistidium crassipilum 

Selected specimens examined: Aggtelek Karst: 7589/1 Comit. Abaúj-Torna. In rupibus 

calcareis apricis “Tohonya-bérc” prope Jósvafõ, 250 m, 01.05.1953, leg. Á. Boros, BP 

110956; : Garadna-Tal, angesprengte schattige, feuchte Kalkwand an 

der Straße ca 2 km östlich Ómassa, N 48° 06’ 43.8”, E 20° 32’ 19.6”, ca 460 m, 07.04.2007, 

leg. T. Pócs and P. Erzberger, herb. Erzberger, (B) 12119; : Comit. 

Heves. In declivibus dumetosis inter vallem rivi Csevice-patak et jugum montis Ágasvár 

supra pag. Tar, 300–600 m, 06.05.1928, leg. Á. Boros, BP 110920; : 

Com. Pest. In rupibus siccis in decl. montis Várhegy prope pag. Csõvár, montes Cserhát, 

17.03.1951, leg. L. Vajda, BP 5641; : Pest county. Babat valley, 

Gödöllõ city, basalt rock, oak forest-grassland transition, N 47.623°, E 19.384°, 200 m, 

11.03.2001, leg. P. Ódor, herb. Ódor, 20010311-10; : Comit. Hont. 

Hung. cent. In valle “Ernõ forrásvölgye” ad pag. Zebegény. Solo trachyt., 16.07.1928, leg. 

J. Szepesfalvi, BP 5709; : In saxis trachyticis rivuli Kõhegy ad 

Pomáz cottus Pest, 24.04.1904, leg. L. Simonkai, BP 5657; : Comit. Nógrád. 



In monte Nagyszál supra Kosd, 10.09.1916, leg. Á. Degen, BP 46951; : 

Hung. centr., com. Pest. In rupibus calcareis montis Pilishegy supra pag. Pilisszentkereszt, 

01.06.1941, leg. L. Vajda, EGR; Comit. Pest. In rupibus calcareis sept. montis 

Oszoly prope Csobánka, 250 m, 31.03.1946, leg. Á. Boros, BP 110884; : 

Comit. Pest. In saxis calcar. pedis montis Zsíros-hegy prope pag. Budajenõ, 350 m, 

21.03.1948, leg. Á. Boros, BP 110887; : Szomód, Betlehem-vadászház, 

tölgyes-EF, árnyas mészkõ, N 47° 42.058’, E 18° 19.496’, 321 m, 26.08.2006, leg. P. 

Szûcs, herb. Szûcs, NYG 7; : Comit. Komárom. In rupibus calc. occid. 

dumetosis montis Lófõ supra pag. Várgesztes, 350 m, 12.05.1935, leg. Á. Boros, BP 110835; 

: Comit. Fejér. In rupibus graniticis montis Zsidó-hegy prope 

Pázmánd, 150–200 m, 27.06.1938, leg. Á. Boros, BP 110832; : Comit. 

Veszprém. In rupibus calc. vallis Cuhavölgy prope Csesznek, 350 m, 07.06.1928, leg. Á. 

Boros, BP 5679; : Hung. occid. Ad muros vinearum mtis “Badacsonyhegy” 

ad lac. Balaton. Solo basalt., 300 m, 11.08.1933, leg. J. Szepesfalvi, BP 39074; 

: Comit. Zala. In rupibus dolomit. sept. montis Rezi-vár prope Rezi, 

300–400 m, 23.03.1962, leg. Á. Boros, BP 110798; : Sopron, Bot. kert, 

muszkovit-szilikát sziklán, 04.02.2004, leg. P. Szûcs, herb. Szûcs, SOP/1; : 

Comit. Vas. In muris in arboreto ad Szeleste, 180 m, 10.06.1962, leg. Á. Boros, BP 

111371; : Comit. Zala. In rupibus calcar. eocen. ad fontem majorem ripae rivi 

Marcal prope Gyepükaján, 170 m, 15.09.1961, leg. Á. Boros, BP 110807; : 

Comitat Somogy Ad muros pr. pag. Inke, 180 m, 23.02. 1956, leg. Á. Károlyi, BP 

59108; : Mánfa. Téglakerítésen, 23.08.1931, leg. A. Visnya, BP 

111391; : Comit. Baranya. In rupestribus ad Sárga-bánya in monte 

Csukma prope pag. Máriagyüd, 200 m, 08.08.1999, leg. B. Papp, BP 166476; : 

Comit. Moson. In muris viae ferreae ad fluv. lajta prope pag. Hegyeshalom, 120 m, 

10.04.1920, leg. Á. Boros, BP 111360; : Comit. 

Csongrád. Sándorfalva. Tegmen lepidarium aggeri ad Sajtos, 83 m, 09.08. 1951, leg. L. 

Timár, BP 111355; : Com. Hajdú. In cippo in sepultura Felsõ-temetõ 

prope Hajdúnánás, 02.04.1941, leg. J. Igmándy, BP 111365; : 

Comit. Komárom. In saxis calcareis deportatis marginis alvei Danubii inter pagos 

Piszke et Lábatlan, 100 m, 15.02.1925, leg. Á. Boros, BP 110860. 

(Thér.) W. A. Weber 

(Figs 15, 16) 

Plants small, forming low tufts. Central strand distinct. Hair-point mostly very short, 

but rarely absent on all leaves, 0–0.15(–0.25) mm, weak and narrow throughout, erect, 

straight, not or very shortly and narrowly decurrent, from nearly smooth to distinctly 

spinulose. Costa and leaf margins smooth (BLOM 1996), but in some Hungarian specimens 

distinctly papillose in apical part (Fig. 15B, C). Lamina smooth, usually unistratose. 

Lamina cells (7.5–)8–9.5(–11) μm wide and not or slightly sinuose in upper part of leaf, 

wider and more sinuose below (Fig. 15E). Perichaetial leaves narrow (Fig. 15D), hardly 

concealing the urn. K+ red. 



A 

B 

F 

H 

G 

K 

C 

D 

I 

E 

J 

Schistidium dupretii. =leaves; , =leafapices; = perichaetial leaf; E =lamina 

cells in lower part of leaf; = capsules in perichaetium; =capsule; , , = exothecial cells 

at urn mouth / mid urn / base of urn; = peristome tooth. Scale bar: A, D, F, G – 2 mm; B, I, 

K – 400 μm; C, E, H, J – 200 μm. [Erzberger 8056, del. Erzberger] 



Sporophytes common, much exposed in lateral view (Fig. 15F). Urn red-brown, 

finely striolate after dehiscence, length / width ratio (1.2–)1.5–1.9–2.3(–2.6) (BLOM 

1996), in Hungarian material 1.85 ±0.3, n = 2. Exothecial cells irregular in size and shape, 

striolae composed of collapsed narrowly elongate cells. Stomata about 6 per urn. Peristome 

teeth (250–)290–380 μm, red, entire or rarely with median cracks in central part, finely 

papillose. 

Characteristic features of this species are the narrow hair-point (if present), which 

appears to be “glued on” to the leaf apex, being much narrower than the chlorophyllose 

part of the apex (Fig. 15A–C, F), and the capsule being well visible between the thin, 

appressed perichaetial leaves. 

: In Hungary, S. dupretii grows on ± exposed, dry limestone rocks or walls, 

but it has also been found on siliceous rock (trachyte); a wide substrate tolerance is also reported 

by BLOM (1996). 

: Schistidium apocarpum, S. crassipilum. 

: 250–375 m a.s.l. 

Distribution of Schistidium dupretii 

: : Ómassa, Vörös-kõ, top of limestone 

wall near stream, 375 m, 07.04.2007, leg. P. Erzberger and T. Pócs, herb. Erzberger, 

(B) 12117; : Comit. Pest Hung. centr. Ad pedem montis Kõhegy versus 

pag. Pomáz. Solo trachytico, 11.04.1928, leg. J. Szepesfalvi, BP 5651; : 

Buda: Sas-hegy, on exposed dolomite, N 47° 28’ 59.5”, E 19° 01’ 09.3”, ca 250 m 

a.s.l., 05.04.2002, leg. P. Erzberger, herb. Erzberger, (B) 8056. 



H. H. Blom 

(Figs 17, 18) 

Plants medium-sized to large, forming mats or decumbent tufts. Central strand absent. 

Hair-point 0.3–1.0 mm, erect, ± pellucid, hyaline but often brownish at insertion 

(Fig. 17A, C), straight and stiff, terete, narrow throughout, not decurrent (the transition 

between the hyaline and the chlorophyllose part of the apex smooth), distantly 

spinulose-denticulate throughout. Costa and leaf margins smooth. Lamina smooth, 

unistratose with bistratose patches in upper part. Lamina cells not or slightly sinuose, predominantly 

isodiametric, in upper part 8–10 μm wide, in lower wider. K+ red. 

Sporophytes common, immersed, but partly visible among the perichaetial leaves. 

Urn straw-yellow to orange, becoming finely striolated when old, oval, widest at middle 

(narrowed towards mouth), length/width ratio 1.6–2.0–2.4 (BLOM 1996), in Hungarian 

material (1.4–)1.87 ±0.12(–2.5), n = 25. Exothecial cells predominantly elongate, 50–70 μm 

long. Stomata 6–8 per urn (8–16 in subsp. wilsonii). Peristome teeth 350–430 μm long, 

orange to red, entire or with narrow submarginal slits in central part, densely and coarsely 

papillose, in lower part smooth or with papillae more distantly arranged. 

S. elegantulum subsp. wilsonii (which has not yet been found in Hungary) differs 

from the typical subspecies in a broader and coarser hair-point that appears rather whitish 

(pellucid in subsp. elegantulum) and smooth (i.e. the spinulae are denser and smaller than 

in subsp. elegantulum and not visible with a hand lens). Other differences include growth 

form (dense tufts versus mats or decumbent tufts in subsp. elegantulum), hair-point length 

in perichaetial leaves (up to 1.0–1.5 mm versus up to 0.7–0.9 mm), costa width at leaf base 

(75–88 μm versus 53–78 μm), costa stratosity at leaf base (5–7 vs 4–5) and number of 

stomata (8–16 vs 6–8). 

In Hungarian plants of S. elegantulum, the hair-point appears smooth when viewed 

in low magnification (as in subsp. wilsonii), due to few / short spinulae as seen in the compound 

microscope. Otherwise, they possess the characteristics of subsp. elegantulum. 

The differentiation between S. elegantulum and S. crassipilum can be very difficult in 

Hungarian plants, although in other geographical regions (e.g. in Germany), S. elegantulum 

poses no problems. Some plants could be considered transitional between the two 

species, e.g. displaying sporophytes typical of S. elegantulum (oval urns with many stomata); 

but leaves characteristic of S. crassipilum. The converse situation is observed as well. Such 

plants have been placed in the latter, variable species during the present revision. 

: S. elegantulum typically grows on ± shaded limestone or dolomite boulders 

and rocks, sometimes on concrete; in Hungary it has rarely also been found on base-rich siliceous 

rock (basalt, andesite). In Scandinavia, S. elegantulum subsp. wilsonii is reported to 

grow on exposed to semi-shaded calcareous rocks, often on wall-tops. 



A B C D E 

F 

H 

G 

Schistidium elegantulum. =leaves; =leafapex;C = subperichaetial leaf; =perichaetial 

leaf; = perichaetial leaf apex; =capsule; = exothecial cells at base of urn; 

= peristome tooth. In A and C the coloured part of the hair-point is stippled. Scale bar: 

A, C, D, F – 2 mm; B, E – 800 μm; H – 400 μm; G – 200 μm. [Erzberger 12125, del. Erzberger] 



: Bryum capillare, Didymodon rigidulus, Leucodon sciuroides, 

Orthotrichum anomalum, Plagiochila porelloides, Pseudoleskeella nervosa, Schistidium 

apocarpum, S. crassipilum, S. helveticum, Tortella tortuosa, Tortula muralis, T. ruralis. 

: 82–910 m a.s.l. 

Distribution of Schistidium elegantulum 

: : Comit. Abaúj-Torna. In 

rupibus andesit. tophac. ad ruinam Regéci-várrom prope Regéc, 600 m, 03.11.1959, leg. Á. 

Boros, BP 111933 sub Grimmia commutata Hüben., rev. P.E. : 

Comit. Abaúj-Torna. In rupibus calcar. sept. silvat. montis Szádvár prope Szögliget, 450 m, 

01.05.1953, leg. Á. Boros, BP 106976; : Szentlélek: Örvény-kõ, niedriger 

Kalkblock in lichtem Quercetum, N48° 07’ 48.4”, E 20° 32’ 22.9”, 766 m, 07.04.2007, leg. P. 

Erzberger, herb. Erzberger, (B) 12125; Répáshuta, Három-kõ, auf Kalkstein in 

lichtem Wald, N 48° 03’ 41.0”, E 20° 28’ 54.1”, 820 m, 08.04.2007, leg. P. Erzberger, herb. 

Erzberger, (B) 12145; : Bergrutsch, auf Kalkstein, N 47° 50’ 24.7”, E 19° 07’ 

49.6”, 350 m, 09.04.2007, leg. P. Szûcs and P. Erzberger, herb. Erzberger, (B) 12179; 

: Ex herb. C. Schilberszky, Farkasvölgy, mészkõ 02.05.1886, unknown collector, 

BP 5624; : Comit. Komárom. In rupibus calcareis montis 

Kõpite-hegy prope pag. Dunaalmás, 150–250 m, 27.04.1942, leg. Á. Boros, BP 110856; 

: Comit. Fejér. In rupibus dolomit. sept. montis Vár-hegy prope 

Csókakõ, 350–400 m, 20.08.1948, leg. Á. Boros, BP 111711; : Comit. 

Veszprém. Bakony. In valle Cuha-völgy, 28.05.1928, leg. Á. Degen, BP 5677; 

: Comit. Zala. In calcareis montis Péter-hegy supra Balatonarács, 200–300 m, 

02.04.1926, leg. Á. Boros, BP 110802; : In monte Szent György-hegy 

prope Tapolcza, 05.05.1912, leg. Á. Degen, BP 5673; : Sopron, Bot. 

kert, 61/7 parcella, betonon, 04.02.2004, leg. P. Szûcs, herb. Szûcs, SOP/2; : 

Comit. Baranya. In rupibus calcar. merid. montis Tubes prope Pécs, 600 m, 



26.04.1962, leg. Á. Boros, BP 111397; : Comit. Baranya. In rupestribus 

in decl. merid. montis Szársomlyó prope pag. Nagyharsány, 200–300 m, 07.02.2000, leg. B. 

Papp, BP 166467; : [between Szeged and Hódmezõvásárhely] 

Algyõ felett, sajtos szék, pyrog. andesit kõ, 22.06.1928, leg. I. Gyõrffy, BP 86585; 

: Comit. Csongrád. Szeged. Sövényháza. Stratura “P…(?)-töltés” ad 1. 

gátõrház, 82 m, 21.04.1951, leg. L. Timár, BP 111353. 

(De Not.) Ochyra 

(Figs 19, 20) 

Plants small to medium-sized, forming dense, sometimes hoary cushions. Central 

strand narrow and indistinct. Hair-point 0.15–0.85 mm, straight or slightly bent, weak and 

flattened, wide or narrow at insertion, not or shortly decurrent, finely and distantly 

denticulate. Costa and leaf margins smooth. Lamina smooth, unistratose or occasionally 

with bistratose spots in upper part. Lamina cells in upper part transversely ovate to shortly 

oblong, not or slightly sinuose, in lower part slightly sinuose, 7–9(–10) μm wide throughout 

except for wider basal cells. Basal marginal cells square to shortly rectangular, with 

± strongly thickened cross-walls, forming a rectangular alar group. Perichaetial leaves much 

larger than vegetative leaves (compare Fig. 19A and C), concave and plicate, with hairpoints 

up to 1.0 mm, wide in lower part and embracing a large portion of the upper lamina, 

but not or very shortly decurrent. K+ yellow. 

Sporophytes common, deeply immersed and often completely hidden in lateral view. 

Seta short, 0.05–0.13(–0.28) mm. Urn yellowish brown to orange brown, with a conspicuous 

bright red rim at the mouth, length/width ratio 0.9–1.3 (BLOM 1996), in Hungarian 

material 1.17, n = 1. Exothecial cells rectangular in lower half (Fig. 19H, I), irregular in 

shape in upper half (Fig. 19G). Stomata small and obscure, 3–6(–8)per urn. Peristome 

teeth rudimentary, 17–25 μm, consisting of short basal segments not or only shortly extending 

beyond the capsule mouth (but sometimes up to 320 μm long: BLOM 1996: p. 177, 

not seen in Hungarian specimens). Operculum mamillate, not rostrate. 

Usually an easily recognised species, with a unique combination of short and wide 

urn (Fig. 19E), rudimentary peristome and mamillate operculum. Superficially resembling 

Grimmia anodon in the lack of a well-developed peristome, but that species grows on calcareous 

rocks, not on siliceous rock as S. flaccidum. 

: In Hungary, S. flaccidum grows on exposed andesite and other volcanic 

rock; this tallies with its preference for siliceous substrata in other parts of its distribution 

area (Europe, North America; BLOM 1996). 

: Bryum moravicum, Grimmia laevigata, G. muehlenbeckii. 

: 320–355 m a.s.l. 



A B C F 

E 

D 

G 

H 

I 

Schistidium flaccidum. =leaf; =leafapex; = perichaetial leaf; = perichaetial 

leaf apex; = capsule; = Exothecial cells at mouth of urn /shortly above mid urn / at 

base of urn (H + I). Scale bar: A, C, E – 2 mm; B, D – 800 μm; G, H – 400 μm; F, I – 200 μm. 

[Erzberger 6343, del. Erzberger] 



S. flaccidum was known in Hungary from two locations, one near 

Pomáz in the Visegrád Mts and one between Nagymaros and Zebegény, on the left side of 

the Danube. Phytogeographically this site, Mt Szentmihály or Mt Ördög, belongs to the 

Visegrád Mts (BOROS 1968). Here the plant was apparently found only once (SZEPESFALVI 

1941), and only a single, very small specimen exists in BP, whereas at the Pomáz site, in the 

year 2000, the population was still present and vigorous. From this site, numerous collections 

exist, the oldest from 1921 (leg. Gyõrffy). S. flaccidum grows on the treeless, sunny 

slopes of the hill Kis Csikóvár, in fissures of andesite rock outcrops. Recently, B. Papp discovered 

a third locality at Szarvaskõ in the Bükk Mts. 

Distribution of Schistidium flaccidum 

: : , Szarvaskõ, vártól ÉK-re, sziklagyep, 

volcanic rocks, N 47° 59’ 35.3”, E 20° 19’ 48.3”, 355 m, 04.06.2008, leg. B. Papp, BP 

175454; : Comit. Hont, Hungariae centr. In fissuris rupium andesit. 

inter pag. Zebegény et Nagymaros, 07.1927, leg. J. Szepesfalvi, BP 5586; Comit. 

Pest. In rupium fissuris montis Kis Csikóvár ad Pomáz, sol. andesit., 320 m, 01.04.1923, 

leg. Á. Degen, BP 163958 (numerous dupl.); Pomáz, Kis Csikóvár, 09.08.2000, leg. P. 

Erzberger, herb. Erzberger, (B) 6343. 

(Schkuhr) Deguchi 

(Figs 21, 22) 

Plants medium-sized to large, ± glossy, in upper part jet-black or olivaceous, forming 

tufts. Central strand distinct. Hair-point short, but rarely absent on all leaves, 0–0.2 mm, 

coarse and ± terete, coarsely spinulose with short and broad, patent to squarrose spinulae. 



A B C D E 

H 

G 

F 

I J K L 

Schistidium helveticum. =leaves; , =leafapices; = perichaetial leaf; = perichaetial 

leaf apex; = capsule; , = exothecial cells in mid urn; , , = peristome teeth; 

= part of peristome. Scale bar: A, D, F – 2 mm; B, E – 800 μm; H, J–L – 400 μm; C, G, I – 

200 μm. [A–F, K, L: Erzberger 8015; G–J: Erzberger 12188, del. Erzberger] 



Costa and margins usually smooth, but sometimes with few papillae immediately below 

apex (Blom, pers. comm. in CASPARI 2004). Lamina smooth, irregularly bistratose or more 

rarely unistratose with bistratose patches in upper part. Lamina cells rounded, esinuose, 

(6–)7–8(–9) μm wide in upper part of leaf, 7–10 μm wide and slightly to strongly sinuose 

in central and lower parts of leaf. K+ red. 

Sporophytes common, deeply immersed, but partly exposed in lateral view. Urn yellowish 

to orange-brown, length / width ratio (1.4–)1.6–1.9–2.3(–2.5) (BLOM 1996), in Hungarian 

material (1.4–)1.6 ±0.14(–1.9), n = 7. Exothecial cells variable in size and shape, in 

central and lower part predominantly elongate, forming an irregular pattern (Fig. 21G, H). 

Stomata absent. Peristome teeth 300–430(–460) μm, orange to reddish, ending in a conical 

or flattened horizontal bar, often split in prongs at top, ± semi-perforated in lower part, 

strongly perforated to cribrose in upper part (Fig. 21I–L). 

Habitat: On often dry, more rarely humid calcareous rocks (limestone, dolomite) in 

usually open vegetation, often exposed to the sun. This habitat preference of Hungarian 

plants is the same as that reported in BLOM (1996). 

: Bryum argenteum, Didymodon acutus, D. luridus, D. rigidulus, 

Grimmia tergestina, Schistidium brunnescens subsp. brunnescens, S. crassipilum, S. elegantulum. 

: 200–600 m a.s.l. 

Distribution of Schistidium helveticum 

: : Comit. Borsod-Abaúj- 

Zemplén. In rupibus calcareis supra vallem Jósva-völgy contra vallem Almás-völgy, 220 m, 

17.08.1988, leg. M. Rajczy, BP 165391; : Eger-Felnémet, Berva-völgy, 

auf exponiertem Kalkfels, 350 m, 03.04.2002, leg. T. Pócs and P. Erzberger, herb. Erzberger, 

(B) 8015, dupl. in EGR; : Vác, Dolomit-Felsrasen Látó-hegy, N 47° 50’ 

11.6”, E 19° 07’ 59.2”, ca 500 m, 09.04.2007, leg. P. Erzberger and P. Szûcs, herb. Erzberger, 

(B) 12188; : Comit. Esztergom. In rupibus calcareis or. montis Öreg-szirt 



prope Kesztölc, 450 m, 14.04.1946, leg. Á. Boros, BP 111736; : Com. 

Pest. In rupibus calcareis montis Telkihegy prope pag. Telki, 22.03.1948, leg. L. Vajda, BP 

58480; : Comit. Esztergom. In rupibus calcareis montis Öregkõ 

prope Bajót, 300–375 m, 05.04.1936, leg. Á. Boros, BP 110846; : Csákberény, 

Ugró-völgy, auf w-exponierten, besonnten niedrigen Dolomitfelsen, 29.04.2001, 

leg. P. Erzberger, herb. Erzberger, (B) 7081; : Comit. Baranya. In 

rupibus calcar. merid. montis Tubes prope Pécs, 600 m, 26.04.1962, leg. Á. Boros, BP 

111398; : Comit. Baranya. In rupestribus calcar. montis Tenkes-hegy 

prope Máriagyüd, 300–400 m, 17.04.1942, leg. Á. Boros, BP 111384. 

(Kindb.) H. H. Blom 

(Figs 23, 24) 

Plants small to medium-sized, forming lax tufts. Central strand absent or indistinct. 

Hair-point short or absent, 0–0.3 mm, decurrent, sharply and strongly spinulose-denticulate. 

Costa distantly papillose in upper (1/2–)2/3–4/5 of the leaf length with papillae increasing 

in height towards the leaf apex (about 8 μm wide and 9–11 μm high in apical part). 

Margins irregular dentate in upper part with teeth pointing in various directions. Lamina 

unistratose, smooth or rarely with few papillae on the abaxial side near apex. Lamina cells 

sinuose, 7–10 μm wide in upper part, (8–)9–10 μm wide in lower parts of leaf. K+ red. 

Sporophytes usually present, mostly shallowly immersed and much exposed in lateral 

view. Urn dark red to reddish-brown, small, 0.7–0.85–1 mm long, length / width ratio 

1.3–1.5–1.7 (BLOM 1996), in Hungarian material (0.9–)1.3 ±0.08(–1.6), n = 33. Exothecial 

cells thin-walled, short transversely rectangular and isodiametric, square, arranged in 

regular perpendicular rows (Figs 1F, G, 23G–I). Stomata large, 6–10 per urn. Peristome 

teeth 320–400 μm, orange, semi-perforated in lower part, from almost entire to strongly 

perforated in upper part. 

Usually an easily known plant with a unique combination of apically papillose costa 

and margins, with the size of papillae increasing towards the apex (Fig. 23B, C) and small 

capsules exhibiting a characteristic pattern of exothecial cells. In moist shaded habitats, 

costal papillae may be confined to the apical part of the leaf. S. apocarpum differs from S. 

lancifolium by its larger size in nearly all aspects (leaf length, width, urn length, width, peristome 

teeth length – compare Figs 4 and 23), but there is a small overlap in these quantitative 

characters. 

In Hungary, S. lancifolium is a typical plant of shaded andesite rocks, with a preference 

for higher altitudes. In Scandinavia, this species is an early pioneer on dry acidic to 

slightly basic (typically greenstone, but very occasionally also limestone) cliffs and boulders 

in relatively dense and moist forests (WEIBULL 2006). 

: On shaded siliceous (andesite, porphyric rock, slate, sandstone) or more 

rarely calcareous rock (dolomite), often near streams or flushes and in ravines, in forests, 

preferably, but not exclusively at higher altitude (700–900 m). 



A 

C 

B 

C 

D 

C 

A 

E 

F 

G 

I 

J 

H 

Schistidium lancifolium. = leaves; = upper parts of leaves; = leaf apices; 

= perichaetial leaf; = upper part of perichaetial leaf; = capsules; , , = exothecial 

cellsinmidurn/atbaseofurn(H+I); = peristome tooth. Scale bar: A, D, F – 2 mm; B, 

E – 800 μm; C, I, J – 400 μm; G, H – 200 μm. [A–F, J: Erzberger 6136; G, I: EGR, Vajda 

1958 (Börzsöny, Rakottyás-bérc); H: EGR, Boros 1957 (Börzsöny, Bacsina-patak), del. 

Erzberger] 



: Apometzgeria pubescens, Brachythecium populeum, B. 

velutinum, Bryum capillare, Ceratodon purpureus, Ctenidium molluscum, Didymodon 

insulanus, D. rigidulus, D. vinealis, Grimmia hartmanii, Hedwigia ciliata, Homalia besseri, 

Isothecium alopecuroides, Lophocolea minor, Mnium stellare, Plagiochila porelloides, 

Rhynchostegium murale, Schistidium apocarpum, S. crassipilum, Seligeria pusilla, Thuidium 

abietinum, Tortella tortuosa, Tortula ruralis. 

: 120–900 m a.s.l. 

Distribution of Schistidium lancifolium 

: : Hungaria bor.-or., com. 

Zemplén. In rupibus andesiticis montis Szappanyos supra vall. Radványpatak völgye 

prope pag. Sárospatak, 11.09.1946, leg. L. Vajda, EGR; : Comit. Borsod. 

In rupibus porphyrit. sept. silvat. montis “Nagy István erõse” prope Nagyvisnyó, 900 m, 

30.09.1950, leg. Á. Boros, BP 110942; : Heves county, Kékes North- 

Forest Reserve, Mátraháza village, andesite rock, montane beech forest, N 47.874°, E 

020.007°, 800–900 m, 10.05.1996, leg. P. Ódor, herb. Ódor, 19960510-2, dupl. in BP; 

: Comit. Nógrád. In rupibus umbrosis montis Rakottyás-bérc supra 

vallem rivi Bacsina-patak prope Királyháza, montes Börzsöny, 20.05.1958, EGR, dupl. BP 

59580; : Szerkövek. Az andezit sziklacsoport északi oldalának 

aljában, 650 m, 11.1984, leg. G. Kis, EGR; : Budapest. In silvis sept. 

montis Csúcs-hegy ad Óbuda, 300–400 m, 15.03.1941, leg. Á. Boros, BP 106900; 

: Comit. Fejér. In rupibus dolom. dumetosis vallis Balog-völgy supra Csákvár, 

250 m, 10.09.1953, leg. Á. Boros, BP 110831; Comit. Fejér. In rupibus dolomiticis 

Papirtás prope Csákberény, 300–400 m, 29.03.1936, leg. Á. Boros, BP 110833; 

Comit. Bereg. Ad saxa andesit. pedis montis Nagy-hegy pr. Tarpa, 120 m, 

15.03.1961, leg. Á. Boros, BP 111358. 



(Figs 25, 26) 

Culm. 

Plants small to large, in upper part often reddish, forming lax or dense tufts or mats. 

Central strand absent or rarely narrow and indistinct. Hair-point 0–1.25 mm, thin, 

flexuose, from shortly to longly and widely decurrent, slightly to strongly spinulose-denticulate 

in lower part, distantly denticulate to smooth in upper part. Costa ± densely papillose 

on abaxial side (papillae 6–8 μm wide and 5–11 μm tall). Margins ± papillose-denticulate 

in upper part of leaf. Lamina unistratose or with bistratose spots in upper part, ± densely 

papillose on abaxial side (papillae 6–7(–9) μm wide and 5–8 μm tall), on the adaxial side 

scattered papillae particularly near the margins. Lamina cells incrassate, sinuose, 8–10 μm 

wide in upper, 8–11 wide in central and lower parts of leaf. K+ red. 

Sporophytes common, from deeply immersed to emergent. Urn dark reddish, widest 

at about middle, length/width ratio 1.5–1.9–2.3(–2.8) (BLOM 1996), in Hungarian material 

1.81, n = 1. Exothecial cells transversely rectangular and square. Stomata 10–17 per urn. 

Peristome teeth (310–)330–500 μm, reddish, gradually narrowed into a long, fine point, 

entire or with few to several median or submarginal perforations. 

When growing in sufficiently sun-lit places, S. papillosum usually (but not always) develops 

red-coloured spots in the lamina of upper leaves, which are diagnostic. In other respects, 

e.g. papillosity, hair-point length, this species is highly variable. 

: In Scandinavia, S. papillosum grows on both acidic and basic rock (from 

granite to limestone, but becoming more calcifuge towards the south) and occurs in dense 

forests as well as at exposed sites in open pastures (BLOM 1996, WEIBULL 2006). It prefers 

acid rocks in Catalonia (CASAS et al. 2001). 

In Hungary, this species was only recently discovered during fieldwork of the first 

author, and is presently known from a single location in the Bükk Mts, in the valley 

Lök-völgy near Felsõtárkány, where it grows in open vegetation on aleurolitic slate near 

the road, associated with S. apocarpum and S. pruinosum. 

: 445 m a.s.l. 

Specimen examined: Bükk Mts: 7988/4 Felsõtárkány, Lök-völgy, aleurolite slate 

rocks at road, N 48° 01’ 51.1”, E 20° 28’ 46.5”, 445 m, 06.04.2007, leg. P. Erzberger and T. 

Pócs, herb. Erzberger, (B) 12097. 



B 

B 

H 

A 

C 

D 

F 

I 

G 

E K J 

Schistidium papillosum. = leaves; = leaf apices; C = perichaetial leaves; , 

= lamina cells at mid leaf / at widest part of leaf; = papillae and lamina cells at abaxial 

side of leaf; =capsule; , , = exothecial cells at mouth of urn / at mid urn / at base of urn; 

= peristome tooth. Scale bar: A, C, G – 2 mm; B, I, K – 400 μm; D, E, F, H, J – 200 μm. 

[Erzberger 12097, del. Erzberger] 



Distribution of Schistidium papillosum 

(Mitt.) H. Perss. 

(Figs 27, 29) 

Plants small to medium-sized, forming dense or lax tufts. Central strand distinct, but 

narrow. Hair-point 0–0.3 mm, frequently yellow or reddish in lower part or throughout, 

thin and narrow, not decurrent. Costa smooth, margins entire or irregularly denticulate just 

below apex. Lamina smooth, in upper and central parts partly bistratose or more rarely unistratose. 

Lamina cells slightly incrassate, collenchymatous, in upper part of leaf ± isodiametric 

and shortly elongate, 9–11 μm wide, in central and lower parts more elongate. 

K+ red. 

Sporophytes occasional to frequent, immersed. Urn yellowish to red-brown, subspherical 

to obovoid (cyathiform when old), length/width ratio 0.8–1.3 (BLOM 1998), not 

measured in Hungarian material. Exothecial cells thin-walled or slightly incrassate, in central 

and lower part of urn predominantly shortly elongate, forming a ± regular pattern. 

Stomata absent. Peristome teeth 380–570 μm, red, tapering to a narrow point, semiperfoated 

in lower part, strongly irregularly perforated in upper part. Spores finely 

granulose, 16–24 μm. 

This is in Hungary the only species of the Rivulare group, which is characterised 

by spores (16–24 μm) larger than those of the Schistidium apocarpum complex 

((8–)9–10.4–12(–19) μm). 



A B C 

D 

F 

E 

G 

I 

H 

Schistidium platyphyllum. = leaves; = perichaetial leaf; = leaf apex; = capsule; 

, = exothecial cells at mouth of urn / at mid urn; , = seta surface cells in upper / 

lower part of seta; = part of peristome. Scale bar: A, B, D – 2 mm; I – 400 μm; C, E–H – 

200 μm. [A–C, E–H: BP 163340; D, I: Erzberger 12625 (Sweden, Abisko), del. Erzberger] 



A 

B 

C 

G 

D 

F 

E 

Schistidium rivulare (Erzberger 6529, Germany: Hessen). = leaves; = leaf 

apex; =capsule; , = exothecial cells at mid urn / at base of urn; = seta surface cells 

in upper part of seta; = peristome tooth. Scale bar: A, C – 2 mm; G – 400 μm; B, D, F – 

200 μm. [Del. Erzberger] 



From S. rivulare (which has not yet been found in Hungary – Fig. 28), it can be distinguished 

by gametophytic characters, e.g. the entire leaf margins (except at apex – compare 

Figs 27C and 28B) and the more shallowly keeled, symmetric leaves (sharply keeled 

and often asymmetrically secund in S. rivulare: Fig. 28A), and sporophytic characters, particularly 

the patterns of exothecial cells and seta surface cells (exothecial cells isodiametric 

and incrassate in S. rivulare: Fig. 28D, E), shortly elongate and rather thin-walled in 

S. platyphyllum (Fig. 27F); seta surface cells ± strongly elongate, incrassate in S. rivulare 

(Fig. 28F), shortly elongate to isodiametric and rather thin-walled in S. platyphyllum: 

Fig. 27G), and by the absence of stomata. 

Habitat: In Scandinavia S. platyphyllum grows on base-rich or calcareous rocks in or 

beside streams (rarely submerged), occasionally in fissures of irrigated ledges and cliffs in 

the mountains, or on regularly inundated sea- or lake-shore boulders (BLOM 1998, 

WEIBULL 2006). In NW Germany it was found mainly on siliceous blocks at the rivers Elbe, 

Weser, Ems, Rhine and some of their tributaries (MEINUNGER and SCHRÖDER 2007). The 

Hungarian habitat on block edging at the banks of the Danube corresponds to the German 

type of lowland habitat on river banks. 

: 100 m a.s.l. 

: In Hungary at present known from a single collection. 

Distribution of Schistidium platyphyllum 

: Comit. Pest. In saxis irrigatis 

Danubii ad urb. Szentendre, 100 m, 11.08.1994, leg. B. Papp and M. Rajczy (sub S. rivulare), 

rev. P. Erzberger 2007, conf. W. Schröder 2007, BP 163340. 

The plants conserved in this specimen were growing on stones in the inundation area 

at the Danube in the small island “Pap-sziget“ near Szentendre (B. Papp, ex verbis). A recent 

search of the site and of similar places near Szentendre and in the Danube valley 

between Szob and Vác by the first author was unsuccessful. 



This record represents a considerable range extension within the European area of S. 

platyphyllum, viz. towards the river Danube in the southeastern part of Central Europe. 

The species was hitherto known from Asia, Africa, America, Greenland, Svalbard (SMITH 

2004), and in Europe from Scandinavia (BLOM 1998), Great Britain (SMITH 2004), and 

Germany (HOMM 1998, SIEMSEN et al. 2003, SCHULZ and DENGLER 2006, MEINUNGER 

and SCHRÖDER 2007). In the meantime, S. platyphyllum has been discovered in South Germany, 

at a tributary of the Danube, too (“Innschleife auf Uferpackungen bei Ebing, 7740/4, 

15.05.1998”, leg. W. S. sub S. rivulare, rev. 01.03.2008; in MEINUNGER and SCHRÖDER 2007 

erroneously under S. rivulare). Its discovery in the Austrian part of the Danube valley is to 

be expected sooner or later. 

(Wilson ex Schimp.) G. Roth 

(Figs 30, 31) 

Plants medium-sized, forming dense, often hoary tufts. Central strand narrow but 

distinct. Hair-point 0.2–1.25 mm, ± coarse, not or shortly and broadly decurrent, densely 

spinulose in lower, more distantly spinulose in upper part. Abaxial side of costa densely 

papillose (papillae 3–6 μm wide and tall), margins ± smooth except at dentate apex. 

Lamina partly bistratose in upper part, densely papillose with low, broad papillae (3–4.5 μm 

wide and tall) on both adaxial and abaxial sides of leaf. Lamina cells rounded, not or 

slightly sinuose in upper part, mostly sinuose in central and lower parts, 6–9 μm wide 

throughout except for wider basal cells. Perichaetial leaves large and conspicuous, broadly 

elliptical or broadly ovate, 0.8–1.25 mm wide, strongly contrasting the lower leaves 

(0.6–0.85(–1.0) mm wide). K+ red. 

Sporophytes common, deeply immersed. Urn dark red to red-brown with a motherof-pearl 

shine, widest at middle (narrowed towards mouth), length / width ratio 

(1.2–)1.3–1.5–1.8(–1.9) (BLOM 1996), in Hungarian material (1.5–)1.6 ±0.05(–1.7), n = 6. 

Exothecial cells predominantly square. Stomata 6–8 per urn. Peristome teeth 

270–390(–450) μm, reddish, in lower part entire or semi-perforated, in upper part mostly 

strongly perforated with narrow slits. 

S. pruinosum differs from other species with papillose lamina in the elliptical perichaetial 

leaves contrasting the narrower lower leaves (compare Fig. 30A and D), the small 

rounded and oval upper leaf cells (Fig. 30E), the irregularly bistratose lamina, the distinct 

central strand and the papillae usually abundant on both sides of the leaf, arranged in 

oblique rows (Fig. 30G, H). 

: In Hungary on rather exposed or more rarely shaded siliceous rocks (andesite, 

aleurolitic slate), in Scandinavia on ± exposed, often warm, base-rich or siliceous (especially 

basalt and porphyric) rocks (BLOM 1998). 

: Hypnum cupressiforme, Schistidium apocarpum, S. papillosum. 

: 200–730 m a.s.l. 



A B J 

D 

C 

I 

M 

G 

H 

E 

F 

L 

Schistidium pruinosum. = leaf; , = leaf apices; = perichaetial leaves; , 

= lamina cells at mid leaf / at lower third of leaf; , = papillae and lamina cells at 

abaxial / adaxial side of leaf; = capsule; , , = exothecial cells at mouth of urn / at mid 

urn / at base of urn; M = peristome tooth. Scale bar: A, D, I – 2 mm; B – 800 μm; C, K, M – 

400 μm; E–H, J, L – 200 μm. [A–H, J–M: BP 171483; I: Erzberger 12095, del. Erzberger] 

K 



Distribution of Schistidium pruinosum 

: : Nagy szárkõ/Regéc, 

665–695 m, 04.04.2004, leg. B. Papp, BP 171483; 7695/3 Hungaria bor.-or., com. Zemplén. 

In rupibus umbrosis in sylvestribus montis Darnó supra pag. Hercegkút, 04.09.1948, leg. L. 

Vajda, EGR; : Felsõtárkány, Lök-völgy, aleurolite slate rocks at road, N 

48° 01’ 51.1”, E 20° 28’ 46.5”, 445 m, 06.04.2007, leg. P. Erzberger and T. Pócs, herb. 

Erzberger, (B) 12095; Lehmschieferfelsen an der Straßenböschung zwischen 

Bükkzsérc und Felsõtárkány, mit Hypnum cupressiforme, N 47° 59’ 43.4”, E 20° 28’ 59.5”, ca 

400 m, 27.03.2008, leg. P. Erzberger and T. Pócs, herb. Erzberger, (B) 12883; : 

Comit. Heves. In rupibus andesit. septen. montis Disznókõ prope Parád, 730 m, 

25.03.1951, leg. Á. Boros, BP 110915; : Comit. Hont, Hung. centr. 

In saxis vallis “Malomvölgy” sub mont. “Nagycsitár” inter pag. Zebegény et Kóspallag, 200 m, 

07.07.1927, leg. J. Szepesfalvi, BP 5723. 

(Nees et Hornsch.) H. H. Blom 

(Figs 32, 33) 

Plants medium-sized to large, forming small or large, typically conspicuously hoary 

tufts. Central strand distinct, mostly broad. Hair-point ± coarse, mostly long and conspicuous, 

(0.2–)0.4–0.8(–1.1) mm, stiff and straight, decurrent, densely and evenly spinulose 

with sharp, strongly protruding spinulae. Costa and margins smooth. Lamina smooth, 

unistratose or less frequently with bistratose spots. Lamina cells mostly elongated and becoming 

shorter towards apex, (8–)9–10(–11) μm wide in upper, 8–11 μm wide in central and 

lower parts of leaf, sinuose to strongly sinuose with incrassate walls. K+ red. 



A 

B 

D 

G 

C 

H 

E 

J 

F 

I 

Schistidium robustum. = leaf; = leaf apex; = perichaetial leaf; = perichaetial 

leaf apex; = lamina cells at widest part of leaf; = capsule; , , = exothecial 

cells at mouth of urn / at lower third / at base of urn; = part of peristome. Scale bar: A, C, F 

– 2 mm; B, D – 800 μm; H, J – 400 μm; E, G, I – 200 μm. [Erzberger 8024, del. Erzberger] 



Sporophytes common, shallowly immersed, much exposed in lateral view. Urn light 

brown, sometimes faintly striolate, widest at about middle, length/width ratio (1.9–)2.1–2.4–3.1 

(BLOM 1996), in Hungarian material 2.15, n = 1. Exothecial cells irregular in size and shape, 

predominantly elongate. Stomata 8–10(–12) per urn. Peristome teeth 300–440 μm, orangered, 

narrow and longly tapering to a very thin and fragile point, frequently perforated. 

In typical plants, the strongly sinuose, incrassate leaf cells, somewhat resembling 

those in Racomitrium, are very characteristic (Fig. 32E). Plants from moist, shaded habitats 

can have less sinuose leaf cells. S. robustum can superficially approach S. elegantulum, 

but differs in the decurrent hair-point (Fig. 32B), usually not pronounced leaf bistratosity, 

a distinct central strand, and in leaf areolation as well as urn shape, long and narrow in S. 

robustum (Fig. 32F). 

: In Hungary as in Scandinavia S. robustum grows on ± dry calcareous rocks 

(limestone, dolomite) in exposed or moderately shaded sites. 

: Schistidium crassipilum. 

: 200–770 m a.s.l. 

Distribution of Schistidium robustum 

Selected specimens examined: Bükk Mts: 8088/1 Felsõtárkány, Mész-völgy, auf 

grobem Kalkschotter, 03.04.2002, leg. P. Erzberger and T. Pócs, herb. Erzberger, (B) 8024; 

: Comit. Pest: in m. Pilis, 19.09.1909, leg. G. Lengyel, BP 110898; 

: Comit. Fejér. In saxis dolom. silvat. versus Várgesztes prope Vérteskozma, 

400 m, 24.04.1932, leg. Á. Boros, BP 110826; : Mons Tobán, ad rupes, 

10.06.1932, collector unknown, BP 26270; : County Veszprém. 

On exposed calcareous rock Szamár-kõ in valley Kígyós-völgy at Balatongyörök, 200 m, 

18.10. 1999, leg. B. Papp BP 166860. 



ARTIFICIAL KEY FOR SCHISTIDIUM IN HUNGARY 

Apart from the 15 taxa found in Hungary, several others have been reported 

from parts of Central Europe (BLOM 1996, 1998, BLOM and LÜTH 

2002, GRIMS 1999), and might therefore occur in Hungary as well, though 

hitherto undetected. Since the highest elevation of present-day Hungary is 

1,015 m a.s.l. (RADÓ 1979), the occurrence of high-mountain and alpine 

taxa is not to be expected. Taxa that have not yet been shown to occur in 

Hungary, but might be expected, should, in our opinion, be included in a 

key for naming taxa of Schistidium collected in Hungary. In addition to 

those taxa that we have found in this revision, our key therefore includes 

the following: S. elegantulum subsp. wilsonii, S. pulchrum, S. rivulare, S. 

spinosum, S. trichodon var. nutans, S. trichodon var. trichodon. 

Taxa that have been shown to occur in Hungary are printed in 

type; taxa not printed in bold might occur in Hungary, but have not yet 

been found. 

Leaf lamina with papillae 

Lamina smooth, but papillae sometimes at leaf margin and dorsal side 

of costa 

Upper lamina in part bistratose, papillae on both (dorsal and ventral) 

sides of lamina; leaves never with red spots; up to 8 stomata per urn; 

central strand distinct 

Lamina unistratose or rarely with bistratose patches in upper part; 

papillae numerous on dorsal side, sparse or missing on ventral side (if 

present, then near margin, costa or apex); hair-point fine, often rather 

long; lamina occasionally with red spots (absent in plants from shaded 

sites); usually more than 8 stomata per urn; central strand absent or indistinct 

(< 6 cells) 

(siliceous rocks, also on old walls and concrete) 



Hair-point coarse, especially well developed in perichaetial leaves; 

costa dorsally hardly projecting; mid-leaf cells 6–9 μm wide and 

rounded, dorsal and ventral side of lamina with numerous papillae arranged 

in rows (examine young leaves!), usually with extended 

bistratose patches; perichaetial leaves very wide and differing in form 

from stem leaves; central strand of 3–10 cells 

(base-rich or siliceous rocks, esp. basalt and porphyric) 

Hair-point fine (but mostly straight), costa dorsally prominent; 

mid-leaf cells 10–12 μm wide and ± sinuose; papillae on both sides of 

lamina, but less numerous on adaxial side; bistratose patches not conspicuous; 

perichaetial leaves only slightly wider than stem leaves; central 

strand of 20–30 cells 

(exposed calcareous rocks, also on old walls and concrete) 

Urn length / width ratio 0.8–1.3, shape not oblong or oblong-cylindrical 

(but S. brunnescens – 17a – with oblong exothecial cells, small spores, 

K+ red and stomata absent sometimes has urns with length / width ratio 

< 1.3) 

Urn length / width ratio 1.3, shape oblong to oblong-cylindrical 

Spores 15–24 μm; hair-point absent on most leaves or very short 

(0–0.3 mm); K+ red; stomata absent or present; leaves 0.5–1.3 mm 

broad (Rivulare group) 

Spores 8–13 μm; hair-point usually present; K+ yellow; stomata present; 

leaves 0.3–0.85 mm broad (Confertum group) 

Exothecial cells incrassate, isodiametric, stomata present; hair-point 

absent; capsule-bearing branches in fascicles of (2–)3–6; leaves 

2.1–3.2 × 0.7–1.3 mm, carinate, slightly asymmetric, apex sometimes 

boat-shaped, leaf margin in upper half of leaf often dentate by projecting 

cells 

S. rivulare (siliceous or calcareous rocks in and beside rivers and 

streams) 



Exothecial cells thin-walled, predominantly oblong; stomata absent; hairpoint 

short or absent, 0–0.3 mm, smooth or finely spinulose- denticulate; 

capsule-bearing branches not in fascicles; leaves 1.4–2.3 × 0.5–1.0 mm, 

obtusely keeled, symmetric, margins entire or irregularly denticulate 

immediately below apex 

subsp. 

(base-rich rocks in or beside 

rivers or streams, rarely submerged) 

Peristome absent or rudimentary (< 25 μm), stomata 3–6 per urn; 

hair-point usually long, finely denticulate; perichaetial leaves plicate; 

operculum mamillate; plants small to medium-sized 

(siliceous rocks) 

Peristome perfect, stomata 3–10 per urn; hair-point short or long, 

coarsely spinulose; operculum rostrate, but rostrum sometimes short; 

plants small 

Hair-point short, 0–0.45 mm, fine, flattened and mostly broadened towards 

insertion; peristome teeth strongly and irregularly perforated, 

pattern of exothecial cells not very irregular, stomata small, 3–8 per 

urn 

(exposed siliceous or base-rich rocks) 

Hair-point usually long, 0–0.8 mm, coarse, terete and narrow throughout, 

not or shortly and narrowly decurrent, in young leaves strongly 

spinulose with some very long spines (22–30(–75) μm); peristome 

teeth almost entire or narrowly perforated in 1(2) rows, exothecial 

cells forming an irregular pattern; stomata 6–8(10) per urn 

S. spinosum (exposed siliceous rocks) 

Leaf margin or dorsal side of costa in upper half of leaf with papillae or 

projecting cells (denticulate) 

Leaf margin and dorsal side of costa smooth (few papillae may occur at 

leaf apex immediately below the hair-point, in S. dupretii also further 

down) 



Exothecial cells in central part of urn predominantly oblong, up to 

60 μm; peristome teeth 300–450 μm long, in freshly deoperculate capsules 

stellate-straight, scarcely twisted, orange; hair-point coarse, of 

variable length; urn usually with 0–4(6) stomata at base, but sometimes 

more, pores rudimentary or lacking; central strand distinct (> 6 cells); 

spores 8–11 μm 

(calcareous rocks, often on concrete) 

Exothecial cells predominantly isodiametric and transversely elongate; 

peristome teeth 300–710 μm long, strongly curved, bent clockwise 

and twisted; stomata with distinct pores present; hair-point fine, 

of variable length; central strand absent or narrow (< 6 cells); spores 

(9)11–15(19) μm 

Peristome teeth forming a dome above the urn mouth, (410)450–700 μm 

long, very finely and longly pointed, red; columella not falling with the 

lid; plants often black, hair-point absent or short (0–0.18 mm) 

S. trichodon (moist calcareous rocks) 

) urn never cyathiform, length / width ratio 1.6–2.4, plants brownish, never 

jet-black 

S. trichodon var. nutans 

) urn often cyathiform, length / width ratio 1.3–2.1, plants often jet-black 

S. trichodon var. trichodon 

Peristome teeth reflexed or spreading, 320–710 μm long, their points 

may be bent upwards, but never form a dome, red-orange; columella 

falling with the lid 

Urn 0.7–1.0 mm long and 0.5–0.65 mm wide, exothecial cells quadrate 

to transversely rectangular, in regular perpendicular rows, peristome 

teeth 320–400 μm; costa strongly papillose on abaxial side in upper 

half of leaf, papillae increasing in length towards leaf apex (up to 9–11 

μm high), leaf margin strongly and irregularly denticulate to serrate in 

upper part of leaf; perichaetial leaves 0.5–0.8 mm wide, small plants 

(shaded siliceous, rarely calcareous rocks) 

Urn 0.9–1.35 mm long and 0.5–0.85 mm wide, exothecial cells isodiametric, 

but pattern less regular; peristome teeth 400–700 μm; leaf margin 

finely denticulate or rarely smooth, costal papillae lower; perichaetial 

leaves 0.8–1.5 mm wide, medium-sized plants 

(calcareous or siliceous substrata) 



Exothecial cells in central and lower part of urn predominantly isodiametric 

and transversely elongate 

Exothecial cells predominantly oblong 

Peristome teeth strongly curved, often semi-perforated in lower part 

and with small perforations in longitudinal rows in upper part, leaves 

mostly falcate to falcato-secund, mostly large plants 

(longipilose plants; calcareous or siliceous substrata) 

Peristome teeth straight or slightly curved, entire or irregularly perforated 

in upper part; leaves usually straight; small to medium-sized 

plants 

Perichaetial leaves broad, 0.8–1.1(1.25) mm wide (urn not or slightly 

exposed in lateral view); hair point structure intermediate between 

fine and coarse: rather coarse and stiff, but remnants of cell walls mostly 

clearly visible, broad (as broad as chlorophyllose part of the apex), up 

to 0.7 mm long, intensely white, urn smooth, plants glossy 

S. pulchrum (on siliceous or calcareous rocks in humid sites) 

Perichaetial leaves narrow, 0.5–0.6 mm wide (urn much exposed in 

lateral view); hair point fine but straight when dry, consisting of few 

cells, distinctly narrower than the chloropyllose part of the apex, up to 

0.15 (0.25) mm long; urn finely striolate after dehiscence, plants dull, 

small, growing in compact cushions 

(exposed calcareous, less often siliceous rocks) 

Basal marginal cells with thick transverse walls, more or less hyaline; 

urn hemispherical to ovate, peristome teeth short (< 340 μm) and 

cribrose (i.e. with numerous pits and holes); hair-point mostly fine and 

long (except S. brunnescens), flattened at base and composed of cells 

with clearly visible lumen 

Basal marginal cells usually without thick transverse walls (examine 

several leaves), sometimes elongate; urn ovate to oblong, peristome 

teeth 220–430 μm; hair-point always coarse, variable in length, terete 

or indistinctly flattened at base 



Stomata present; plants on siliceous substrate; hair-point flat and fine, 

short; exothecial cells in mid-urn 20–42 μm long 

(exposed siliceous or base-rich rocks) 

Stomata absent; plants of calcareous rocks; hair-point coarse, well developed 

in perichaetial leaves, often absent or short in normal leaves; 

exothecial cells up to 50–80 μm long, forming a regular pattern 

(dry, exposed calcareous rocks) 

) Leaves ovate-lanceolate to ovate-triangular, often with shoulders, 1.85–2.6 mm, 

usually with longitudinal ridge-like striae on one or both sides of costa; costa 

58–90 μm wide in central part of leaf subsp. 

) Leaves oblong, ovate or short ovate-triangular, 1.1–2.0 mm; shoulders and longitudinal 

ridge-like striae absent; costa 38–60 μm in central part of leaf 

subsp. 

Urn with 6–10 stomata; leaves acuminate; hair-point usually conspicuous, 

0.3–1 mm 

Urn with 0–4 stomata; leaves not acuminate; hair-point conspicuous 

or less than 0.2 mm 

Central strand absent, urn usually ovate in shape, with 6–8(–16) stomata; 

exothecial cells with hardly incrassate longitudinal walls; hairpoint 

of normal leaves terete at base, not decurrent; upper lamina cells 

isodiametric, not sinuose, not distinctly incrassate; lamina with irregular 

bistratose patches 

(shaded calcareous rocks) 

) Hair-point broad, very finely but densely spinulose with only slightly protruding 

spinulae (appearing smooth with a hand lens); peristome teeth straight, 

patent to squarrose; costa in lower part of leaf 75–88 μm wide, 5–7 stratose; 

plants forming tufts; stomata 8–16 per urn S. elegantulum subsp. wilsonii 

) Hair-point narrow, distantly spinulose with longer spinulae (visible with a 

hand lens); peristome teeth curved, erect to erecto-patent with tips curved inward; 

costa in lower part 53–78 μm wide, 4–5 stratose; plants forming mats or 

decumbent tufts; stomata 6–8 per urn 

subsp. 

Central strand always distinct (> 20 cells); urn usually cylindrical in 

shape, with 8–10 stomata; exothecial cells with irregularly thickened 

longitudinal walls; hair-point usually flattened at base, decurrent; upper 

lamina cells sinuose, incrassate, bright yellowish, especially in lower 

lamina distinctly incrassate and sinuose; lamina rarely with bistratose 

patches 

(calcareous rocks) 



Plants green, olive or brownish, rarely black, in loose tufts or decumbent; 

hair-point often > 0.2 mm, slightly flattened at base and often 

decurrent; exothecial cells not forming an irregular pattern, stomata 

usually few (0–6) 

(calcareous rocks, often on concrete) 

Plants usually black, glossy, in compact cushions; hair-point absent or 

short (up to 0.2 mm), coarse and coarsely spinulose with short and 

broad, patent to squarrose spinulae, terete, not decurrent; exothecial 

cells forming an irregular pattern, stomata absent 

(= singarense) (calcareous rocks) 

FREQUENCY AND RARITY OF TAXA IN HUNGARY 

Fig. 34 shows the proportion of each taxon in a total of 448 sites of all 

Schistidium specimens examined. The most frequent taxa are: S. crassipilum 

(43.5%), S. brunnescens subsp. brunnescens (16.3%), S. apocarpum (14.1%), 

S. lancifolium (7.6%), S. elegantulum (7.6%), and S. helveticum (4.0%). The 

other taxa are known from less than 8 sites each: S. robustum (7 sites), S. pruinosum 

and S. brunnescens subsp. griseum (6 sites each), S. dupretii (4 sites), S. 

flaccidum (3 sites), S. confertum (2 sites), S. confusum, S. papillosum and S. 

platyphyllum (1 site each). 

From a glance at the geological map of Hungary (RADÓ 1979) it is evident 

that, apart from large parts of the country (in the lowlands) covered by 

quaternary deposits and thus less inviting for colonisation by saxicoles, 

there is (in the mountains and hills) a high diversity in both carboniferous 

and siliceous rocks, i.e. potential sites for taxa of Schistidium. 

Schistidium has been recorded in 30 out of 43 bryogeographical regions 

in Hungary (after Boros 1968). This clearly does not reflect the true 

distribution, since weedy species like S. crassipilum, which colonise walls 

and concrete, are to be expected in all regions. Among the regions without 

Schistidium records (Cserehát Hills, Putnok Hills, Karancs–Medves Mts, 

Fertõ Hills, Vend Region, Õrség, Hetés, Göcsej, Zselic, Külsõ-Somogy, Little 

Hungarian Plain, Nyírség, Dráva Region), there are even some siliceous 

mountains (Karancs–Medves Mts). They were not visited by Hungarian 

bryologists as frequently as the other mountain areas, since they were 

thought to be less interesting and poorer in species (BOROS 1968). 



However, for the study of Schistidium distribution in Hungary, these areas 

may still hide unexpected treasures. 

Species like S. brunnescens, S. crassipilum, S. elegantulum and S. helveticum 

with a strong substrate preference for calcareous rocks naturally 

show their maximum number of sites in the carboniferous mountains (Aggtelek 

Karst, Bükk, Pilis, Buda, Gerecse, Vértes, Bakony, Mecsek, Villány 

Mts). On the other hand, among the more frequent species, S. apocarpum 

and S. lancifolium, both have most growth sites in the Börzsöny Mts (and 

less in the Zemplén, Visegrád, Mátra and Bükk Mts), thus demonstrating a 

preference for siliceous substrates. The distribution maps of the less frequent 

species give immediate evidence of their respective substrate preferences: 

S. robustum, S. brunnescens subsp. griseum, S. confusum and S. dupretii 

predominantly grow on calcareous rocks (Bükk, Pilis, Buda, Vértes, 

Bakony Mts), whereas S. pruinosum, S. papillosum, S. confertum and S. flaccidum 

are found exclusively on siliceous rocks (Zemplén, Bükk, Mátra, 

Börzsöny, Visegrád Mts). 

Frequency of Schistidium taxa in Hungary 



However, these frequency data are not without bias. This becomes obvious 

when considering the number of sites in different regions (Table 2): a 

large number of collection sites of Schistidium taxa can in fact indicate a 

high diversity of growth sites in the region, but it could also just be the consequence 

of more thorough bryological collecting. In this context, it is interesting 

to regard the two regions with the highest numbers of sites: the 

Bükk Mts (62 sites) and the Buda Mts (43 sites). In the case of the Bükk Mts 

species number is also highest (12 taxa), therefore this seems to be a centre 

of diversity for Schistidium in Hungary. On the other hand, the high number 

of collection sites in the surroundings of Budapest, where species 

number is only 8, might be indicative of bryological activity concentrated 

around the capital. The high diversity of the Bükk Mts is certainly due to 

the presence of different geological bedrocks (calcareous as well as siliceous), 

but perhaps other factors, e.g. the altitudinal range (up to 900 m), low density 

of human settlements and the large area protected as a national park 

are also important. 

Number of growth sites of Schistidium taxa as recorded on specimen labels in 

regions of Hungary with 10 or more sites. 

Region (as defined in Boros 1968) No. of sites No. of Schistidium taxa 

Bükk Mts 62 12 

Buda Mts 43 8 

Aggtelek Karst 34 5 

Gerecse Mts 34 5 

Vértes Mts 33 6 

Pilis Mts 33 5 

Börzsöny Mts 28 5 

Balaton Uplands 24 4 

Visegrád Mts 23 5 

Bakony Mts 18 4 

Zemplén Mts 13 3 

Mátra Mts 12 5 

Mecsek Mts 12 4 

Villány Mts 10 5 

As regards rare species, it is important to distinguish between species 

that are probably under-recorded and others that are truly rare. We consider 

S. platyphyllum and S. papillosum, both with a single growth site 



known at present, to be under-recorded. Although a first search along the 

banks of the Danube to find more S. platyphyllum was unsuccessful, it is 

hardly conceivable that there should be no more potential sites along this 

river in Hungary, considering the occurrence of this species along German 

lowland rivers. S. papillosum was discovered only recently at its only known 

growth site during a search of siliceous substrata for Schistidium. Although 

a few excursions were undertaken into such areas (in the Bükk and Börzsöny 

Mts), there remain many potentially rewarding places with siliceous 

bedrock in these mountains as well as in the Zemplén, Mátra, Karancs– 

Medves, Börzsöny and Visegrád Mts to be explored in more detail. 

On the other hand, there are reasons to believe that S. confusum, with 

only one site, and in particular S. confertum and S. flaccidum, with two and 

three sites, respectively, might be truly rare. S. confusum and S. flaccidum 

are considered to be rare in Europe, and S. confertum is rare in Fennoscandia, 

but more widespread in Central and Southern Europe (BLOM 

1996). In Hungary, S. confusum has been discovered only recently in a single 

population, in spite of plenty of potential growth sites on calcareous substrate 

throughout the country. The species circumscription of S. confertum 

and S. flaccidum has been stable within the last hundred years, and was 

known to Hungarian bryologists of former times. Their populations have 

been detected about fifty (S. confertum) and a hundred (S. flaccidum) years 

ago. No new site of either of them was found during fieldwork of the first author, 

but a single new location of S. flaccidum has been discovered recently 

(B. Papp, unpublished). 

The remaining taxa with only few records: S. pruinosum, S. dupretii, S. 

robustum, and S. brunnescens subsp. griseum are certainly not frequent, but 

may also be under-recorded, since there are two or three recently discovered 

sites for each of them. 

It is interesting to compare frequency and rarity of Hungarian taxa 

with data from other Central European countries, e.g. Austria (GRIMS 1999), 

Baden-Württemberg (HOLZ 2000) and Germany (MEINUNGER and 

SCHRÖDER 2007). As in Hungary, S. crassipilum and S. apocarpum are frequent 

and widely distributed in the above-named countries. Nearly as frequent is 

S. elegantulum in Baden-Württemberg, much less so in other parts of Germany, 

and in Austria. In Scandinavia, this species has a southwestern distribution 

(BLOM 1996), and perhaps some oceanic affinity is reflected in the 



fact that it becomes rarer towards continental parts of Europe like Hungary. 

It may be significant that most of its Hungarian records are in the 

Bükk Mts, the highest among the calcareous mountains, and receiving 

most precipitation (RADÓ 1979). On the other hand, S. brunnescens subsp. 

brunnescens, one of the most frequent taxa in Hungary, may appear to be 

more continental, since records are fewer in Germany and Austria. S. 

lancifolium, which in Hungary is among the five most frequent species, is 

definitely even more continental; it is not known from Baden-Württemberg 

and rare in Germany and Austria. The characterization of its German 

and Austrian habitats corresponds well to our observations in Hungary, 

and where elevation is given in the Austrian records, it ranges from 800 to 

1,700 m a.s.l. S. helveticum, which is not rare in the south-western and central 

parts of Germany and in Hungary, appears to be very rare in Austria, 

maybe due to scarcity of xerothermic habitats. 

For the remaining taxa, Hungarian records are few, but in the light of 

data from Germany and Austria, this may in some cases not reflect their 

true frequency. S. papillosum, S. robustum and S. dupretii are frequent in 

both countries, and we have at present no idea why they appear to be much 

rarer in Hungary. S. pruinosum is not infrequent in Baden-Württemberg 

and a few other, mainly western parts of Germany, but rare in Austria and 

perhaps also in Hungary. The taxa of the Confertum group, S. confertum 

and S. flaccidum, which are thought to be truly rare in Hungary, are rare and 

red-listed in Baden-Württemberg and Germany (in the highest threat categories, 

2 and 1, respectively, of all extant Schistidium taxa in MEINUNGER 

and SCHRÖDER 2007); in Austria, however, S. confertum is moderately frequent 

in the siliceous parts of the Alps, whereas S. flaccidum is rare to occasional 

(H. Köckinger, pers. com.). S. confusum is rare in Germany as well as 

in Austria. 

CONSERVATION 

It seems premature to draw conclusions with respect to threat status 

according to the IUCN criteria (IUCN 2001), since much of the data is still 

insufficient for this purpose. To improve the state of knowledge, a more 

comprehensive field survey of Hungary with special focus on Schistidium 

would be desirable. This would probably lead to the discovery of several 



new locations, and maybe even new taxa. It would also be necessary to 

re-visit the sites of old records and check if the species are still there. Some 

future field research should also aim at population dynamics, not only of 

rare or less frequent species, but also of species of intermediate frequency 

viz. S. elegantulum and S. helveticum, because we do not yet understand, 

why they are not much more widespread, given the many additional potential 

growth sites in the calcareous parts of Hungary. Thus S. elegantulum 

has most of its records in the Bükk Mts, but strikingly few in the other limestone 

and dolomite areas, none at all in the Pilis Mts. Does this reflect a 

preference for limestone vs. dolomite, or are there climatic reasons, or is it 

just an artefact of insufficient mapping? Similarly, it is not clear, why S. helveticum 

has not (yet?) been found in the Bakony Mts or the Balaton Uplands. 

Many of the Schistidium growth sites mapped during this revision lie 

already in protected areas like the Bükk National park, the Aggtelek National 

Park and the Duna–Ipoly National Park. 

However, since this protected status does not prevent the sometimes 

negative consequences to habitat quality of forestry management, 

it is important to make the localities of important populations known to 

the authorities concerned. In this context, we would like to draw attention 

to the sites of S. flaccidum (Bükk Mts, Visegrád Mts) and the two only sites 

of S. confertum (Mátra and Börzöny Mts). The status of S. confertum is uncertain. 

A short search at the surroundings of Holló-kõ (Börzsöny Mts) by 

the first author was unsuccessful; no search has been made at Disznó-kõ 

(Mátra Mts) for this species. 

It is hoped that this survey of Schistidium in Hungary, where this genus 

is well represented, has outlined some of its interesting taxonomic, floristical 

and ecological features, and that it will help and stimulate future research. 

* * * 

Acknowledgements – The authors are grateful to Dr H. H. Blom for examining a critical 

specimen, to the curators and directors of the public herbaria and the owners of private 

herbaria mentioned in the Introduction for the loan of specimens, to Dr L. Lõkös for the 

preparation of maps, to PD Dr H. Kürschner for the opportunity to use a Leitz drawing apparatus, 

to R. McF. Craig for correcting the English and to Mag. H. Köckinger for helpful 

comments. Special thanks are due to Prof. emer. Dr T. Pócs for his genuine interest in this 

work and his active participation in the search for interesting potential growth sites of 

Schistidium in the Bükk Mts. 



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(Received 21 November, 2007)

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