A taxonomic study of genus Brachythecium Schimp. (Brachytheciaceae, Bryophyta) in the Mediterranean region

Nova Hedwigia Vol. 95 issue 3 4, 295 318 Stuttgart, November 2012 Article A taxonomic study of genus Brachythecium Schimp. (Brachytheciaceae, Bryophyta) in the Mediterranean region J. David Orgaz, María Jesús Cano and Juan Guerra Departamento de Biología Vegetal, Facultad de Biología, Universidad de Murcia, 30100 Murcia, España. E-mail: jdavid.orgaz@um.es With 6 figures Orgaz, J.D., M.J. Cano & J. Guerra 2012: A taxonomic study of genus Brachythecium Schimp. (Brachytheciaceae, Bryophyta) in the Mediterranean region. Nova Hedwigia 95: 295 318. Abstract: The moss genus Brachythecium (Brachytheciaceae) is taxonomically revised for the Mediterranean area, based on morphological information. Fourteen species are recognised. Lectotypes for Brachythecium geheebii Milde, Hypnum glareosum Bruch ex Spruce and Hypnum mildeanum Schimp. are designated. A key to the species of the Mediterranean region as well as descriptions, illustrations and distribution data are provided. New records for Bosnia-Herzegovina, Cyprus, France and Slovenia reported. Key words: Brachytheciaceae, Brachythecium, Bryophytes, distribution, Mediterranean area, taxonomy, typifications. Introduction The genus Brachythecium Schimp. represents one of the most complex and diverse genera in terms of morphological variation in the family Brachytheciaceae. It includes approx. 150 species in the world (Crosby et al. 1999, Frey & Stech 2009), distributed mainly in temperate areas. It is mainly characterised by small to large plants, usually concave and plicate, lanceolate to ovate leaves, autoicous or dioicous sexual conditions, rough or smooth setae and a conic operculum. In recent years the taxonomical status of genus Brachythecium has changed drastically, since Ignatov & Huttunen (2002) carried out molecular studies in Brachytheciaceae using three molecular markers (chloroplastic psbt-h and trnl-f and nuclear ITS2). According to these studies, species previously placed in Brachythecium are now placed in three different genera (Brachytheciastrum Ignatov & Huttunen, Brachythecium and Sciuro-hypnum (Hampe) Hampe) within Brachytheciaceae. 2012 J. Cramer in Gebr. Borntraeger Verlagsbuchhandlung, Stuttgart, www.borntraeger-cramer.de Germany. DOI: 10.1127/0029-5035/2012/0062 0029-5035/2012/0062 $ 6.00 295

More recently, molecular studies on the genera Brachytheciastrum (Vanderpoorten et al. 2005; Vanderpoorten & Goffinet 2006) and Sciuro-hypnum (Draper & Hedenäs 2009) supported the monophyly of both genera and transferred some species previously placed in Brachythecium to Brachytheciastrum and Sciuro-hypnum. Except for the treatment of Brachythecium in diverse regional flora or compendia (Casas et al. 2006, Cortini-Pedrotti 2006, Kürschner & Frey 2011), there has been no taxonomic study focused on Brachythecium in the Mediterranean area, and many members of this genus remain poorly understood. In recent years we have been engaged in studies of the Mediterranean Brachytheciaceae, especially in the genus Brachythecium s.l. (Orgaz & Blockeel 2010, Orgaz et al. 2010, Orgaz et al. 2011); here we present the first taxonomic revision that has been carried out for the genus Brachythecium in the Mediterranean region, as circumscribed in 'Material and methods'. Material and methods This study is based on a revision of more than 1600 specimens from the herbaria B, BCB, BCN, BM, BP, BUCA, C, CAIA, CAME, E, FCO, FI, GDAC, GJO, GZU, H, HBG, HUJ, JE, L, LEB, LISU, M, MA, MACB, MAN, MGC, MHA, MO, MUB, MW, NY, O, OXF, PAMP, PC, S, SALA, SOM, UPS, VAL, VIT, W, WU and the private herbaria of T.L. Blockeel (Sheffield, U.K.), V. Mazimpaka and F. Lara (Madrid, Spain), J. Martínez Abaigar (Logroño, Spain) and R. Skrzypczak (Montbrison, France). The geographic area covered includes Albania, Algeria, Andorra, Bosnia-Herzegovina, Bulgaria, Croatia, Cyprus, Egypt, France, Greece, Israel, Italy, Jordan, Lebanon, Libya, Macedonia, Malta, Montenegro, Morocco, Portugal, Romania, Serbia, Slovenia, Spain, Syria, Tunisia and Turkey. We have considered as Mediterranean countries those bordered on Mediterranean sea, plus four countries (Bulgaria, Jordan, Macedonia and Romania) that although are not bordered on Mediterranean sea, have, al least in some areas, climate Mediterranean influence. The global distribution of the species is summarized based on the following checklists and floras: Africa (O Shea 2006), Asia (Hu et al. 2008, Ignatov et al. 2006, Iwatsuki 2004), Oceania (Fife 1995, Hedenäs 2002), North America (Anderson et al. 1990), South America (Churchill et al. 2000, Matteri 2003, Müller 2009). Microscopic examinations and measurements were made using an Olympus-BH2 light microscope and microphotographs were obtained with a Spot inside 3.5 camera mounted on this microscope. Under 'Selected specimens studied', one collection per country is listed when the number of specimens studied is very large. If the number of samples specimens studied is small, all are listed. Taxonomic treatment Brachythecium Schimp., Byol. Eur. 6: 5. 1853. Lectotype: Brachythecium rivulare Schimp., designated by Robinson (1962). Plants small to large, dark green to yellowish, occasionally brownish-yellow or brownish-green. Stems postrate to ascending, regularly or irregularly branched; hyalodermis absent, sclerodermis developed, central strand present. Stems usually erect, occasionally julaceous ascending or subjulaceous. AxillAry hairs 2 4 celled, with 1 2 basal brownish cells and 1 2 apical hyline cells. PseudoPArAPhylliA foliose, triangular to orbicular, rarely lanceolate. rhizoids irregularly branched, reddish to 296

brownish, smooth. Stem leaves erect to patent when dry, occasionally imbricate, lanceolate to broadly ovate-lanceolate, slightly plicate to strongly plicate, occasionally not plicate, concave or somewhat concave, straight or slightly falcate, occasionally strongly falcate or curved, gradually or abruptly narrowed into acute to piliferous apex, slightly to longly decurrent; margins entire to strongly serrulate, recurved or not at base, occasionally recurved throughout. CostA single, reaching 0.4 0.95 way up leaf, ending in a spine or not. laminal Cells linear, thick- or thin-walled, smooth, rarely prorate; AlAr Cells differentiated or not, quadrate to rectangular, thin or thickwalled, occasionally inflated, forming a well defined group or not, opaque or pellucid, ascending up along margins or not, not excavate. Branch leaves smaller than stem leaves, lanceolate to ovate; margins entire to serrate, occasionally strongly serrate; lamina cells occasionally prorate. Autoicous or dioicous. Perichaetial leaves with a broad basal part abruptly narrowed into a large and usually reflexed acuminate apex, with a short costa, sometimes without costa. Seta straight or slightly flexuose, rough or smooth, reddish to brownish. Capsule inclined to horizontal, occasionally erect, reddish to brownish; urn ovate, occasionally oblong, straight to curved. Annulus caduceous, separating in fragments. Peristome double; exostome of 16 lanceolate to triangularlanceolate teeth; external surface of exostome with vertical and horizontal trabeculae; basal part cross-striolate; medium part papillose to densely papillose-verrucose; upper part strongly papillose-verrucose, sometimes with disperse papillae; internal surface of exostome generally with vertical and horizontal trabeculae, slightly papillose to papillose; endostome with 16 slightly keeled segments scarcely papillose, CiliA 1 3(4), usually nodose, occasionally subapendiculate or apendiculate; basal membrane as high as segments, usually with disperse papillae. operculum conic. CAlyPtrA cucullate, naked. Spores sphaerical, slightly to strongly papillose. Key to the species of Brachythecium in the Mediterranean area 1 Costa ending more than 0.8 of leaf length...1. B. geheebii 1' Costa ending less than 0.8 of leaf length... 2 2 Alar cells weakly differentiated, forming a poorly defined group, or undifferentiated... 3 2' Alar cells clearly differentiated, forming a more or less well defined group... 5 3 Stem leaves usually broadly ovate to ovate-lanceolate... 2. B. rutabulum 3' Stem leaves triangular to lanceolate... 4 4 Stem leaves usually triangular; margins entire or almost so... 3. B. mildeanum 4' Stem leaves usually lanceolate; margins clearly denticulate, at least at apex... 4. B. capillaceum 5 Alar cells inflated... 5. B. rivulare 5' Alar cells not inflated... 6 6 Stem leaves usually abruptly narrowed into an acuminate or piliferous apex... 7 6' Stem leaves gradually narrowed into an acuminate or acute apex... 9 7 Stem leaves generally with acuminate apex, usually slightly falcate; alar cells pellucid; seta smooth... 6. B. albicans 7' Stem leaves with piliferous apex, not falcate; alar cells opaque; seta rough... 8 8 Alar cells ascending up along leaf margins; lamina cells 7 10 µm wide... 7. B. tommasinii 8' Alar cells not ascending up along leaf margins; lamina cells 6 7 µm wide... 8. B. cirrosum 297

9 Leaf margins entire or subentire... 10 9' Leaf margins denticulate to serrulate, at least at apex... 11 10 Dioicous; pseudoparaphyllia triangular; alar cells thin-walled, forming a pellucid group...... 9. B. glareosum 10' Autoicous; pseudoparaphyllia orbicular; alar cells thick-walled, forming an opaque group...... 10. B. turgidum 11 Stem and branch leaves curved or falcate... 11. B. erythrorrhizon 11' Stem and branch leaves straight or slightly falcate... 12 12 Seta rough, sometimes weakly so; branch leaves with prorate cells on dorsal face... 12. B. campestre 12' Seta smooth; branch leaves without prorate cells on dorsal face... 13 13 Alar cells thick-walled, forming an opaque group; dioicous; leaf apex acute or shortly acuminate, acumen (0.25)0.3 0.5(0.6) mm long... 13. B. laetum 13' Alar cells thin-walled, forming a pellucid group; autoicous; leaf apex longly acuminate, acumen (0.45)0.5 0.9(1) mm long... 14. B. salebrosum Description of species 1. Brachythecium geheebii Milde, Hedwigia 8: 161. 1869. Fig. 1.1 3 Homalothecium geheebii (Milde) Wigh, Bot. Not. 126: 323. 1973. Type: PolAnd. Schlesien, Porphyrfelsen bei Görbersdorf, July 1868, Milde s.n. (lectotype: S!, here designated). Stems postrate, (20)30 60 mm long, regularly branched. PseudoPArAPhylliA triangular, occasionally orbicular, ending in a small apiculus, 0.35 0.5 0.25 0.4 mm. Stem leaves triangular, occasionally lanceolate, 1.25 1.75 0.55 0.75 mm, usually strongly plicate, slightly concave, straight, gradually narrowed into an acuminate apex, acumen 250 500 µm long, leaf base slightly decurrent; margins entire or subentire, usually recurved at base. CostA reaching 0.80 0.95 way up leaf, rarely ending in a spine. laminal Cells in mid-leaf 30 65 7 9 µm, thin-walled, not prorate; AlAr Cells differentiated, quadrate to subquadrate, 12 15 10 13 µm, thick-walled, not inflated, forming a well defined group, ovate, opaque, not ascending up along leaf margins. Branch leaves usually lanceolate; laminal cells without prorate cell endings. Dioicous. Seta rough throughout. habitat And distribution: Brachythecium geheebii grows between 450 and 2100 m a.s.l. (above sea level) in rock crevices and on ground in meadows or scrublands, mainly in mountain ranges. It is known from Europe and Asia. In the study area this species has been found in Bulgaria, Bosnia-Herzegovina, France, Greece, Romania and Turkey. Records from Croatia, Macedonia, Montenegro, Serbia and Slovenia (Sabovljević et al. 2008) and Italy (Aleffi et al. 2008) remain unconfirmed by us. notes: This species is easily recognised by its triangular leaves with costa reaching 0.80 0.95 way up leaves and the alar cells forming an opaque group. Brachythecium laetum could be confused with B. geheebii but the costa in B. laetum is shorter, never reaching 0.80 way up leaf. The protologue of Homalothecium geheebii reads "In Silesia prope Görtersdorf (Milde 1868); kleine Schneegrube (Schulze); Buchberg im Isergeb. (Limpr.); Rhöngebirge, 298

Fig. 1. 1 3. Brachythecium geheebii (1 3 Blockeel 36/330). 1. Leaf apex. 2. Leaf base with alar cells. 3. Stem leaf. 4 6. Brachythecium erythrorrhizon (4 6 MUB 38469). 4. Leaf apex. 5. Leaf base with alar cells. 6. Stem leaf. Scale bar: 1 = 80 µm; 2, 5 = 0.1 mm; 3 = 0.25 mm; 4 = 60 µm; 6 = 0.3 mm. copiose, etiam c. fr. (Geheeb.). Silva nigra: ad montem Feldberg (Sickenberger)". We have found two syntypes of this name in S, where the original herbarium of Milde is kept, one of them collected by Milde in Poland and the other collected by Limpricht in Germany. We designate as lectotype the sample collected in Poland because it is better preserved. specimens studied: bulgaria: Montana, Stara Reka river valley above Zanozhene quarter, Vurshets, 29-6-2006, Ganeva s.n. (SOM 9830). bosnia-herzegovina: Republika Srpska, Turov stan auf der Treskavica planina, 5-8-1904, Glowacki s.n. (GJO). FrAnCe: Alsace, Vosges, col de la Schlucht, 1983, Philippi s.n. (L). greece: East Macedonia and Thrace, Lepida forest, Tsoukali Rema, ca 1 km north of road junction (left fork) 3 km NE of the forestry station, 20-6-2007, Blockeel 36/330 (Herb. T.L. Blockeel). romania: Cluj, Hidegszamos, Somesul-Rece, 27-9-1962, Boros s.n. (BP 179191). turkey: Artvin, Ardanuc, Kordevan dag, Yalnizcam Daglari, at Kütül Yayla, 28-6-1957, Davis & Hedge 30404h (E). Trabzon, Pontic mts., Altindere Valley Natural Park, Trabzon town, Maçka village, Sumela Monastery, on Shaded volcanic rocks, 18-7-2001, Papp 01/2 (BP). 299

2. Brachythecium rutabulum (Hedw.) Schimp., Bryol. Eur. 6: 16. 1853. Fig. 2.1 3 Hypnum rutabulum Hedw., Sp. Musc. Frond.: 276. 1801. Type: "the lower right plant on a sheet in herb. Hedwig-Schwaegrichen" [lectotype: G, designated by Hedenäs (1996)]. Stems usually postrate, 35 75 mm long, regularly branched. PseudoPArAPhylliA broadly ovate to triangular, 0.50 0.75 0.30-0.45 mm. Stem leaves broadly ovate to ovate-lanceolate, occasionally triangular, 1.7 2.7 0.8 1.4 mm, slightly plicate, concave, straight, gradually or abruptly narrowed into an acuminate apex, sometimes twisted, acumen 300 700(800) µm long, leaf base slightly to longly decurrent; margins denticulate to serrulate, especially at apex, occasionally recurved at base. CostA reaching 0.40 0.60 way up leaf, not ending in a spine. laminal Cells in midleaf 75 120 7 10 µm, thin-walled, not prorate; AlAr Cells scarcely differentiated, shortly rectangular to rectangular, 18 35 10 22 µm, thin-walled, not inflated, forming a weakly defined group, pellucid, not ascending along leaf margins. Branch leaves ovate-lanceolate, occasionally lanceolate; laminal cells without prorate cell endings Autoicous. Seta rough throughout. habitat And distribution: Brachythecium rutabulum grows as an epiphyte on trees, on the ground in slopes, in forest meadows, on rocks, river banks and generally on wet and shady places, from 0 to 2225 m a.s.l. It has been found in Africa, Asia, Europe, North America, Oceania and South America. In the study area this species is known from Albania, Bulgaria, Bosnia-Herzegovina, Croatia, France, Greece, Italy, Lebanon, Morocco, Montenegro, Portugal, Slovenia, Spain, Tunisia and Turkey. Records from Argelia (Ros et al. 1999), Malta (Frahm & Lüth 2008), Macedonia, Romania and Serbia (Sabovljević et al. 2008) and Syria (Kürschner & Frey 2011) remain unconfirmed by us. notes: Brachythecium rutabulum is a very variable species since it can grow in many different habitats. Nevertheless it can be easily recognised by the ovate-lanceolate leaves with denticulate margins, the alar cells scarcely differentiated, and the rough seta. Specimens that grow on river and spring banks often have inflated basal cells, which is the cause of their misidentification with B. rivulare. Normally, the differentiation of these species is easy because B. rivulare usually has an acute leaf apex, and B. rutabulum has a longly acuminate apex. However, some specimens are very difficult to identify. selected specimens studied: AlbAniA: Korab, 1-5-1959, Korpoh s.n. (BP 64437). bulgaria: Blagoevgrad, Bulgaria australis, prope via Katuntsi-Petrovo, 14-3-2003, Ganeva & Natcheva s.n. (SOM 8067). bosnia-herzegovina: Republika Sprska, Hladne vode, bei Cajnica, 14-5-1904, Glowacki s.n. (GJO). CroAtiA: Primorje-Gorski, Insel Sansego, 16-5-1911, Brunnthaler & Ginzberger s.n. (WU). FrAnCe: Rhône-Alpes, Isère, defilé de Maupas, 1-6-2011, Cano 6300 (MUB). greece: Central Greece, Eubea, below the Drimona waterfalls, between Drimona and Kerasia, 3-3-2000, Blockeel 29/259 (Herb. T.L.Blockeel). italy: Sicily, Palermo, Madonie, monte Daino, 3-6-1990, Düll s.n. (JE). lebanon: Der Kubil, near Choneifar, 7-3-1943, Davis 5382 (E). morocco: Marrakech-Tensift- Al Haouz, Alto Atlas, Toubkal, Subida desde Arnt hacia el refugio de Neltner, 19-6-1998, Cano et al. s.n. (MUB 8398). montenegro: Northwestern part of Skadarsko jezero, at Rijeka Crnojevica, 10-10-2003, Papp 03/90/1 (BP). PortugAl: Tras-os-Montes e Alto Douro, Bragança, Mirandela, Ponte de Barreiro, 9-10-2008, Garcia s.n. (LISU 231184). slovenia: Abhang des Bachergebirges, gegen Maria Rast bei Marburg/Maribor, 29-4-1876, Breidler s.n. (GJO). spain: Albacete, Riópar, nacimiento del río Mundo, 6-3-2008, Guerra & Orgaz s.n. (MUB 27375). 300

Fig. 2. 1 3. Brachythecium rutabulum (1 3 MUB 27375). 1, 2. Alar cells; 3. Stem leaf. 4 6. Brachythecium mildeanum (4 6 BCB 25091). 4. Leaf apex. 5. Alar cells. 6. Stem leaf. 7 9. Brachythecium capillaceum (7 9 MUB 36898). 7, 9. Stem leaves. 8. Alar cells. Scale bar: 1, 4, 7 = 80 µm; 2, 5 = 60 µm; 3, 6, 9 = 0.35 mm; 8 = 50 µm. 301

3. Brachythecium mildeanum (Schimp.) Schimp., Bot. Zeitung (Berlin) 20: 452. 1862. Fig. 2.4 6 Hypnum mildeanum Schimp., Syn. Musc. Eur.: 694. 1860. Type: PolAnd. Bei Nimkau, Bruch s.n. (lectotype: S!, here designated). Stems slightly ascending, 35 60(70) mm long, irregularly branched. PseudoPArAPhylliA orbicular, ending in a small apiculus, 0.3 0.45 0.3 0.5 mm. Stem leaves usually triangular, occasionally ovate-triangular, 1.9 2.8 0.8 1.2 mm, not plicate, occasionally slightly plicate, concave, straight, gradually narrowed into an acuminate apex, acumen 450 700 µm long, leaf base slightly to longly decurrent; margins entire, rarely with scattered teeth near apex, sometimes recurved at base. CostA reaching 0.50 0.65 way up leaf, smooth or occasionally with an indistinct spine. laminal Cells in mid-leaf 55 95 7 10 µm, thin-walled, not prorate; AlAr Cells weakly differentiated, shortly rectangular to rectangular, occasionally subquadrate, 15 35 12 20 µm, thin-walled, not inflated, forming a poorly delimited group, pellucid, not ascending along leaf margins. Branch leaves triangular, 1.5 2.1 0.8 1.1 mm; laminal cells without prorate cell endings. Autoicous. Seta smooth. habitat And distribution: Brachythecium mildeanum occurs in flooded meadows and on river banks between 10 and 2400 m a.s.l. It occurs in Africa, North America, Asia, Australia and Europe. In the study area it is known from Bulgaria, Bosnia, Croatia, France, Greece, Israel, Italy, Portugal, Slovenia, Spain and Turkey. Records from Montenegro, Romania and Serbia (Sabovljević et al. 2008) remain unconfirmed by us. notes: This species is characterised by the triangular leaves with entire margins and alar cells not well differentiated and the smooth seta. Brachythecium rutabulum is similar to B. mildeanum but it can be distinguished by the ovate or ovate-lanceolate leaves with denticulate margins and the rough seta. B. capillaceum is another species that is similar to B. mildeanum, but the leaves in the former are usually lanceolate and the margins are denticulate. The protologue of Hypnum mildeanum reads "in fossis aquosis loci turfosi 'Bruch' dicti prope Nimkau Silesiae ubi clar. Milde detexit atque pulchris speciminibus comunicavit". We have not found any syntype in BM, where original herbarium of Schimper is kept. However, we have found three syntypes for this name in Milde s herbarium in S. For two of them the locality is illegible although the date is clear. The locality of the third sample is clear but no date is provided. We have designated as lectotype this third sample, since the locality and the collector are the same as in the protologue, the sample is deposited in Milde s herbarium, and it is the only sample which has sporophytes. selected specimens studied: bulgaria: Sofia, Sredna Gora mts., Koprivstica, 0,6 km SE from the hill Goljam Bogdam, towards the Maly Bogdan mt., 2-7-2002, Hájkova s.n. (SOM 8106). bosniaherzegovina: Federation of Bosnia and Herzegovina, an der Drina, bei Gorazde, 15-8-1904, Glowacki s.n. (GJO). CroAtiA: Primorje-Gorski Kotar, Insel Veglia, von Verbenica, zum Pale von Bresca, 4-4- 1908, Baumgartner s.n. (W). FrAnCe: Poitou-Charentes, Charente-Maritime, Benon, 16-4-1953, Pierrot s.n. (O). greece: Epirus, Ioannina, Katara Pass, by roadside south west of Vitsa, 4-1990, Blockeel 19/206 (Herb. T.L.Blockeel). israel: Dan Valley, river Dan, Ein El Bard, Hefez s.n. (HUJ). italy: Lombardia, Milano, 6-9-1896, Corti s.n. (S). PortugAl: Estremadura, Porto de Mós, Serra dos Candeiros, Arrimal, Lagoa, 5-7-1982, Casas et al. s.n. (LISU 154247). slovenia: Nasse Stellen in der Verbina, bei Videm, 4-4-1885, Glowacki s.n. (GJO). spain: Zamora, Galende, Ilanes, 940 m, 7-9-1999, Heras & Infante 972/99 (VIT). turkey: Bolu, Gölu, about 30 km SW of the town Bolu, 11-11-1974, Engelmark & Nyholm s.n. (S). 302

4. Brachythecium capillaceum (F.Weber & D.Mohr) Giacom., Ist. Bot. Reale Univ. Reale Lab. Crittog. Pavia, Atti 4: 268. 1947. Fig. 2.7 9 Hypnum salebrosum var. capillaceum F.Weber & D.Mohr, Bot. Taschenbuch: 313. 1807. Brachythecium salebrosum var. capillaceum (F.Weber & D.Mohr) Lorentz, Bryol. Notizb.: 66. 1865. Type: not found. Brachythecium rotaeanum De Not., Cron. Briol. Ital. 2: 19. 1867. Type: not found. Stems postrate, 35 65 mm long, irregularly branched. PseudoPArAPhylliA triangular, 0.4 0.6 0.3 0.4 mm. Stem leaves lanceolate, occasionally triangular, 1.7 2.6 0.5 0.9 mm, usually plicate, concave, straight, gradually narrowed into an acuminate apex, acumen 400 800 µm long, leaf base slightly decurrent; margins denticulate, at least towards apex, recurved or not at base. CostA reaching 0.50 0.70 way up leaf, not ending in a spine. laminal Cells in mid-leaf 50 95 7 9 µm, thin-walled, not prorate; AlAr Cells scarcely differentiated, quadrate to shortly rectangular, 15 25 10 16 µm, thin walled, not inflated, forming a not well defined group, pellucid, not ascending up along leaf margins. Branch leaves lanceolate, 1.4 1.9 0.45 0.65 mm; laminal cells without prorate cell endings. Autoicous. Sporophyte not seen. habitat And distribution: Brachythecium capillaceum grows on the ground in slopes in forest, tree bases and fallen trunks between 2000 and 2060 m a.s.l. It is known from North America, Asia and Europe. In the study area it occurs in Italy and Romania. Records from Bosnia-Herzegovina (Sabovljević et al. 2008) and Slovenia (Martinčič 2003) remain unconfirmed by us. notes: Brachythecium capillaceum is mainly characterised by the lanceolate leaves with denticulate margins and alar groups poorly delimited, the smooth seta and the autoicous sexual condition. B. salebrosum is similar to B. capillaceum in the shape of the leaves, the denticulate margins, the smooth seta and autoicous sexual condition. However, in B. salebrosum the alar cells are clearly differentiated. Corley et al. (1981) synonymized B. capillaceum with B. roteanum and many other authors have accepted this proposal (Ignatov 1998, Hill et al. 2006, Aleffi et al. 2008). We were not able to find the type material of these taxa although, after a carefully study of samples of both species from North America and East Asia, we conclude that all of them correspond with the same species and, consequently, we accept the synonymy. specimens studied: italy: Trentino-Alto Adigio, Meran, 29-11-1883, Reyer s.n. (S). romania: Cluj- Napoca, an Waldbäumen bei Langental, 2-9-1870, Barth. s.n. (S). Prahova, Sinaia, Monte Bucegi, 6-8-2009, Guerra et al. s.n. (MUB 36898). 5. Brachythecium rivulare Schimp., Bryol. Eur. 6: 17. 1853. Type: FrAnCe. In Vogeso inferiore, 11-1845, Schimper s.n. [lectotype: BM!, designated by Hedenäs (1996)]. Fig. 3.4 6 Stems postrate to slightly ascending, 30 70 mm long, regular o irregularly branched. PseudoPArAPhylliA orbicular to broadly triangular, 0.50 0.75 0.50 0.65 mm. Stem leaves broadly ovate to narrowly ovate, occasionally triangular, 1.5 2.2 0.8 1.4 mm, slightly plicate, somewhat concave, straight, gradually or abruptly narrowed into an acute or occasionally shortly acuminate apex, acumen (100)150 250(350) µm long, leaf base longly decurrent; margins denticulate to serrulate, specially at apex, occasionally 303

Fig. 3. 1 3. Brachythecium albicans (1 3 MUB 16669). 1, 3. Stem leaves. 2. Alar cells. 4 6. Brachythecium rivulare (4 6 MUB 27374). 4, 6. Stem leaves. 5. Alar cells. Scale bar: 1, 4, 3, 6 = 0.35 mm; 2 = 50 µm; 5 = 40 µm. subentire, not recurved at base. CostA reaching 0.50 0.65 way up leaf, rarely ending in a spine. laminal Cells in mid-leaf 55 120(140) 7 11 µm, thin walled, not prorate; alar cells differentiated, shortly rectangular to rectangular, 20 50 15 30 µm, thinwalled, inflated or strongly so, forming a well defined group, pellucid, not ascending up along leaf margins. Branch leaves ovate-lanceolate to triangular, 1.1 1.75 0.5 0.9 mm; laminal cells without prorate cell endings Dioicous. Seta rough throughout. habitat And distribution: Brachythecium rivulare occurs mainly on river and stream banks, sometimes submerged in the water, as epiphytic in submerged trunks and in flooded meadows from 10 to 3000 m a.s.l. It is known from Africa, North America, Asia, Australia and Europe. In the study area it has been found in Algeria, Andorra, Bulgaria, Bosnia-Herzegovina, Cyprus, Croatia, Egypt, France, Greece, Italy, Lebanon, Morocco, Macedonia, Montenegro, Portugal, Serbia, Slovenia, Spain and Turkey. Records from Albania and Romania (Sabovljević et al. 2008) and Syria (Kürschner & Frey 2011) remain unconfirmed by us. notes: Brachythecium rivulare is easily recognised by the ovate leaves with inflated alar cells and denticulate margins and the dioicous sexual condition. 304

This species is very variable in the length and shape of the leaves, the length and shape of the leaf apex and the length of laminal cells. We have observed that the leaves of specimens which grow totally or partially submerged have a tendency to be broadly ovate and with shortly acute apex, whereas those living on river banks but not in direct contact with water have narrowly ovate leaves with longly acute apex. Specimens which live in flooded meadows usually have triangular leaves, with shortly acuminate apex and almost subentire leaf margins. selected specimens studied: AlgeriA: Kabylie de Collo, dpt. de Constantine, Vallé de l Oued, 28-5-1944, Feldmann s.n. (PC). AndorrA: subida al Stany Enrodat, 17-10-2001, Fuertes & Oliván s.n. (MACB 81132). bulgaria: Sofia, Rila mountains, Mount Malyovitsa, 16-6-1967, Een s.n. (S). bosnia-herzegovina: Federation of Bosnia and Herzegovina, Bugojno, 3-9-1904, Glowacki s.n. (GJO). CyPrus: In Opympo Cypri, 1862, Unger s.n. (W). CroAtiA: Lika-Senj, Velebit, Bacic, Plovni Dabar, 26-7-1908, Degen s.n. (W). egypt: Garbiyah, Nile Delta, Damat, 18-4-1980, Refai s.n. (CAIA). FrAnCe: Auvergne, Puy-De-Dôme, Chambon sur Lac, 10-8-1989, Greven s.n. (L). greece: Peloponnese, Fteroti river, near Kamutsa village in Taigetos mts., 2-4-1995, Papp 95/39/1 (BP). italy: Lombardia, Bellagio, 10-9-1913, Suse s.n. (H). lebanon: Mount Lebanon, 22-6-1897, Bornmüller s.n. (S). Morocco: Marrakech-Tensift-Al Haouz, Alto Atlas, Toubkal, Refugio de Nelther, 20-6-1998, Cano et al. s.n. (MUB 8415). macedonia: Sar Planina, Ljuboten, 3-7-1963, Jalas s.n. (H). montenegro: Lovcen mts., along the road form Njegusi village to the mausoleum of Njegos, 11-10-2003, Papp 03/93/1 (BP). PortugAl: Beira Baixa, Pemanacor, Serra da Malcata, 14-5-2005, Cano 2433 (MUB). serbia: Central Serbia, Jablanica, Vlasina, prope Crkvena mahala, 7-1910, Podpera s.n. (S). slovenia: Frauheimer Garten, bei Marburg/Maribor, 28-10-1900, Glowacki s.n. (GJO). spain: León, Puebla de Lillo, bajada del puerto de las Señales, río Porma, 10-7-2008, Orgaz s.n. (MUB 28399). turkey: Mugla, Okukludag, 2-4-1971, Ayedem et al. s.n. (S). 6. Brachythecium albicans (Hedw.) Schimp., Bryol. Eur. 6: 23. 1853. Fig. 3.1 3 Hypnum albicans Hedw., Sp. Musc. Frond.: 251. 1801. Type: "the left specimen on a sheet in herb. Hedwig-Schwaegrichen" [lectotype: G, designated by Hedenäs (1996)]. Stems ascending, 25 55(65) mm long, irregularly branched. PseudoPArAPhylliA triangular, 0.5 0.65 0.3 0.4 mm. Stem leaves broadly ovate-lanceolate to triangularovate, 1.6 2.5(2.8) 0.65 1.2(1.5) mm, somewhat plicate, concave, slightly falcate, occasionally straight, abruptly or rarely gradually narrowed into an acuminate or occasionally piliferous apex, acumen 0.5 0.9(1) mm long, leaf base longly decurrent; margins entire or subentire, usually recurved at base. CostA ending 0.45 0.65 way up leaf, occasionally ending in an indistinct spine. laminal Cells in mid-leaf 40 90 7 10 µm, thin-walled, not prorate; AlAr Cells well differentiated, quadrate to subquadrate, occasionally shortly rectangular, 12 30 12 22 µm, thin-walled, not inflated, forming a well defined group, triangular, pellucid, ascending up along leaf margins. branch leaves ovate-lanceolate to lanceolate, 1.2 1.9 0.5 0.8 mm; laminal cells without prorate cell endings Dioicous. Seta smooth. habitat And distribution: Brachythecium albicans grows on the on ground and slopes in forest, dunes and sandy zones and in general in humid places with soil accumulation, between 0 and 2600 m a.s.l. It is known from Africa, North America, South America, Asia, Oceania and Europe. In the study area this species has been found in Algeria, Bulgaria, Bosnia-Herzegovina, France, Greece, Italy, Montenegro, Portugal, Romania, Serbia, Slovenia, Spain and Turkey. Records from Albania, Croatia and Macedonia (Sabovljević et al. 2008) remain unconfirmed by us. 305

notes: Brachythecium albicans is easily recognised by the leaves abruptly narrowed into a long and often slightly falcate acumen, the alar cells ascending up along the leaf margins, the dioicous sexual condition and the smooth seta. Although it is easily recognised, it is a very variable species mainly as regards leaf size, the apex and lamina cell length and the falcateness of the acumen. Occasionally, Brachythecium albicans has a piliferous leaf acumen like B. cirrosum and B. tommasinii but in these two species the acumen more distinctly differentiated and not falcate, the alar groups are opaque and the seta is rough. selected specimens studied: AlgeriA: près Tizi, 6-1926, Maire s.n. (PC). bulgaria: Plovdiv, eastern Rhodopes, along a river near the village Dolno Bryagovo, 16-3-2004, Ganeva s.n. (SOM 9030). bosnia-herzegovina: Republika Srpska, Matnaca, bei Teslic, bz. Teianj, 22-7-1904, Glowacki s.n. (GJO). FrAnCe: Bretagne, Morbihan, Île d Houat, le Tal, 27-1-1989, Boudier 2021 (L). grecee: Ionian Islands, Insula Kephalonia, in montis Aenos, 13-5-1926, Bornmüller s.n. (W). italy: Abruzzos, Aquila, 1-1910, Pellegrini s.n. (FI). montenegro: Durmitor mts., Durmitor NP, Zabljak, between Momcilov grad restoran and Jablan bara, 8-10-2004, Papp s.n. (BP 178886). PortugAl: Beira Baixa, Serra da Malcata, cruzamento para a capela da Senhora do Bom Sucesso, 30-3-1999, Sim-Sim & Garcia s.n. (LISU 196257). serbia: Central Serbia, Prope Vlasina, 7-1910, Podpera s.n. (S). romania: Sibiu, montes Fagaras, lago Balea, 8-8-2009, Guerra et al. s.n. (MUB 36095). slovenia: Schieferfelsen, bei Wuchern /Vuhred, an der Drau, 8-9-1881, Breidler s.n. (GJO). spain: Lérida, Arrós, Valle de Arán, Saut deth Pish, 15-6-2010, Guerra et al. s.n. (MUB 34286). turkey: Bursa, Northern part of Uludag, 9-7-1978, Nyholm s.n. (S). 7. Brachythecium tommasinii (Sendtn. ex Boulay) Ignatov & Huttunen, Arctoa 11: 268. 2002. Fig 4.4 6 Hypnum tommasinii Sendtn. ex Boulay, Fl. Cryp. Est Muscin.: 225. 1872. Eurhynchium tommasinii (Sendtn. ex Boulay) Molendo, Ber. Naturhist. Vereins Passau 10: 224. 1875. Cirriphyllum tommasinii (Sendtn. ex Boulay) Grout, Bull. Torrey Bot. Club 25: 226. 1898. Type: FrAnCe. Ad lapides Jurasi prope Fleurier, Martius, Mougeot & Nestler s.n., Stirp. Crypt. Vogeso-rhenanae, nº1227, 1850 [lectotype: H!, designated by Karttunen (1990)]. Cirriphyllum vaucheri Loeske & M.Fleisch, Allg. Bot. Z. Syst. 13: 22. 1907. Eurhynchium vaucheri Schimp., Syn. Musc. Eur.: 556. 1860. nom. illeg. non E. vaucheri (Lesq.) Schimp. nec Hypnum vaucheri Lesq. Type: not found. Stems generally ascending, 35 65 mm long, irregularly branched. PseudoPArAPhylliA triangular, 0.5 0.75 0.4 0.5 mm. Stem leaves ovate, 1.8 2.8 0.8 1.2 mm, plicate, occasionally only slightly plicate, concave, abruptly narrowed into a piliferous apex, acumen (0.5)0.6 1100 µm, leaf base longly decurrent; margins entire or subentire, towards apex slightly denticulate, generally recurved at base. CostA ending 0.5 0.65 way up leaf, smooth or rarely ending in an inconspicuous spine. laminal Cells in midleaf 35 80 7 10 µm, thin-walled, not prorate; AlAr Cells differentiated, quadrate to shortly rectangular, 12 30 10 22 µm, thick-walled, not inflated, forming a well defined group, triangular-ovate, somewhat opaque, ascending up along leaf margins. branch leaves ovate-lanceolate to lanceolate, 1.25 2 0.4 0.8 mm; laminal cells without prorate cell endings. Dioicous. Seta rough throughout. habitat And distribution: Brachythecium tommasinii grows on the ground or on rocks, mainly calcareous ones, in forests, between 600 and 1900 m a.s.l. This species is endemic to Europe. In the study area it occurs in Andorra, Bosnia-Herzegovina, Bulgaria, Croatia, France, Greece, Italy and Montenegro. Records from Romania and Serbia (Sabovljević et al. 2008) remain unconfirmed by us. 306

Fig. 4. 1 3. Brachythecium cirrosum (1 3 VIT 37712). 1. Stem leaf. 2. Leaf base with alar cells. 3. Leaf apex. 4 6. Brachythecium tommasinii (4 6 MUB 4413). 4. Branch leaf. 5. Alar cells. 6. Leaf apex. Scale bar: 1 = 0. 35 mm; 2 = 70 µm; 3 = 200 µm; 4 = 0.3 mm; 5 = 50 µm; 6 = 100 µm. notes: Brachythecium tommasinii is characterised by the ovate stem leaves, abruptly narrowed into a pilifeous acumen, the lamina cells broader than 7 µm, the alar cells ascending up along leaf margins, the dioicous sexual condition and the rough seta. Brachythecium cirrosum is similar, but can be differentiated by its lamina cells that are narrower than 7 µm, and the alar cells do not ascend up along the leaf margins. In several works and checklist of the study area, and in many of the consulted herbaria, this species is called Cirriphyllum tenuinerve (Lindb.) Wijk & Margad. However, the basionym Hypnum tenuinerve Lindb is invalid since it was published as a synonym of Eurynchium vaucheri Schimp. (Kartunnen 1990). selected specimens studied: bosnia-herzegovina: Federation of Bosnia and Herzegovina, Krupa, quelle bei Pazarić, 30-7-1904, Glowacki s.n. (GZU). bulgaria: Gabrovo, Central Balkan mts., Bulgarka Nature park, 27-6-2008, Roussakova s.n. (SOM 9871). CroAtiA: Lika-Senj, Velebit, 16-6-1912, Baumgartner s.n. (S). FrAnCe: Aquitaine, Pyrénées, Ste. Engrâce, Gorges d Ehujarré, 27-7-1992, Greven s.n. (L). greece: Epirus, mt. Timfi, on steep ascent in forest to Stani Katsanou west of Vrisochori, 12-6-2008, Blockeel 37/261 (Herb. T. L. Blockeel). italy: Trentino-Alto Adigio, Carbonare, South of Trento, 20-7-1985, Greven s.n. (L). Veneto, Monte Baldo, Venturi s.n. (H). 307

montenegro: Peruĉica potok am Südfuß des Kom, 13-8-1896, Glowacki s.n. (GZU). slovenia: Wocberg bei Rohitsch, 25-8-1885, Glowacki s.n. (GZU). spain: Burgos, Espinosa de los Monteros, macizo de Castro Valnera, camino al Bernacho, 11-6-2008, Guerra et al. s.n. (MUB 34413). 8. Brachythecium cirrosum (Schwägr.) Schimp., Syn. Musc. Eur.: 696. 1860. Fig. 4.1 3 Hypnum cirrosum Schwägr., Reise Glockner 2: 365. 1804. Cirriphyllum cirrosum (Schwägr.) Grout, Bull. Torrey Bot. Club 25: 223. 1898. Type: not found. Stems generally postrate, (35)45 80 mm long, regular o irregularly branched. Pseudo- PArAPhylliA orbicular, ending in a short apiculus, occasionally broadly triangular, 0.4 0.75 0.4 0.7 mm. Stem leaves ovate to broadly ovate, 2.2 3.1(3.8) 0.8 1.3 mm, slightly plicate, strongly concave, straight, abruptly narrowed into a piliferous apex, acumen 0.75 1.3(1.4) mm long, leaf base shortly decurrent or occasionally longly so; margins subentire, occasionally slightly denticulate towards apex, usually recurved at base. CostA reaching 0.4 0.6 way up leaf, smooth. laminal Cells in midleaf 40 90 6 7 µm, thin-walled, not prorate; AlAr Cells differentiated, quadrate to subquadrate, rarely rectangular, 12 25(30) 10 20 µm, thick-walled, not inflated, forming a well defined group, ovate, somewhat opaque, not ascending up along leaf margins. Branch leaves ovate, 1.8 2.6 0.7 1 mm; laminal Cells without prorate cell endings. Dioicous. Seta rough throughout. habitat And distribution: This species occurs from 1200 to 2320 m a.s.l., on rocks, mainly siliceous ones, in mountainous areas. Occasionally it may appear growing on humid soils in montane zones. It is known from North America, Asia and Europe. In the study area this species has been found in Andorra, Croatia, France, Italy, Romania, Slovenia and Spain. Records from Bosnia-Herzegovina, Macedonia and Montenegro (Sabovljević et al. 2008), remain unconfirmed by us. notes: Brachythecium cirrosum is easily recognised by the ovate or broadly ovate, strongly concave leaves, abruptly narrowed into a piliferous acumen, the lamina cells narrower than 7 µm, the alar cells not ascending up along the leaf margins, the dioicous sexual condition and the rough seta. selected specimens studied: AndorrA: Serrat Melizà, damunt de la Font Roja, 14-7-1978, Casas s.n. (BCB 51071). CroAtiA: Lika-Senj, Velebit, Badanj, 22-6-1912, Baumgartner s.n. (W). FrAnCe: Franche-Comté, Dép. Doubs, ca. 15 km NW of the swiss town Vallorbe, slopes of the mont d Or, 25-7-1986, Dirkse s.n. (L). Midi-Pyrénees, Hautes Pyrénées, Aragnouet, vallée d Aure, Barroude, 21-9-2003, Infante & Heras 1491/03 (VIT 31712). Provenze-Alpes-Côte d Azur, Haute Alpes, vallée de Ségur prés Ristolas, Husnot s.n. (O). Rhône-Alpes, La Moucherolle, Alpes de l Isère, Ravaud s.n. (L). italy: Friuli-Venezia-Giulia, Küstenland, bei Görz, 12-6-1903, Baumgartner s.n. (W). romania: Prahova, Sinaia, monte Bucegi, 6-8-2009, Guerra et al. s.n. (MUB 36096). slovenia: Julische Alpen, aufstien zum Triglav, 13-9-1898, Baumgartner s.n. (W). spain: Gerona, Val de Núria, 1-9-1956, Casas s.n. (BCB 46315). 9. Brachythecium glareosum (Bruch ex Spruce) Schimp., Bryol. Eur. 6: 23. 1853. Fig. 5.4 6 Hypnum glareosum Bruch ex Spruce, Musci Pyren. nº29. 1847. Type: FrAnCe. Bédat (B.-de- Bigorre), etiam ad saxa in valle de Béost prope les Eaux Bonnes, 1847, Spruce s.n., Musci Pyren. nº29 (lectotype: BM!, here designated). 308

Fig. 5. 1 3. Brachythecium turgidum (1 3 MUB 34287). 1. Stem leaf. 2. Pseudoparaphyllia. 3. Leaf base with alar cells. 4 6. Brachythecium glareosum (4 6 MUB 28645). 4. Stem leaf. 5. Pseudoparaphyllia. 6. Alar cells. Scale bar: 1, 4 = 0.3 mm; 2, 5 = 0.2 mm; 3 = 100 µm; 6 = 50 µm. Stems postrate or slightly ascending, (35)50 85 mm long, irregularly branched. PseudoPArAPhylliA triangular, 0.45 0.85 0.3 0.55 mm. Stem leaves broadly lanceolate to ovate-lanceolate, 2.4 3.9(4.4) 0.7 1.2 mm, generally strongly plicate, slightly concave, straight, gradually narrowed into an acuminate apex, acumen 900 1300(1500) µm long, leaf base slightly to longly decurrent; margins entire or subentire, rarely denticulate at apex, usually not recurved at base. CostA reaching 0.4 0.6 way up leaf, sometimes ending in an unconspicous spine. laminal Cells in mid-leaf 50 100 µm, thin-walled, not prorate; AlAr Cells differentiated, quadrate or subquadrate, occasionally rectangular, 15 32 12 25 µm, thin-walled, not inflated, forming a defined group, shortly ovate, pellucid, not ascending up along leaf margins. Branch leaves lanceolate 1.9 2.7 0.9 1.2 mm; laminal cells without prorate cell endings. Dioicous. Seta smooth. habitat And distribution: Brachythecium glareosum grows between 100 and 2300 m a.s.l. on soils in forest or scrublands, on slopes, on rocks in forests and occasionally as epiphytic on tree bases. It is known from Africa, Asia and Europe. In the study area 309

this species has been found in Bosnia-Herzegovina, Cyprus, Croatia, France, Italy, Montenegro, Portugal, Serbia, Slovenia, Spain and Turkey. Records from Andorra (Sérgio et al. 2006), Bulgaria, Greece and Romania (Sabovljević et al. 2008) and Morocco (Ros et al. 1999) remain unconfirmed by us. This species is new to Cyprus. notes: Brachythecium glareosum is easily recognised by its ovate-lanceolate and longly acuminate leaves with entire or almost entire margins, the alar cells forming a pellucid group not ascending up along leaf margins, the smooth seta, its dioicous sexual condition and the triangular pseudoparaphyllia. Brachythecium turgidum has leaves similar to B. glareosum and the two could sometimes be difficult to separate. However, B. turgidum is autoicous, the pseudoparaphyllia are orbicular and the alar group is opaque. If the humidity conditions are favourable, e.g. in dense forests, B. glareosum could get leaves that are longer than 4 mm. There are various syntypes of this species, all of them exsiccate specimens. We have designed as lectotype, the sample kept in BM. selected specimens studied. bosnia-herzegovina: Republika Srpska, Belsbucje bei Teslic, bz Tesanj, 23-7-1904, Glowacki s.n. (GJO). CyPrus: Troodos mts, along Kryos potamos stream, at pano Platres village,18-9-2001, Papp s.n. (BP 177877b). CroAtiA: Lika-Senj, Dinarische Alpen, 6-7-1911, Baumgartner s.n. (S). FrAnCe: Midi-Pyrénees, Ariège, Salau, from Mines d Anglade to cirque de Anglade, 1400 m, 12-5-1996, Greven s.n. (L). italy: Veneto, lago di Garda, Ferrara di Monte Baldo, Madona della Corona, 26-5-1964, Cedercrentz s.n. (H). montenegro: Krivosije, Westseite der V. Subra, 7-6-1911, Baumgartner s.n. (W). serbia: Central Serbia, Branjska Banja, 7-1910, Podpera s.n. (S). slovenia: Auf Erdaufwürten im Walde bei Rohitsch/Rogaska, 20-8-1885, Glowacki s.n. (GJO). spain: Cuenca, Hoz de Beteta, 7-7-2008, Orgaz s.n. (MUB 28400). turkey: Rize, Hemsin, Meydan Kobaca, Mollaveysa, 3-9-1952, Davis 21363 (E). 10. Brachythecium turgidum (Hartm.) Kindb., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 9: 294. 1888. Fig. 5.1 3 Hypnum salebrosum var. turgidum Hartm., Handb. Skand. Fl. 2: 309. 1838. Type: sweden. Herjed fjellen, 1836, Thedenius s.n. (holotype: UPS!). Stems usually ascending, 45 90 mm long, irregularly branched. PseudoPArAPhylliA orbicular, obtuse or shortly apiculate, 0.4 0.6 0.4 0.5 mm. Stem leaves ovatelanceolate, occasionally lanceolate, 2.2 3.2 0.7 1.2 mm, strongly plicate, concave, straight, gradually or occasionally abruptly narrowed into an acuminate apex, acumen (0.6)0.7 1. mm long, leaf base slightly decurrent; margins entire or subentire, usually recurved, at leaf base, occasionally recurved towards leaf apex. CostA reaching 0.5 0.65 way up leaf, occasionally ending in an inconspicuous spine. laminal Cells in midleaf 50 90 8 10 µm, thin-walled, not prorate; AlAr Cells differentiated, quadrate to subquadrate, 10 25(30) 10 20(25) µm, thick-walled, not inflated, forming a well defined group, shortly ovate, somewhat opaque, not ascending up along leaf margins. Branch leaves lanceolate to ovate-lanceolate, 2 2.5 0.65 0.9 mm; laminal cells without prorate cell endings. Autoicous. Seta smooth. habitat And distribution: Brachythecium turgidum grows between 1900 and 2900 m a.s.l. on soils or rocks in humid mountainous areas. It is known from North America, 310

Asia and Europe. In the study area this species has been found in Bosnia-Herzegovina, France, Italy, Slovenia and Spain. This species is new to Bosnia-Herzegovina, Slovenia and France. notes: Brachythecium turgidum is not very frequent in the study area, where it is only found in high mountains. It is characterised by the ovate-lanceolate, entire leaves, with opaque alar cells, the orbicular pseudoparaphyllia, the smooth seta and the autoicous sexual condition. specimens studied. bosnia-herzegovina: Republika Srpska, Cajnica/Cajnice, 13-8-1904, Glowacki s.n. (GJO). FrAnCe: Midi-Pyrénées, Hautes Pyrénées, Aragnouet, vallé d Aure, Barroude, 21-9-2003, Infante & Heras 1483/03 (VIT). Rhône-Alpes, Saboie, massif du Mont-Cenis, col de la Masse, 27-7-1965, Een s.n. (S). italia: Lombardia, Valtellina, Vollaccia sotto Motta Grande, presso lago di Foscagno, 2-8-1924, Mazzuchelli s.n. (FI). slovenia: Krain, Hochmoor "Za mlako", auf der Poklula, bei Koprivnik, 22-7-1907, Glowacki s.n. (GJO). spain: Lérida, Boí, Muntanyó de Llacs, 6-7-1959, Monserrat s.n. (BCB 50872). Lérida, Valle de Arán, valle de Varradós, cabaña de Mont, 15-6-2010, Guerra et al. s.n. (MUB 34287). Lérida, Tor, bordes de Palanca, 8-7-1973, Casas s.n. (BCB 46469). 11. Brachythecium erythrorrhizon Schimp., Bryol. Eur. 6: 18. 1853. Fig. 1.4 6 Type: norway. Bogstadaas ved Christiania, 1827, Blytt s.n. (lectotype: BM!, designated by Orgaz et al. (2012b); isolectotype: O!). Stems usually ascending, 10 30 mm long, regularly branched. PseudoPArAPhylliA broadly triangular, 0.3 0.5 0.25 0.35 mm, occasionally slightly falcate. Stem leaves triangular-ovate to lanceolate, 1.3 1.7 0.45 0.7 mm, usually plicate, slightly concave, falcate to curved, rarely straight, gradually narrowed into an acuminate apex, acumen 300 500(600) µm long, leaf base somewhat decurrent; margins slightly denticulate to serrulate, especially at apex, recurved or not at base. CostA reaching 0.5 0.75 way up leaf, generally ending in a conspicuous spine. laminal Cells in mid-leaf 35 100 7 9 µm, thick-walled, not prorate; AlAr Cells differentiated, quadrate to subquadrate, 10 18 8 16 µm, thin-walled, occasionally thick-walled, not inflated, forming a well defined group, triangular, sometimes opaque, slightly ascending along leaf margins. Branch leaves lanceolate, 0.9 1.4 0.3 0.5 mm; laminal cells without prorate cell endings. Dioicous. Seta rough throughout. habitat And distribution: Brachythecium erythrorrhizon grows on ground, rocks or epiphytic on trees in forests, between 1300 and 2750 m a.s.l., outside the Mediterranean this species tends to grow in calcareous habitats. It is known from North America, Asia and Europe. In the study area, this species has been found in Bosnia-Herzegovina, France, Italy, Romania, Slovenia, Spain and Turkey. Records from Montenegro and Serbia (Sabovljević et al. 2008) remain unconfirmed by us. notes: Brachythecium eythrorrhizon is easily to identify, characterized mainly by its ovate-triangular to lanceolate leaves, usually curved or at least slightly falcate, with denticulate margins and a dioicous sexual condition. Occasionally, it can be confused with poorly developed specimens of B. salebrosum, but it can be distinguished from B. salebrosum by straight or occasionally only slightly falcate, ovate-lanceolate or triangular leaves and the plants are autoicous. B. albicans usually has leaves with a falcate acumen, for which reason it may be confused with B. erythrorrhizon. However, it is easy to distinguish because the leaves and leaf acumen of B. albicans are longer, its costa does not end in a spine and the leaves are not curved. 311

specimens studied: bosnia-herzegovina: Bosnia and Herzegovina Federation, Stit-planina, bei Busovaca, 1500 m, 1-8-1904, Glowacki s.n. (GJO). FrAnCe: Hautes-Alpes, Le Monêtier-les Bains, Col du Galibier, 9-9-1997, R. Skrzypczak 1034 (Herb. R. Skrzypczak). Savoie, Col de Iseran, 1-9- 2009, R. Skrzypczak 9345 (Herb. R. Skrzypczak). italy: Emilia-Romagna, Monte Prado, 15-9-1992, Velluti s.n. (CAME). romania: Distr. Buzau, Mt. Siriu, Dirohu, s.c. (BUCA 10810). slovenia: Triglav-Gebiete, 26-8-1900, Glowacki s.n. (GJO). spain: Lérida, Tavascán, Estany Romedo Baix, 16-7-2011, Guerra et al. s.n. (MUB 38468, MUB 38469). turkey: Bolu, Bolu-Gerede, Aktas Forest, 12-10-1981, Cetin s.n. (S). 12. Brachythecium campestre (Müll.Hal.) Schimp., Bryol. Eur. 6: 16. 1853. Fig. 6.1 3 Hypnum rutabulum var. campestre Müll.Hal., Syn. Musc. Frond. 2: 368. 1851. Type: not found. Stems slightly ascending, 30 60 mm long, irregularly branched. PsudoPArAPhylliA triangular, 0.5 0.65 0.35 0.45 mm. Stem leaves erect ovate-lanceolate to triangular lanceolate, 1.9 2.5(2.8) 0.7 1,1 mm, weakly plicate to strongly plicate, concave, straight or slightly falcate, gradually narrowed into an acuminate apex, acumen (0.4)0.5 0.8 mm long, leaf base slightly decurrent, margins denticulate to serrulate, specially towards apex, usually recurved, mostly at base. CostA ending 0.5 0.65 way up leaf, usually ending in a strong spine. laminal Cells in mid-leaf 45 105 7 10 µm, thick-walled, sometimes prorate; AlAr Cells differentiated, subquadrate to shortly rectangular, 10 25 8 15 µm, thick-walled, not inflated, forming a defined group, triangular, opaque, ascending up along leaf margins. Branch leaves lanceolate, sometimes triangular-lanceolate, 1.3 2.2 0.4 0.8 mm; laminal Cells strongly prorate. Autoicous. Seta rough throughout, occasionally weakly so. habitat And distribution: Brachythecium campestre grows on soils in decidous forest, outside the Mediterranean with a clear tendency to grow in calcareous habitats. Occasionally it could be found as epiphytic on tree bases; between 940 and 1960 m a.s.l. It has been found in Africa, North America, Asia, Europe and Oceania. In the study area this species is known from Greece, Italy, Romania and Slovenia. Records from Algeria (Ros et al. 1999), Bulgaria, Bosnia-Herzegovina and Croatia (Sabovljević et al. 2008), France (Augier 1966) and Turkey (Kürschner & Erdağ 2005) remain unconfirmed by us. notes: The identification of this species may be very difficult if the samples do not present sporophytes since there are not many characters to distinguish it from other similar ones such as B. laetum or B. salebrosum. However, some characters that could be useful to identify B. campestre are the ovate-lanceolate or ovate-triangular leaves, the rough seta, the autoicous sexual condition and the branch leaves with prorate laminal cells. Brachythecium laetum can be distinguished from B. campestre by its dioicous sexual condition, the smooth seta, the branch leaves do not have prorate laminal cells and the alar cells do not ascend up along leaf margins. The differences between B. salebrosum and B. campestre are not easy to observe and it is sometimes impossible to distinguish them, especially if the samples of B. campestre have no sporophytes. B. salebrosum can be distinguished by the smooth seta, the usually pellucid alar cells and the branch leaves normally without prorate lamina cells. 312

Fig. 6. 1 3. Brachythecium campestre (1 3 MUB 36099). 1. Leaf apex. 2. Alar cells. 3. Stem leaf. 4 6. Brachythecium salebrosum (4 6 MUB 28385). 4. Leaf apex. 5. Alar cells. 6. Stem leaf. 7 9. Brachythecium laetum (7 9 MUB 34285). 7. Leaf apex. 8. Alar cells. 9. Stem leaf. Scale bar: 1, 4, 7 = 80 µm; 2, 5, 8 = 50 µm; 3, 6, 9 = 0.35 mm. specimens studied: greece: East Macedonia, Rhodope mountains, Elatia forest, 3-5 km north of the forestry station, upper part of Stavronema valley, 17-6-2007, Blockeel 36/175 (Herb. T.L.Blockeel). Epus, mt. Timfi, just below the Astrakas saddle, on the west side, near the refuge, 8-6-2003, Blockeel & Sabovljević 32/234 (Herb. T.L.Blockeel). italy: Friuli-Venezia Giulia, Von Castellnuovo, bei Sagrado, 23-12-1903, Bertagnolli s.n. (GJO). Lombardia, Torre d Isola, 11-11-1898, Artaria s.n. 313