Arctoa (2002) 11: 245-296

BRACHYTHECIACEAE (BRYOPHYTA) – A FAMILY OF SIBLING GENERA BRACHYTHECIACEAE (BRYOPHYTA) – ÑÅÌÅÉÑÒÂÎ ÐÎÄΖÁËÈÇÍÅÖΠMICHAEL S. IGNATOV1 & SANNA HUTTUNEN2 ÌÈÕÀÈË Ñ. ÈÃÍÀÒÎÂ1 & ÑÀÍÍÀ ÕÓÒÒÓÍÅÍ2

Abstract

The history of familial and infrafamilial circumscriptions of the Brachytheciaceae is briefly reviewed and a new classification of the family is proposed, based largely on a phylogenetic analysis of molecular and morphological characters. The family includes four subfamilies and 41 genera; the number of species is estimated to be 250-350. Some genera are established or re-established: Brachytheciastrum gen. nov. – for the group of species around Brachythecium velutinum; Eurhynchiastrum gen. nov. for Eurhynchium pulchellum; Eurhynchiadelphus gen. nov. for Eurhynchium eustegium; Sciurohypnum (Hampe) Hampe for the group of species around Brachythecium oedi podium, B. reflexum, B. populeum; Oxyrrhynchium (B. S. G.) Warnst., nom. conserv. prop. for the group of species around Eurhynchium hians; Remyella C. Müll. for a group of Malesian and Oceanian species, usually referred to Rhynchostegiella. Camptothecium and Trachybryum are not accepted and are placed in Homalothecium; Steerecleus and Scleropodiopsis are not accepted and are placed in Rhynchostegium. The Chinese endemic species known as Platyhypnidium patulifolium is transferred to Donrichardsia.

Abstract

Ðàññìàòðèâàåòñÿ èñòîðèÿ ïðåäñòàâëåíèé îá îáúåìå ñåìåéñòâà Brachytheciaceae è âõîäÿùèõ â íåãî ðîäîâ è ïðåäëàãàåòñÿ íîâàÿ ñèñòåìà ñåìåéñòâà íà îñíîâå àíàëèçà ìîðôîëîãè÷åñêèõ è ìîëåêóëÿðíûõ äàííûõ. Ñåìåéñòâî âêëþ÷àåò ÷åòûðå ïîäñåìåéñòâà, 41 ðîä è 250-350 âèäîâ. Ðÿä ðîäîâ îïèñàí â êà÷åñòâå íîâûõ è åùå ðÿä ðîäîâ âîññòàíîâëåí: Brachytheciastrum gen. nov. – äëÿ âèäîâ, áëèçêèõ ê Brachythecium velutinum; Eurhynchiastrum gen. nov. äëÿ Eurhynchium pulchellum; Eurhynchiadelphus gen. nov. äëÿ Eurhynchium eustegium; Scuirohypnum (Hampe) Hampe äëÿ âèäîâ, áëèçêèõ ê Brachythecium oedipodium, B. reflexum, B. populeum; Oxyrrhynchium (B. S. G.) Warnst., nom. conserv. prop. äëÿ âèäîâ, áëèçêèõ ê Eurhynchium hians; Remyella C. Müll. äëÿ ãðóïïû âèäîâ èç Ìàëåçèè è Îêåàíèè, îáû÷íî îòíîñèìûõ ê Rhynchostegiella. Camptothecium è Trachybryum îòíåñåíû â ñèíîíèìû ê Homalothecium; Steerecleus è Scleropodiopsis – â ñèíîíèìû ê Rhynchostegium. Êèòàéñêèé ýíäåìèê Platyhypnidium patulifolium îòíåñåí ê ðîäó Donrichardsia.

CONTENTS Subfam. Homalothecioideae ...... 258 1.Early attempts to understand the 1. Homalothecium ...... 258 Brachytheciaceae (1812-1879) ...... 246 2. Brachytheciastrum ...... 259 2.Further attempts to understand Brachythe- 3. Eurhynchiastrum ...... 260 ciaceae (late XIXth cent. – late XXth cent.) . . 249 Subfam. Brachythecioideae ...... 262 2A. Studies at the species level ...... 249 4. Kindbergia ...... 262 2C. Studies at the generic level ...... 249 5. Bryhnia ...... 263 2C. Studies at the familial level ...... 250 6. Eurhynchiadelphus ...... 263 3.Current attempt to understand 7. Myuroclada ...... 264 Brachytheciaceae (2002) ...... 256 8. Scleropodium ...... 264 3A. The circumscription of the Brachytheciaceae . 256 9. Brachythecium ...... 265 3B. Infrafamilial classification ...... 257 10. Unclejackia ...... 268 3C. A new generic system of Brachytheciaceae . . 257 11. Brachytheciella ...... 268

1 — Main Botanical Garden of Russian Academy of Sciences, Botanicheskaya, 4, Moscow 127276 Russia – Ðîññèÿ 127276 Ìîñêâà, Áîòàíè÷åñêàÿ, 4, Ãëàâíûé áîòàíè÷åñêèé ñàä ÐÀÍ 2 — Botanical Museum, P. O. Box 47, University of Helsinki FIN 00017 Finland 246 MICHAEL S. IGNATOV & SANNA HUTTUNEN

12. Sciurohypnum ...... 269 are discernible, is that of Bridel (1812). In this Subfam. Rhynchostegielloideae . . . . 271 second edition of “Muscologia Recentiorum...”, 13. Cirriphyllum ...... 271 Bridel, who included most of the pleurocarps 14. Okamuraea ...... 271 in the genus Hypnum, presented a rather elab- 15. Oxyrrhynchium ...... 272 orated subdivision of this genus, based on four 16. Donrichardsia ...... 273 17. Nobregaea ...... 274 characters of the branching pattern and foli- 18. Eurhynchiella ...... 274 age. Some sections of Hypnum that he estab- 19. Rhynchostegiella ...... 275 lished are similar to volume of modern genera 20. Helicodontium ...... 276 (e. g. ‘fissidentoidea’ ~ Plagiothecium, and 21. Clasmatodon ...... 277 ‘tamariscina’ ~ Thuidium). The Brachytheciaceae 22. Flabellidium ...... 278 in modern sense (except Homalothecium that 23. Homalotheciella ...... 278 was kept in Leskea due to the modified peristome) 24. Mandoniella ...... 279 composed mostly1 6 of the 28 sections that Bridel 25. Schimperella ...... 279 recognized within Hypnum, namely [specific ep- 26. Juratzkaeella ...... 280 27. Aerolindigia ...... 280 ithets are given here in modern nomenclature]: 28. Remyella ...... 281 Sect. Praelonga [foliis distichis, nudis, ramis 29. Meteoridium ...... 281 vagis, ramulis subcompressis]: hians, stokesii, spe- 30. Squamidium ...... 282 ciosum, schleicheri, confertum, rotundifolium (cf. 31. Zelometeorium ...... 282 the more commonly used concept of Eurhynch- Subfam. Rhynchostegioideae . . . . . 283 ium s.l. and Rhynchostegium); 32. Platyhypnidium ...... 283 Sect. Illecebra [foliis spiralis imbricatis, ap- 33. Rhynchostegium ...... 284 pressis, ramis simpliciter pinnatis vel subpinna- 34. Eriodon ...... 285 tis, ramulis teretibus obtusis]: purum, touretii, 35. Aerobryum ...... 286 36. Pseudoscleropodium ...... 287 cirrosum (Pseudoscleropodium, Scleropodium, 37. Bryoandersonia ...... 287 Brachythecium); 38. Eurhynchium ...... 287 Sect. Muralia [foliis spiralis imbricatis, ap- 39. Plasteurhynchium ...... 288 pressis, ramis confertis erectis]: murale (Rhyn- 40. Palamocladium ...... 289 chostegium); 41. Scorpiurium ...... 289 Sect. Velutina [foliis spiralis imbricatis, pat- Epilogue ...... 290 entibus omnibus, ramis confertis erectis]: veluti- Appendix 1 ...... 292 num, starkei, reflexum (cf. small-sized species of Acknowledgements ...... 293 Brachythecium s.l.); Literature cited ...... 293 Sect. Plumosa [foliis spiralis imbricatis, pat- The Brachytheciaceae is a family that is of- entibus omnibus, ramis pinnatis vel subpinna- ten considered taxonomically difficult. This tis]: plumosum, salebrosum, lutescens, aureum, paper gives a brief overview of the history of populeum (cf. some narrow-leaved Brachy- familial and generic concepts, and provides the thecium and Homalothecium species); nomenclatural consequences that are derived Sect. Rutabula [foliis spiralis imbricatis, pat- from phylogenetic inference based on molecu- entibus omnibus, ramis vagis, ramulis subteret- lar and morphological characters (Huttunen ibus]: rutabulum, rivulare, piliferum, touretii, & Ignatov, 2003). megapolitanum, rusciforme (cf. broad-leaved 1. EARLY ATTEMPTS TO UNDERSTAND species of Brachythecium, Cirriphyllum, Scle- THE BRACHYTHECIACEAE (1812-1879) ropodium, Rhynchostegium, Platyhypnidium). The circumscriptions of many widespread This classification is of certain interest for boreal and temperate genera of pleurocarpous the following discussion, because one can see here mosses, and of the Brachytheciaceae in particu- the division of Brachytheciaceae into six more or lar, were established in “Bryologia Europaea...” less natural groups based on character states (Bruch & al., 1851-1855). However, already in 1 – some placements, however, look odd from the current earlier publications of the XIXth century some point of view, for example Brachythecium acuminatum groupings of species were proposed. One of the (sub. Hypnum oxycladon) was in one section with Orthothecium, Tomentypnum, Pleurozium, Calliergon, and first classifications, wherein the Brachytheciaceae Calliergonella, obviously due to strongly appressed leaves. Brachytheciaceae (Bryophyta) – a family of sibling genera 247

Table 1. Number of species in the larger genera of Brachytheciaceae in four publications for Europe (Bruch & al., 1851-55) and the world. In Crosby & al. (1999) the species number given by two values: (recently revised and confirmed species, marked **** in Crosby & al., l.c.) /(all still not synonymized species) ) [roughly speaking: good species/ good plus dubious species]. B.S.G. Jaeger & Sauerbeck Brotherus Crosby & al. 1851-55 1876-79 1925 1999 Homalothecium 3 14 18 8/14 Camptothecium 3 9 17 (in Homalothecium) Brachythecium 24 100 224 94/179 Scleropodium 2 3 8 5/9 Eurhynchium 17 62 31 25/54 Rhynchostegium 10 87 135 30/133 that are quite inappropriate in their application sis Muscorum Europaeorum” (1876). Pleuro- to certain taxa (e. g. ramis vagis is not the case of carps were divided into 6 tribes: Fontinaliace- Hypnum stokesii and H. piliferum, and ramis ae, Neckeraceae, Hookeriaceae, Fabroniaceae, pinnatis is quite a rare case in H. lutescens, etc.). Leskeaceae, Hypnaceae, and the latter included Thus, this classification was essentially intuitive 5 families: 1. Pterigynandreae; 2. Orthothecieae and the subsequent attempt to explain which (Pylaisia, Entodon, Platygyrium, Isothecium, etc., morphological characters were involved was not and Homalothecium); 3. Camptothecieae (Campto- very successful. The usage of these diagnostic char- thecium); 4. Brachythecieae (Brachythecium, Eu- acters to find a correct place for new species rhynchium, Rhynchostegium, Scleropodium, and would be an impossible task. also Hyocomium, Ptychodium, Thamnium, etc.); Bruch, Schimper & Guembel in “Bryologia 5. Hypneae [names are in that day style]. The Europaea...” established several important gen- Brachytheciaceae, as we know them now, fell era of the Brachytheciaceae: Homalothecium into 3 families, due to the differences in sporo- 1851, Rhynchostegium 1852, Brachythecium 1853, phytic characters. The family Hypneae was al- Camptothecium 1853, Scleropodium 1853, and most equal to the present volume of Eurhynchium 1854. Since the idea of the primary Hypnaceae + Amblystegiaceae + Plagiothe- importance of peristome structure still dominat- ciaceae + Hylocomiaceae. The differences be- ed at that time, Homalothecium was placed in tween Hypneae and Brachythecieae include the tribe Orthothecieae (capsule usually straight, characters of the seta (sometimes rough in peristome intermediate between Hypnum and Brachythecieae), capsule (longer in Hypneae) Neckera), whereas all other genera were in Hyp- and areolation (cells often narrowly linear in neae (differs from Orthothecieae in gametophyte Hypneae). It is obvious, that these differences texture (“consistentia”), capsule and peristome). are not enough to delimit these two groups. But The diagnosis of genera was based mainly on it is no less obvious, that Schimper (1876) felt sporophytic characters, though gametophytic ones intuitively how to divide them very well – no were supplemented: Rhynchostegium (opercu- species of the Brachytheciaceae (as we know it lum longly rostrate), Brachythecium (differs from now) was placed in Hypneae (though some non- Rhynchostegium by its conic operculum), Brachytheciacean taxa fell into Brachythecieae). Eurhynchium (intermediate between Rhyncho- Two genera of the Brachytheciaceae were placed stegium and Brachythecium in operculum shape; in two other families: Homalothecium – due differs from Rhynchostegium by shorter cells to its reduced peristome, and Camptothecium and leaf texture), Camptothecium (similar to – due to its transitional characters between Homalothecium in gametophytic characters, but Homalothecium and Brachythecium. its capsule is more like that in Brachythecium), Lindberg (1879) in “Musci Scandinavici in Scleropodium (differs from Brachythecium by Systemate novo naturali dispositi” did the next its narrow flexuose cells and julaceous foliage). step towards improving the classification of the The characters of branching and foliage (used Brachytheciaceae. He used a nomenclature quite by Bridel) were not mentioned. distinct from that of Schimper: Hypnum sensu The family Brachytheciaceae was established Schimp. = Stereodon sensu Lindb., Orthothecie- by Schimper in the second edition of “Synop- ae pp. + Camptothecieae + Brachythecieae = Hyp- 248 MICHAEL S. IGNATOV & SANNA HUTTUNEN

Table 2. List of genera of Brachytheciaceae (* – described as not a member of Brachytheciaceae) *Helicodontium Schwaegr., 1830 (gen. et sp. nov./ South America). *Clasmatodon Hook. & Wils., 1842 (gen. et sp. nov./ North America). *Pleuropus Griff., 1842, nom. illeg., later homonym (gen. et sp. nov./ Asia and Africa). *Eriodon Mont., 1845 (gen. et sp. nov./ Patagonia and Chile). *Homalothecium B. S. G., 1851 (segregated from Leskea/ Europe & al.). [*Anisodon B.S.G., 1852 (gen. et sp. nov./ Germany). – synonymized with Clasmatodon, but probably erroneously, see discussion under the latter genus below]. Rhynchostegium B. S. G., 1852 (segregated from Hypnum/ Europe & al.). Brachythecium B. S. G., 1853 (segregated from Hypnum/ Europe & al.). Camptothecium B. S. G., 1853 (segregated from Hypnum/ Europe & al.). Scleropodium B. S. G., 1853 (segregated from Hypnum/ Europe & al.). Eurhynchium B. S. G., 1854 (segregated from Hypnum/ Europe & al.). *Fiorinia Schimp., 1865, nom. nud. et illeg., later homonym (gen. et sp. nov./ Italy). *Dubyella Schimp., 1866 (replacing Fiorinia). [*Phlogostomum Hampe, 1867, nom. illeg. (replacing Anisodon)]. Pancovia Neck. ex J.J. Kickx, 1867, nom. illeg. (many groups of the family). Sciuro-hypnum (Hampe) Hampe, 1874 (Hypnum sect. Sciuro-Hypnum Hampe, 1867, segregated from Brachyth- ecium and Camptothecium). Scorpiurium Schimp., 1876 (gen et sp. nov. / France and Algeria). Rhynchohypnum (Hampe) Hampe, 1878 (Hypnum subgen. Rhynchohypnum Hampe, 1852, various areas). Oticodium (C. Muell.) Kindb., 1888 (Hypnum sect. Oticodium C. Muell., 1875. Malesia). Bryhnia Kaurin, 1892 (gen. et sp. nov. / Norway). Myuroclada Besch., 1893 (segregated from Hypnum / Japan). Rhynchostegiella (B.S.G.) Limpr., 1896 (Rhynchostegium sect. Rhynchostegiella B.S.G., 1852). Palamocladium C. Muell., 1896 (replacing Pleuropus, nom. illeg.). Remyella C. Muell., 1896, nom. reject. (gen. et sp. nov./ Hawaii). Cirriphyllum Grout, 1898 (segregated from Eurhynchium). Burnettia Grout, 1903, nom. illeg., later homonym (segregated from Pterigynandrum / Japan). Achrolepis (Lindb.) Cardot, 1904, nom. illeg., based on the same type as ealier name [Myuroclada] (Myurel- la subgen. Achrolepis Lindb., 1872 / Japan). Oxyrrhynchium (B.S.G.) Warnst., 1905 (Eurhynchium subg. Oxyrrhynchium B.S.G. 1854). Paramyurium (Limpr.) Warnst., 1905 (Eurhynchium subgen. Paramyurium Limpr. 1896). Okamuraea Broth., 1906 (gen. et sp. nov./ Japan). Homalotheciella (Card.) Broth., 1908 (Homalothecium sect. Homalotheciella Card., 1899). *Chionobryum Glow., 1913 (segregated from Bryum (sic!)/ Italy). Flabellidium Herzog, 1916 (gen. et sp. nov./ Bolivia). Mandoniella Herzog, 1916 (segregated from Helicodontium). Eurhynchiella Fleisch., 1923 (Rhynchostegium sect. Leptorhynchostegium C. Muell., 1899). Pseudoscleropodium (Limpr.) Fleisch., 1923 (Scleropodium sect. Pseudoscleropodium Limpr., 1896). *Platyhypnidium Fleisch., 1923 (segregated from Eurhynchium, Rhynchostegium, and Hygrohypnum). Schimperella Thér., 1926 (gen. et sp. nov./ Madagaskar). Chamberlainia Grout, 1928 (segregated from Brachythecium). *Rigodiopsis Dix. et Thér., 1932. (gen. et sp. nov./ Japan). *Helicodontiadelphus Dixon, 1936 (gen. et sp. nov./ Australia). Bryoandersonia H. Robinson, 1962 [1963] (segregated from Cirriphyllum / North America). Cratoneurella H. Robinson, 1962 [1963] (segregated from Brachythecium). Stokesiella H. Robinson, nom. illeg., later homonym, 1967 (segregated from Bryhnia or Eurhynchium). Trachybryum (Broth.) Schof., 1968 (Camptothecium sect. Trachybryum Broth., 1908). Juratzkaeella Buck, 1977 (segregated from Juratzkaea, a genus of Stereophyllaceae / China). *Donrichardsia H. Crum et L. E. Anderson, 1979 (segregated from Hygroamblystegium or otherwise Eu- rhynchium / North America). Kindbergia Ochyra, 1982 [replacing Stokesiella H. Robinson, 1967]. Steerecleus H. Robinson, 1987 (segregated from Rhynchostegium). Nobregaea Hedenaes, 1992 (gen. et sp. nov./ Madeira). Scleropodiopsis Ignatov, 1998 (gen. et sp. nov./ Altai). Brachytheciella Ignatov, 1999 (gen. et sp. nov./ Papua New Guinea). Unclejackia Ignatov, Norris et Koponen, 1999 (segregated from Chaetomitrium). Brachytheciaceae (Bryophyta) – a family of sibling genera 249 num sensu Lindb. The latter genus included five valid orthographic variants [Anisodon and subgenera: 1. Scleropodium; 2. Pankowia (~Eu- Phlogostomum are included with questionmark, rhynchium s. l.); 3. Rhynchostegium; 4. Brachy- but see comments under Clasmatodon below]. thecium; 5. Pleuropus (species referred now to Note, that Brachytheciacean genera started to Homalothecium). This seems to be the first case appear before “Bryologia Europaea”, but many wherein all the Brachytheciaceae were put to- of them were attributed to the family later gether, in one “genus” (Lindbergs’ equivalent (those are marked with an asterisk). for familial level), and wherein genera described Most of the genera, that appeared after “Bry- from the tropics were included in the classifica- ologia Europaea”, were monospecific. Only six of tion (Pleuropus). This circumscription of the them included more than 5 species (Oxyrrhynch- family was used with some relatively small ium, Rhynchostegiella, Bryhnia, Cirriphyllum, modifications up to the recent time. Chamberlainia, Steerecleus) and all of them lat- 2. FURTHER ATTEMPTS TO UNDERSTAND er were much disputed in regard to the position BRACHYTHECIACEAE (LATE XIXTH CENT. – of some species within or outside their boundaries LATE XXTH CENT.) (i. e. none of six got a widely recognized repu- The further studies of the Brachytheciaceae tation as a stable and clearly delimited group). can be overviewed at three levels: specific, ge- The discussions on three largest genera of neric, and familial ones. the family, concerned the following: (A) Brachythecium was claimed to be too 2A. STUDIES AT THE SPECIES LEVEL heterogeneous and was subdivided into 4-8 sec- Two main tendencies were important at this tions, first introduced by Kindberg (1897), and level. Since the second half of the XIX century then used with just a little alternations (Broth- and until the 1920s, exotic collections were ac- erus, 1905-1909, 1925; Takaki, 1955b; Robinson, tively studied, resulting in descriptions of many 1962; McFarland, 1988; Ignatov, 1998). new species (Tabl. 1: note, that most of taxa were However, only one attempt was made to split sorted into the three largest genera, established on it. Robinson (1962) transferred almost half of European material). Since the second half of XXth the North American species to Chamberlainia, a century, a number of regional revisions of the fam- genus established originally for only one pecu- ily were undertaken, reducing the number of spe- liar species, Chamberlainia acuminata [and two cies, for example in Japan and its adjacent areas synonyms of this species]. Some years later, how- (Takaki, 1955ab, 1956; Noguchi, 1991), North Amer- ever, Robinson (1987), suggested that the previ- ica (Robinson, 1962), Fennoscandia (Nyholm, 1965), ous broad concept of Brachythecium is better. British Isles (Smith, 1978), East India and adja- (B) Rhynchostegium was recognized as a cent areas (Gangulee, 1978), Eastern North Amer- separate genus by most authors. Robinson (1987) ica (Crum & Anderson, 1981), Himalayas (Vohra, found that the group of tropical American spe- 1983), Madeira (Hedenäs, 1992), Latin America cies and the North American R. serrulatus are (McFarland, 1988), Altai Mts. (Ignatov, 1998), Papua not related to the European R. confertum, the New Guinea (Ignatov & al., 1999), West Indies type of the genus, and segregated them to Steere- (Buck, 1999), Australia (Hedenäs, 2002). cleus. McFarland (1994) and Buck (1999) how- Though the exact number and the volume ever did not accept the latter genus. The aquatic of many species remains obscure, it is clear now species where placed in the genus Platyhypnid- that the number of morphologically distinct ium, which was included by most of authors in species of the Brachytheciaceae in the world is Rhynchostegium and, by some, in Eurhynchium. between 250 and 350. The most recent revisions, however, kept it as a 2B. STUDIES AT THE GENERIC LEVEL genus of its own (McFarland, 1994; Ignatov & At this level deviating taxa (especially those al., 1999; Buck, 1999; Hedenäs, 2002). found outside Europe or rare in Europe) were (C) Eurhynchium in the original description recognized as separate genera, including some included already two subgenera – Eurhynchi- subgeneric units that were raised up to the um (in modern sense: striatum, striatulum, circi- generic level. This can be seen from the Table 2, natum, pulchellum, etc.) and Oxyrrhynchium which includes 50[-52] genera that are based (hians, praelongum; and species traditionally re- on species of Brachytheciaceae, excluding in- 250 MICHAEL S. IGNATOV & SANNA HUTTUNEN ferred to Cirriphyllum: piliferum, cirrosum, crass- Myuroclada, Cirriphyllum, Eurhynchium, Scor- inervium, flotowianum, tommasinii). The latter piurium, Oxyrrhynchium, Rhynchostegium, and subgenus was considered as a separate genus by Rhynchostegiella. He correctly added (as has Warnstorf (1905), Fleischer (1923) and Broth- turned out in this study) Eriodon, Okamuraea, erus (1905-1909, 1925), mostly for species around and Homalotheciella, and incorrectly (in the O. hians. However almost all subsequent authors same sense) Stenocarpidium, Ptychodium, (cf. Takaki, 1956; Robinson, 1962; Buck, 1988) Catagoniopsis, and Rigodium. did not agree with this concept, and returned to Fleischer (1923) correctly excluded Steno- the original broad concept of the genus. The group carpidium, Rigodium, and Ptychodium from the of species around E. praelongum was transferred Brachytheciaceae and incorrectly excluded by Robinson (1962) to the genus Bryhnia, but Platyhypnidium and Okamuraea. later he segregated them as a new genus Stokesiel- Brotherus (1925) incorrectly included Ishibaea, la (Robinson, 1967). The latter appeared to be Sciuroleskea, and Lepyrodontopsis; incorrectly illegitimate, and was replaced by Kindbergia retained Tomentypnum and Catagoniopsis in the (Ochyra, 1982). Kindbergia was accepted by Law- Brachytheciaceae, and incorrectly retained Oka- ton (1971) and Crum & Anderson (1981), while muraea outside the Brachytheciaceae. most authors kept this group in Eurhynchium Grout (1931) correctly included Platy- (cf. Noguchi, 1991; Buck, 1988, Hedenäs, 1992, 2002; hypnidium (within Eurhynchium), and incor- Ignatov, 1998). Some authors united Eurhynchium rectly kept Isothecium and Pseudoisothecium. (or part of it) and Rhynchostegium under the Robinson (1962) incorrectly included Con- former (Grout, 1931, Koponen, 1987) or latter ardia and Isothecium. (McFarland, 1994) name. Buck (1988) incorrectly included Rozea, Some species were moved from one genus to Stenocarpidiopsis, and Bryostreimannia. another many times: Hypnum ornellanum was There is no need to comment all these trans- placed in Brachythecium, Cirriphyllum, and fers. They were numerous and arguments for Scleropodium; Hypnum praelongum in Eurhynch- them were short if any. ium, Oxyrrhynchium, Bryhnia, and Kindbergia, etc. In the Northern Hemisphere the main problem Summing up, despite various attempts to make was the border between Brachytheciaceae and the three biggest genera of Brachytheciaceae Amblystegiaceae. Important steps forwards were (Branchythecium, Eurhynchium, Rhynchostegium) taken by Hedenäs (1989a), who found [among smaller and more homogeneous, no attempt got a others] a set of characters to separate the Brachy- wide recognition, except for some cuttings off theciaceae+Hylocomiaceae+Ctenidiaceae from the monotypic (Myuroclada, Bryoandersonia, etc.) or Amblystegiaceae+Thuidiaceae+Hypnaceae (at more rarely oligotypic genera. The reason for this least the main groups of these families). Among is that the infraspecific variation in many species these characters are the appearance of the costa is very broad, with some extremes extending across and the plicateness in perichaetial leaves, the generic boundaries. Thus, the affinities of some shape of the stomata pore, the capsule shape, the species remain ambiguous. Robinson (1962, p. 78) pattern of exostome teeth border tapering, etc. expressed the situation as follow: “Particularly Thus, Hedenäs excluded from Brachythecia- important at the specific level are cases where no ceae Isothecium (Hedenäs, 1992), Tomentypnum single specimen displays the full range of poten- (Hedenäs, 1987), Conardia (Hedenäs, 1989b), all tial variation. Individual specimens of such high- correctly according to our studies. These publi- ly variable species often show a very limited and cations of Hedenäs install the reliable and sharp distinctive morphology”. This is quite applicable border between Brachytheciaceae and Amblys- at the generic level as well. tegiaceae. Though recently Amblystegiaceae got 2C. STUDIES AT THE FAMILIAL LEVEL a more definite and narrow circumscription The attempts to circumscribe the family were (Vanderpoorten & al., 2002), the ideas of He- many: denäs remain valid, because the pool of genera Brotherus (1905-1909) included into the around Amblystegiaceae + Leskeaceae + Calli- Brachytheciaceae its core genera, Homalotheci- ergonaceae + Rhytidiaceae + etc. is still a unit um, Camptothecium, Brachythecium, Bryhnia, quite distant from the Brachytheciaceae. Brachytheciaceae (Bryophyta) – a family of sibling genera 251

Almost simultaneously with the resolution The few exceptions were, however, found. The of the limits of the Brachytheciaceae in the bo- genus Trachypus, which clearly belongs to the real flora, the problem of its limits in the trop- Meteoriaceae clade on the basis of phylogenetic ics was highlighted. inference and also overall morphology (Buck, (1) In revisions of the genus Lindigia (Mete- 1994, 1999), has pseudoparaphyllia that differ oriaceae) L. capillacea was transferred to Brachy- from the Brachytheciacean+Meteoriacean type. theciaceae by Menzel (1991), to the new genus Lindigia debilis and Neobarbella comes have Aerolindigia, and by Visnadi & Allen (1991), this type of pseudoparaphyllia, but the former to Rhynchostegiella. was found outside the group of these two fam- (2) Duthiella, a genus commonly placed in ilies in the present phylogenetic analysis, while Meteoriaceae or Trachypodaceae was placed in the latter – in analysis of Quandt & al. (2003b). the Brachytheciaceae by Noguchi (1991). While a close relationship between Brachy- (3) Buck (1994) re-evaluated the Meteori- theciaceae and Meteoriaceae was found, the bor- aceae and found it very difficult to separate der between them appeared problematic. Since from Brachytheciaceae. Later Buck (1999) they were not considered closely related and were placed these families close to each other. placed even in different orders (the first author (4) Ignatov, Koponen & Norris (1999) found who placed them closely was Buck, 1999) – their similar Brachytheciaceae/Meteoriaceae problems relationship has not been a subject for a compar- in New Guinea, around species of Unclejackia, ative studies. Meteoriaceae were understood by Brachytheciella, and Rhynchostegiella. having pendent growth (almost never so in (5) Ignatov (1999) found that members of Brachytheciaceae in a traditional sense), often both the Brachytheciaceae and the Meteoriaceae papillose laminal cells (smooth in the Brachythe- have similar pseudoparaphyllia pattern, which ciaceae), seta often short (long in Brachytheci- separate these two families from all other pleuro- aceae), calyptra often hairy (very rarely so in carps. Ignatov (l. c.) confirmed the exclusion from Brachytheciaceae); a modified hygrocastique the Brachytheciaceae of Isothecium, Conardia, and peristomes (Brachytheciaceae have usually more Tomentypnum, and did not confirm the posi- or less complete xerocastique peristomes with ex- tion in this family of Rozea, Sciuroleskea, Steno- ostome teeth striolate below and an endostome carpidiopsis, Catagoniopsis, and Lepyrodontop- with a high basal membrane, broad and broadly sis. The position of Trachypus in Meteoriaceae perforated segments, and long cilia). However, was not confirmed [which however, appeared these characters are not consistent across taxa. to be an exceptional case – the genus fell well The single special study of Brachytheciaceae within Meteoriaceae, both in molecular and over- based on sequence data (Stech & Frahm, 1999), all morphological studies (Quandt & al., 2003a; included eight species, mostly of the Eurhynch- Huttunen & Ignatov, 2003; also Figs. 1,2)]. ium group. It was found that Eurhynchium However due to the limited number of gen- pumilum is more closely related to Rhynchost- era studied, Ignatov (1999) failed to find any egiella jacquinii, whereas Eurhynchium stria- genera outside Brachytheciaceae+Meteoriaceae tum is neither related to E. (Oxyrrhynchium) that can be included in this group of two fam- hians, nor to E. (Kindbergia) praelongum. A ilies based on pseudoparaphyllia pattern. number of species were included also in the Soon afterwards, however, molecular data analyses of Buck & al. (2000), Goffinet & al., showed that Brachytheciaceae includes also Clas- (2001), Stech & Frahm (2001) and Tsubota & matodon (Buck & Goffinet, 2000), Helicodontium al. (2002). (Buck & al., 2000), Okamuraea (Tsubota & al., Our analysis of sequence and morphological 2002), and Donrichardsia (Stech & Frahm, 1999; data is published separately (Huttunen & Igna- Vanderpoorten & al., 2002). All these rather un- tov, 2003). Consensus of three most parsimoni- expected transfers were supported by our subse- ous trees based on combined analysis of molec- quent pseudoparaphyllia studies. This high con- ular and morphological data is presented in Fig. gruence of molecular and morphological data is 1. This result is obtained using direct optimi- especially pleasant, raising the trust to molecular zation method as implemented in phylogenet- data by morphologists and perhaps vice versa. ic software POY (Gladstein & Wheeler, 2001, 252 MICHAEL S. IGNATOV & SANNA HUTTUNEN

Leskea polycarpa Rozea andrieuxii Rozea subjulacea Hypnum cupressiforme Ctenidium molluscum Isothecium alopecuroides Lembophyllum clandestinum Bestia longipes Dolichomitriopsis diversiformis Meteoriella soluta Hylocomium splendens Rhytidiadelphus triquetrus Lindigia debilis Leucodon sciuroides Plagiothecium denticulatum Amblystegium serpens Campylium stellatum Trachypus bicolor Chrysocladium retrorsum Meteorium polytrichum Duthiella wallichii Aerobryidium filamentosum Pseudospiridentopsis horrida Floribundaria floribunda Barbella flagellifera ETEORIACEAE Trachycladiella aurea M Scorpiurium circinatum Scorpiurium deflexifolium Plasteurhynchium striatulum Aerobryum speciosum Pseudoscleropodium purum Eurhynchium angustirete Eurhynchium striatum Palamocladium euchloron Palamocladium leskeoides Rhynchostegium psilopodium Platyhypnidium ri parioides Rhynchostegium confertum Rhynchostegium pallidifolium Rhynchostegium serrulatum HYNCHOSTEGIOIDEAE

Rhynchostegium rotundifolium R Rhynchostegium murale (North Urals) Rhynchostegium murale (China) Rhynchostegium murale (South Urals) Meteoridium remotifolium Squamidium brasiliense Zelometeorium patulum Bryoandersonia illecebra Rhynchostegiella tenella

Rhynchostegiella teneriffae RACHYTHECIACEAE

Rhynchostegiella macilenta B Platyhypnidium austrinum Okamuraea brachydictyon Brachythecium koponenii Cirriphyllum crassinervium Cirriphyllum piliferum Aerolindigia capillacea Rhynchostegiella brachypodia Clasmatodon parvulus Helicodontium capillare Homalotheciella subcapillata Eurhynchium pumilum Eurhynchium vagans (China) HYNCHOSTEGIELLOIDEAE Donrichardsia macroneuron R Platyhypnidium patulifolium Eurhynchium savatieri Eurhynchium hians Eurhynchium vagans var. bergmaniae

BRACHYTHECIOIDEAE, HOMALOTHECIOIDEAE (see next page) Brachytheciaceae (Bryophyta) – a family of sibling genera 253

RHYNCHOSTEGIOIDEAE, RHYNCHOSTEGIELLOIDEAE and outgroup taxa (see previous page) Myuroclada maximowiczii Scleropodium obtusifolium Brachythecium collinum Brachythecium falcatulum Brachythecium velutinum Eurhynchium pulchellum Homalothecium aureum Homalothecium laevisetum Homalothecium nuttalii Homalothecium megaptilium

Homalothecium fulgescens Brachytheciastrum Homalothecium lutescens OMALOTHECIOIDEAE

Homalothecium philippeanum H Homalothecium sericeum Eurhynchium eustegium Bryhnia novae-angliae (Europe) Bryhnia novae-angliae (Altai) Kindbergia praelonga (China) Kindbergia praelonga (Colombia) Kindbergia praelonga ‘serricuspis’ Brachythecium plumosum Brachythecium populeum Cirriphyllum flotovianum Brachythecium hylotapetum Brachythecium oedipodium Brachythecium reflexum Brachythecium ornellanum

Brachythecium glaciale Sciurohypnum Brachythecium latifolium Bryhnia novae-angliae (China) ß !!! Brachythecium frigidum Brachythecium rutabulum Brachythecium rivulare (Finland) Brachythecium rivulare (China) Brachythecium cirrosum (Yakutia) Brachythecium cirrosum (China) Brachythecium complanatum Brachythecium salebrosum (South Russia) RACHYTHECIOIDEAE

Brachythecium ‘subalbicans’ B Brachythecium mildeanum Brachythecium albicans Brachythecium erythrorrhizon Brachythecium salebrosum (Finland) Brachythecium turgidum Cirriphyllum tommasinii Brachythecium geheebii (Urals) Brachythecium geheebii (Poland) Brachythecium udum

Brachythecium acuminatum Brachythecium Brachythecium laetum Brachythecium glareosum Brachythecium lamprocarpum Unclejackia longisetula ß !!! Brachythecium ruderale Brachythecium buchananii (China) Brachythecium buchananii (China)

Fig. 1 (both this and opposite pages). Strict consensus tree of 3 most parsimonious trees (L=3546; CI=14; RI=88) from POY analysis of combined morphological and sequence data with gap cost 1. [Brachyth- ecium ‘subalbicans’ = Brachythecium cf. salebrosum from China, similar to the type of B. subalbicans Broth., nom. illeg. (its holotype described and illustrated by Wang & Hu, 1998); Kindbergia praelonga ‘serricuspis’ – very thiny phenotype of the species, earlier considered as an independent species Eurhynch- ium serricuspis C. Muell. See comments on Bryhnia novae-angliae and Unclejackia longisetula (marked with !!!) under those genera. 254 MICHAEL S. IGNATOV & SANNA HUTTUNEN

Leskea polycarpa 100 Rozea andrieuxii >15 Rozea subjulacea 73 Leucodon sciuroides 7 Lindigia debilis Bestia longipes 100 14 100 Dolichomitriopsis diversiformis 4 79 Isothecium alopecuroides 100 6 Lembophyllum clandestinum 79 Hypnum cupressiforme

5 Ctenidium molluscum 96 Plagiothecium denticulatum 9 Amblystegium serpens 53 >15 Campylium stellatum 1 86 Meteoriella soluta 68 8 60 Hylocomium splendens 1 1 Rhytidiadelphus triquetrus 81 Trachypus bicolor 99 Chrysocladium retrorsum 7 15 Meteorium polytrichum 99 61 Duthiella wallichii 15 76 Aerobryidium filamentosum 92 3 5 Pseudospiridentopsis horrida 7 95 Floribundaria floribunda

ETEORIACEAE 94 Barbella flagellifera 7 M

8 Trachycladiella aurea Bryoandersonia illecebra 51 64 Pseudoscleropodium purum 6 100 Scorpiurium circinatum 2 68 14 Scorpiurium deflexifolium 7 Plasteurhynchium striatulum 88 Palamocladium euchloron 1 8 Palamocladium leskeoides 79 Eurhynchium angustirete 1 3 Eurhynchium striatum Aerobryum speciosum Platyhypnidium riparioides Rhynchostegium confertum Rhynchostegium psilopodium 84 Rhynchostegium rotundifolium

5 75 Rhynchostegium pallidifolium HYNCHOSTEGIOIDEAE 4 Rhynchostegium serrulatum R Rhynchostegium murale (China) 52 Rhynchostegium murale (North Urals) 3 Rhynchostegium murale (South Urals) Myuroclada maximowiczii 91 100 Meteoridium remotifolium 9 100 >15 Squamidium brasiliense 14 Zelometeorium patulum 99 Donrichardsia macroneuron

94 9 Platyhypnidium patulifolium RACHYTHECIACEAE Eurhynchium hians B 10 4 60 Eurhynchium pumilum Eurhynchium savatieri 93 62 Eurhynchium vagans var. bergmaniae 6 Eurhynchium vagans (China) Platyhypnidium austrinum Oxyrrhynchium 100 Rhynchostegiella tenella 68 100 Rhynchostegiella teneriffae 14 1 11 Rhynchostegiella macilenta 78 Okamuraea brachydictyon 76

8 Cirriphyllum crassinervium HYNCHOSTEGIELLOIDEAE 6 6 Brachythecium koponenii R 52 5 Cirriphyllum piliferum 3 86 Aerolindigia capillacea 97 6 Rhynchostegiella brachypodia Clasmatodon parvulus 7 90 Helicodontium capillare 5 Homalotheciella subcapillata BRACHYTHECIOIDEAE, HOMALOTHECIOIDEAE (see next page) Brachytheciaceae (Bryophyta) – a family of sibling genera 255

RHYNCHOSTEGIOIDEAE, RHYNCHOSTEGIELLOIDEAE, Myuroclada and outgroup taxa (see previous page) Scleropodium obtusifolium Brachythecium collinum 66 Brachythecium falcatulum 7 Brachythecium velutinum 4 Eurhynchium pulchellum 4 52 Homalothecium aureum 58 Homalothecium laevisetum 4 5 69 6 Homalothecium nuttalii 7 Homalothecium megaptilium 71 Homalothecium fulgescens Brachytheciastrum 4 64 Homalothecium lutescens 4 OMALOTHECIOIDEAE 76 Homalothecium philippeanum H 3 4 Homalothecium sericeum Cirriphyllum tommasinii Brachythecium acuminatum 57 Brachythecium complanatum 4 Brachythecium glareosum Brachythecium laetum Brachythecium salebrosum (Finland) Brachythecium turgidum 99 Brachythecium geheebii (Urals) 9 Brachythecium geheebii (Poland) 68 Brachythecium albicans 2 Brachythecium erythrorrhizon 99 Brachythecium salebrosum (South Russia) 9 Brachythecium ‘subalbicans’ 68 Brachythecium mildeanum 1 Brachythecium udum Brachythecium lamprocarpum 1 Unclejackia longisetula 100 Brachythecium cirrosum (Yakutia) 65 13 Brachythecium cirrosum (China) 5 Brachythecium buchananii (China) 94 Brachythecium buchananii (China) 6 Brachythecium ruderale 83 Brachythecium frigidum 53 6 Bryhnia novae-angliae (China)

RACHYTHECIOIDEAE

Brachythecium rutabulum B 1 100 75 10 Brachythecium rivulare (China) 1 Brachythecium rivulare (Finland) Eurhynchium eustegium 99 71 Bryhnia novae-angliae (Europe) 12 Bryhnia novae-angliae (Altai) 1 Kindbergia praelonga (Colombia) 99 87 Kindbergia praelonga (China) 11 4 Kindbergia praelonga ‘serricuspis’ Brachythecium plumosum 100 12 66 Brachythecium populeum 96 3 Cirriphyllum flotovianum 9 92 Brachythecium hylotapetum 77 10 Brachythecium oedipodium 2 86 Brachythecium reflexum Brachythecium ornellanum 5 79 4 88 Brachythecium glaciale Sciurohypnum 4 Brachythecium latifolium

Fig. 2 (both this and opposite pages). Strict consensus tree of 779 most parsimonious trees (L=2436; CI=29; RI=67) from Nona analysis based on POY character optimization on combined morphological and sequence data with gap cost 1. Jackknife values for clades are presented above and bremer support below branches. See also comments to Fig. 1. 256 MICHAEL S. IGNATOV & SANNA HUTTUNEN

Meteoridium, Zelometeorium, and Squamidium, that have been traditionally placed into Me- teoriaceae. Their position in the Meteoriaceae is, however, somewhat odd due to their smooth laminal cells, which make them different from most of the taxa in this family. According to Buck & Goffinet (2000) and also our results, they should be included in the Brachytheci- Fig. 3. Pseudoparaphyllia of Don- aceae. Thus, the Meteoriaceae are quite nicely richardsia macroneuron (x 350) separated from Brachytheciaceae by having (left) and scheme of psuedopara- phyllia arrangement around branch papillose laminal cells. Problems with the clas- initial (from Ignatov, 1999)(above). sification of the Meteoriaceae are discussed by Quandt & al. (2003a) and Huttunen (in prep.). for details see Huttunen & Ignatov, 2003). Due As defined here, the family Brachytheciace- difficulties to calculate branch support values ae is monophyletic and can be surprisingly eas- in POY, jackknife and bremer support for clades ily delimited by only two morphological charac- were obtained from Nona analysis of the POY ters: Brachythecium-like pseudoparaphyllia alignment (Fig. 2). This Nona tree is the main pattern (Fig. 3) and laminal cells without pa- used for the subsequent discussion. However, pillae (with two exceptions of Neobarbella and clades lacking support are collapsed in Nona Lindigia, which otherwise share this definition, tree (Fig. 2), but not in the optimal tree result- but according to the present results do not be- ing from the POY analysis (Fig. 1). Hence our long to the family). Some Brachytheciaceae have discussion will refer to the results from POY papilla-like prorae in cell corners, which are to- analyses (Fig. 1) when addressing the affinities tally different from round “true” papillae above and circumscription of clades that are poorly cell luminae or cell walls in Meteoriaceae. De- resolved in Nona topology (Fig. 2). limited in this way, the Brachytheciaceae in- 3. CURRENT ATTEMPT TO UNDERSTAND cludes 41 genera with 250-400 species. BRACHYTHECIACEAE (2002) * * * 3A. THE CIRCUMSCRIPTION OF THE Crosby & al. (1999) included in the Brachy- BRACHYTHECIACEAE theciaceae seven genera which do not belong The Meteoriaceae and Brachytheciaceae form neither to Brachytheciaceae, nor to Meteori- a monophyletic group in POY analysis (Fig. 1), aceae according to morphological data (and but support for a shared ancestry in Nona ana- in some cases also molecular data – those lysis is lacking (Fig. 2). There are only two spe- marked by asterisk): Bestia*, Bryostreiman- cies, Lindigia debilis (Mitt.) Jaeg. and Neobar- nia, Isothecium*, Pseudopleuropus, Puiggari- bella comes (Griff.) Noguchi (the latter not shown opsis, Rozea*, Stenocarpidiopsis. Ignatov's in the present analysis, but see Quandt & al., (1999) statement that Squamidium does not 2003b), with the pseudoparaphyllia pattern of the have pseudoparaphyllia of the Brachytheci- Brachytheciaceae and Meteoriaceae, which do not acean type was based on misidentified mate- appear within the Brachytheciaceae + Meteoriaceae rial. Also the hypothesis that Meteoriella solu- clade. These exceptions will be discussed elsewhere. ta is a member of the Brachytheciaceae (Igna- Because both Lindigia and Neobarbella are not tov l. c.) is currently being reexamined and regarded as members of the Brachytheciaceae, they the results will be discussed separately. are not discussed in the present paper. Goffinet & al. (2001) found Hildenbrandtiel- The Brachytheciaceae-Meteoriaceae clade la pachyclada Besch. in the most basal position (Fig. 1) includes two independent lineages. The of the Brachytheciaceae clade (in analysis of first clade consists of typical Meteoriaceae sen- rps4 data). However, this species has narrowly su lato including former Trachypodiaceae (Buck, lanceolate pseudoparaphyllia, suggesting [among 1994, Quandt & al., 2003a). The other one in- other morphological data] that its previous po- cludes Brachytheciaceae taxa together with a sition within Pterobryaceae (cf. Buck & Goffi- few genera, such as Aerobryum, Aerolindigia, net, 2000) is likely to be corrrect. Brachytheciaceae (Bryophyta) – a family of sibling genera 257

3B. INFRAFAMILIAL CLASSIFICATION 1 to 6 cells high, below the leaf insertion and around Previous authors usually did not subdivide branch primordia (outside of pseudoparaphyllia). the family into subfamilies or tribes. Limpricht Smooth laminal cells – since the evaluation of a (1896) sorted European taxa into Brachyste- smooth vs. a slightly prorate state, as we believe, de- pends much on microscope quality, we do not sepa- giae (with short operculum) and Eustegiae rate them. Only the cases where projections are high, (with rostrate operculum), and some subsequent tooth-like, and rather common, they are contrasted authors followed him (e. g. Roth, 1905). The to former condition. Defined in this way, all genera application of such descriptive names is allowed have smooth cells except Bryhnia, Kindbergia, Eu- by the ICBN for taxa above the family level, rhynchiadelphus, and Plasteurhynchium. without comments on what to do with such Spore size is categorized in two states: “spores taxa at familial and infrafamilial levels. We large” means that at least a few spores are 25 mm or suggest to ignore these Limpricht' subfamilies, larger; otherwise spores are called small. Most spe- m as they do not refer to any generic name. cies have spores ranging between 14 and 22 m. The presence of very small spores (10-14 mm) in some The present analysis of Brachytheciaceae reveals taxa is specifically underlined. four well-supported clades, which are centered by Subalar cells. The highest differentiation among Homalothecium, Brachythecium, Oxyrrhynchium laminal basal cells in the Brachytheciaceae occurs + Rhynchostegiella, and Rhynchostegium. Of 98 in the area where the line of the leaf insertion makes specimens of ca. 90 species from 33 genera of maximal turn to the decurrency. Cell differentiation Brachytheciaceae, two species were found in rath- starts from this point and sometimes differentiated er indefinite position, Myuroclada maximowiczii cells reach the leaf margin, thus allowing to use the and Scleropodium obtusifolium. They are prelimi- term “alar cells” (=cells of basal leaf angles). How- nary placed and discussed within the Brachy- ever, in many cases (Eurhynchium, Bryhnia, etc.) the group of differentiated cells remains quite far from thecioideae. Eight genera (Brachytheciella, Nobre- leaf angle. We suggest to call them “subalar cells”. gaea, Eurhynchiella, Flagelledium, Mandoniella, Schimperella, Juratzkaeella, Eriodon), very rare and BRACHYTHECIACEAE Schimp., Syn. mostly monospeciefic and known mostly from the Musc. Eur., ed. 2. XCV sic [CXV], 637. 1876, holotypes of type species, were not included in mo- “Brachythecieae”. Type: Brachythecium B. S. G. lecular studies; their placements are based on Camptotheciaceae Schimp., Syn. Musc. Eur., morphology and are considered as preliminary. ed. 2. XCV, 635. 1876, “Camptothecieae”. Type: 3C. A NEW GENERIC SYSTEM OF Camptothecium B. S. G. BRACHYTHECIACEAE Donrichardsiaceae Ochyra, Lindbergia 11: 7. The genera as circumscribed below are supple- 1985. Type: Donrichardsia Crum et Anderson. mented with lists of species; the “species includ- Diagnostic characters: first (earliest in dif- ed” sections are based on specimens checked by ferentiation) pseudoparaphyllium [=juvenille ourselves. These lists of species serve to illustrate branch leaf, scaly leaf] faced downward, sec- our concepts of genera and must not be consid- ond and third – at the angle 120° to the first. ered as complete and taxonomically and nomen- Laminal cells without papillae above central claturally verified. They are based largely on part of lumina. Stomatal pore round. Crosby & al. (1999) list, which summarized data Besides these character states there are no from revisions, check-lists, and old publications. morphological features that consistently charac- Generic descriptions and distribution are terize members of Brachytheciaceae, and at the based only on species from “species included”, same time excluding members of other families. not from problematic species (exception of Hence the family description is rather lowly in- Bryhnia specially discussed under that genus). formative: Stem creeping, ascending to erect or pendent, sometimes with stolons; differentiation NOTES ON TERMINOLOGY: into primary and secondary stems absent or Axillary hairs – number of cells refers to whole present; central strand present or absent; branch- axillary hair, not only to upper cells. Paraphyllia – there is no big similarity between ing irregular to pinnate. Foliage julaceous or com- paraphyllia of the Amblystegiaceae or the Leskeace- planate, very dense to very lax; stem leaves com- ae and that of the Kindbergia (Fig. 4). However, monly ovate-lanceolate, sometimes orbicular, some- seems there is no better term for longitudinal ridges, times narrowly linear; costa present, rarely ab- 258 MICHAEL S. IGNATOV & SANNA HUTTUNEN sent; laminal cells linear, elongate, rhombic, ovate Trachybryum (Broth.) W. B. Schofield, J. or round, thin- or thick-walled; alar cells not dif- Hattori Bot. Lab. 31: 222. 1968. – Camptoth- ferentiated, or small and opaque, or large and ecium sect. Trachybryum Broth., Nat. Pfl. 1(3): inflated. Branch leaves similar to stem leaves or 1140. 1908. Type: Camptothecium megaptilum differentiated. Autoicous, dioicous, polyoicous, syn- Sull. (= T. megaptilum (Sull.) W. B. Schofield.) oicous, or phyllodioicous. Perichaetial leaves Oticodium (C. Müll.) Kindb., Enum. Bryin. smooth or plicate, costate or ecostate. Seta short Exot. 27. 1888. – Hypnum sect. Oticodium C. (capsule immersed to emergent) or long, smooth Müll., Linnaea 39: 469. 1875. Type: Hypnum or rough. Capsule erect, inclined, horizontal, or celebesiae C. Müll. (= O. celebesiae (C. Müll.) pendent. Operculum conic to longly, obliquely Kindb.) = Homalothecium laevisetum Sande Lac. rostrate. Annulus separating by fragments, of 2-3 Plants large to medium-sized, green to golden rows of large cells, or absent. Peristome xerocas- or pale yellow. Stem prostrate to ascending, with tique or hygrocastique, in latter case variously re- central strand, densely foliate, irregularly to regu- duced; exostome present to reduced (Clasmat- larly pinnately branched, branches straight or odon), when developed proximally cross-striolate, curved, densely foliate. Axillary hairs 3-6-celled, papillose or smooth. Endostome completely de- upper cells often relatively short. Pseudoparaphyl- veloped (with 3-4 shortly appendiculate cilia) to lia acute or acuminate. Stem leaves erect-appressed reduced to short basal membrane (free segments when dry, erect to erecto-patent when wet, lan- lacking, some segment material adherent to teeth, ceolate or narrowly triangular, being widest just cilia none). Spores small or large. Calyptra cucul- above leaf base, acuminate, strongly longitudinal- late to somewhat mitrate, naked to hairy. ly plicate; margin serrulate to subentire, some- SUBFAM. HOMALOTHECIOIDEAE times with sharp backward recurved teeth in SUBFAM. NOV. alar region; costa ending above mid-leaf to per- Plantae minutae vel robustae. Folia caulina ova- current, at end smooth or toothed; laminal cells ta vel anguste lanceolata, acuta vel acuminata; elongate to linear, thick-walled; basal cells short, folia ramulina acuta, acuminata vel obtusa; costa small and thick-walled, forming opaque area across crassa vel tenuis; cellulae elongatae vel lineares. the base; alar cells isodiametric, forming more or Autoica, dioica, synoica, polyoica vel phyllodio- less well delimited group. Branch leaves not dif- ica. Seta scabra vel laevissima. Capsula erectis vel ferentiated or somewhat narrower and with a horizontalis. Peristomium perfectum vel imper- more serrate margin and frequently larger than fectum. Operculum rostratum vel conicum. stem leaves. Dioicous or phyllodioicous. Pericha- Type species: Homalothecium B. S. G. etial leaves with straight or more rarely reflexed acumina. Seta rough or smooth. Capsule erect, sym- Plants often rather rigid, rich green to rich metric or more or less curved to inclined. Annu- golden, more rarely pale. Leaf margin toothed to lus separating by fragments. Operculum highly the base, more rarely subentire; costa frequently conic and gradually tapered to rostrum or more ending in spine; laminal cells narrowly linear; rarely conic. Peristome hygrocastique, rarely xero- basal cells frequently small, thick-walled; alar cells castique, perfect to modified; exostome teeth be- small, subquadrate, in subquadrate group. Auto- low cross striolate, papillose, or smooth; endos- icous, dioicous, synoicous, polyoicous, phyllodio- tome perfect (with high basal membrane, broad icous. Seta rough or smooth. Operculum conic or segments and cilia more than half of segment rostrate. Peristome complete to strongly modified. length) to strongly reduced (low basal membrane Calyptra smooth, rarely hairy. Xeromesophytic. with rudimentary segments). Spores small. Ca- Genera included: Homalothecium, Brachy- lyptra naked or rarely hairy. Xeromesophytic. theciastrum, Eurhynchiastrum. Distribution: Eurasia and North America. 1. Homalothecium B. S. G., Bryol. Eur. 5: 91. (a) Homalothecium is monophyletic and well 1851. Type: H. sericeum (Hedw.) B. S. G. (= supported if the species sometimes considered in Leskea sericea Hedw.). Campthothecium and Trachybryum are included. Camptothecium B. S. G., Bryol. Eur. 6: 31. (b) The genus Camptothecium was segregated 1853. Type: C. lutescens (Hedw.) B. S. G. (= from Homalothecium based principally on the Hypnum lutescens Hedw.). better developed to almost complete peristome. Brachytheciaceae (Bryophyta) – a family of sibling genera 259

However, the type of Homalothecium, H. sericeum, H. pinnatifidum (Sull. et Lesq.) Lawt. appears in phylogenetic analysis more closely re- H. sericeum (Hedw.) B. S. G. lated to Camptothecium (Homalothecium) lute- Hofmann (1998) published a world-wide re- scens, the type of Camptothecium, than to the Amer- vision of the genus, wherein she discusses eight ican species with modified peristome, like H. nut- more species, which she excludes from the cir- talii. The results of our analysis show that species cumscription, but without offering alternative with rather well-developed peristome are basal in systematic placements (mostly in Brachy- the Homalothecium clade, and that taxa with thecium). The affinities of one species, Camp- modified peristomes are grouped by their geograph- tothecium subhumile Broth., remain unclear, as ic distribution. This supports the conclusion of Ig- the type was not found in H-BR. natov & al. (1998) based on SEM studies, that there are at least two independent lineages of peris- 2. Brachytheciastrum gen. nov. Type: B. ve- tome modification in the genus. Note, that the North lutinum (Hedw.) Ignatov et Huttunen (= Hyp- American pattern of peristome modification is dif- num velutinum Hedw.). ferent from that in Eurasian species. Plantae minutae vel mediocres. Folia erecta (c) The only species of Trachybryum, T. vel falcata, lanceolata vel lanceolata-ovata, mar- megaptilum appears well nested within Homalo- gine toto serrulata vel serrata; costa ad medium thecium in all analyses. This species is peculiar vel ultra evanida vel percurrenta; cellulae lin- in its large size and quite regular pinnate branch- eares, cellulae alares subquadrates, crassiuscules. ing. However, pinnate branching is not a unique Autoica, synoica vel polyoica. Seta scabra vel lae- case in the genus. For example, the North Ameri- vissima. Capsula horizontalis, subventricosis. Peris- can H. nuttalii and H. pinnatifidum are always tomium perfectum. Operculum conicum. very regularly pinnate, although sometimes not Plants small to medium-sized, green to golden, conspicuously so because of curved branches. more rarely pale yellow. Stem prostrate, with cen- (d) Homalothecium laevisetum is found tral strand, densely or loosely foliate, irregularly together with the rest of the genus in most of the to at places regularly pinnately branched, branches analyses, although the chloroplast genes suggest straight, densely or loosely, julaceously or com- a position within a Rhynchostegium-clade, close planately foliate. Axillary hairs 2-3(-4)-celled, to Palamocladium leskeoides. The problem needs upper cells relatively short and blunt. Pseudopara- additional studies. However, it is worth to notice phyllia acute. Stem leaves erecto-patent, erect, that this species is probably the most isolated in imbricate or falcate-secund, lanceolate to ovate, the genus based on morphology – it has a strong- acuminate, slightly longitudinally plicate to ly reduced peristome, and the shoots (in south- smooth; margin serrulate to subentire, costa end- ern populations) are sometimes attenuate, becom- ing above mid-leaf to percurrent, at end smooth ing stoloniferous. If further studies support the or toothed; laminal cells elongate to linear, thick- isolated position of this species, Oticodium would walled; alar cells isodiametric, forming more or be the correct generic name for this segregate. less well delimited subquadrate group. Branch (e) Brachythecium geheebii was transferred leaves somewhat narrower, with more serrate mar- to Homalothecium by Wigh (1973). This spe- gin and back of costa, and usually smaller than cies is commented under Brachythecium. stem leaves. Autoicous, more rarely synoicous to Species included: polygamous. Perichaetial leaves with reflexed Homalothecium aeneum (Mitt.) Lawt. acumina. Seta rough or smooth. Capsule inclined H. arenarium (Lesq.) Lawt. to horizontal, often ventricose, remains greenish H. aureum (Spruce) H. Robinson until almost mature. Annulus separating by frag- H. fulgescens (Mitt.) Lawt. ments. Operculum conic. Peristome xerocastique, H. laevisetum Sande Lac. perfect. Spores small, mostly, ca. 10-13 mm. Calyp- H. lutescens (Hedw.) H. Robinson tra naked. Xeromesophytic or mesophytic. H. megaptilum (Sull.) H. Robinson Distribution: subcosmopolite. H. nevadense (Lesq.) Ren. et Card. (a) This group of species was recognized by H. nuttallii (Wils.) Jaeg. Brotherus (1925) and many other authors as the H. philippeanum (Spruce) B. S. G. subgenus or section Velutina. In the optimal trees 260 MICHAEL S. IGNATOV & SANNA HUTTUNEN presented here (Figs. 1, 2) Brachytheciastrum is B. microcollinum (E. B. Bartram) comb. nov. resolved as a paraphyletic group, although this – Brachythecium microcollinum E. B. Bartram, result is highly dependent on minor variation in Farlowia 2(3): 318. 1946. alignment of sequence data and hence also on gap B. paradoxum (Hook. et Wils.) comb. nov. cost used in POY analysis as well as treatment of – Hypnum paradoxum Hook. et Wils., London 9 bp inversion region on psbT-N spacer region J. Bot. 3: 554. 1844. (Huttunen & Ignatov, 2003). In the analyses with B. trachypodium (Funck ex Brid.) comb. nov. higher gap costs than one and in those analyses – Isothecium trachypodium Funck ex Brid., where inversion region is omitted or altered, the Bryol. Univ. 2: 766. 1827. three species of Brachytheciastrum involved in B. velutinum (Hedw.) comb. nov. – Hypnum the present analysis appear as a clade. Although velutinum Hedw., Sp. Musc. Frond. 272. 1801. more detailed studies are needed to understand Species referred to the genus based on lit- the relationships between species, the isolated po- erature descriptions [new combinations are not sition of this group with regard to Brachytheci- done, as we did not studied specimens]: um, even in its broadest sense, is clear. Thus, it is Brachythecium appleyardiae McAdam et A. J. with only a little hesitation that we propose to E. Smith (McAdam & Smith, 1981), B. bellicum recognize it as a separate genus. Our decision is Buck, W. R., J. A. Jiménez, R. M. Ros et M. J. also supported by several earlier classifications Cano (Buck & al., 2001), B. bolanderi (Lesq.) where this group has been recognized as an own Jaeg. (Lawton, 1971); B. curvatulum (Broth.) entity (e.g. Brotherus, 1925, Ignatov, 1998). Par. (Gangulee, 1978; Vohra, 1983); B. fendleri (b) The close relationship of the Brachyth- (Sull.) Jaeg. (McFarland, 1994), B. occidentale ecium velutinum group to Homalothecium has (Hampe) Jaeg. (McFarland, 1994), B. percur- never been suggested earlier. These taxa share rens Hedenäs (Hedenäs, 1992), B. praelongum only a few character states, which are, however, Schimp. ex C. Müll. (McFarland, 1988), B. sap- not present in all species: plants often quite rich porense (Besch.) Takaki (Takaki, 1955b). golden, cells of axillary hairs uniformly short, Species of Brachytheciastrum which status leaves rather rigid, margin sharply serrulate al- needs to be confirmed [new combinations are most to the base, costa often ending in spine and not done, as these taxa might be proofed to be sometimes toothed near its end; laminal cells just a synonyms]: narrowly linear, alar cells thick-walled, subquad- Brachythecium leiopodium Broth. (~ falcatulum) rate, in subquadrate group. B. olympicum Jur. (~ velutinum) (c) Most species of Brachytheciastrum have B. venustum De Not. (~ velutinum) small plants, but a few Central Asian and Hi- malayan species (e. g., B. falcatulum and B. kash- 3. Eurhynchiastrum gen. nov. Type: E. pul- mirense) are of about the same size as Brachythe- chellum (Hedw.) Ignatov et Huttunen (= Hyp- cium salebrosum. num pulchellum Hedw.). Species included: Plantae minutae vel mediocres. Folia cauli- B. collinum (Schleich. ex C. Müll.) comb. na ovata-triangulata vel ovata-lanceolata, mar- nov. – Hypnum collinum Schleich. ex C. Müll., gine serrata vel subintegra; costa ultra medium Syn. Musc. Frond. 2: 429. 1851. evanida; cellulae lineares, cellulae alares sub- B. dieckei (Roell) comb. nov. – Brachytheci- quadrates, crassiuscules; folia ramulina ovata- um dieckei Roell, Hedwigia 36 Beibl. 2: 41. 1897. lanceolata, obtusa. Phyllodioica vel dioica. Seta B. falcatulum (Broth.) comb. nov. – Hyp- laevissima. Capsula horizontalis. Peristomium num falcatulum Broth., Acta Soc. Sci. Fenn.: perfectum. Operculum rostratum. 24(7): 34. 1899. Plants small to medium-sized, green to brown- B. kashmirense (Broth.) comb. nov. – Hyp- ish-green. Stem prostrate, with central strand, num kashmirense Broth., Acta Soc. Sci. Fenn.: densely or loosely foliate, irregularly densely 24(2): 37. 1898. branched, branches straight to often curved, dense- B. leibergii (Grout) comb. nov. – Brachyth- ly julaceously foliate. Axillary hairs 3(-4)-celled. ecium leibergii Grout, Mem. Torrey Bot. Club 6: Pseudoparaphyllia acute. Stem leaves erect to 197. 1897. erecto-patent, ovate-triangular to ovate-lanceolate, Brachytheciaceae (Bryophyta) – a family of sibling genera 261

LEAF INSERTION

2

100 mm

1

LEAF BASE PSEUDOPARAPHYLLIA

BRANCH INITIAL LEAF BASE LEAF DECURRENCY

4 Fig. 4. Kindbergia praelonga (Hedw.) Ochyra (from British Columbia, L. & T. Ahti 15159, H): 1 – Front view on stem under leaf insertion: series of shorter cells are low paraphyllia; 2 – transverse section of stem with paraphylla; 3 – semilateral view on stem, showing very low, 1-2 cell high (under leaf insertion) and more tall, 3-4 cell high paraphyllium close to place, where leaf sharply turns to its decurrent part; 4 – semilateral view on stem, showing low paraphyllium between the place, where leaf sharply turns to its decurrent part and group of narrowly triangular 3 pseudoparaphyllia around branch initial. Scale bar same for 1-4. acuminate, not plicate; margin serrulate to sub- chellum in the Homalothecioideae clade strongly entire, costa ending above mid-leaf in spine; lam- supports excluding it from Eurhynchium, a inal cells linear, moderately thick-walled; alar genus based on E. striatum, a member of the cells isodiametric, thick-walled, forming a more Rhynchostegioideae clade. Although our data or less well delimited subquadrate group. Branch offer little support for either Eurhynchiastrum leaves blunt, and smaller and narrower than stem or Brachytheciastrum, these two genera are quite leaves; costa with prominent spine. Dioicous or distinct morphologically. Among others, the phyllodioicous. Perichaetial leaves with reflexed former has smooth seta (rough in most species acumina. Seta smooth. Capsule inclined to hori- of Brachytheciastrum), and rostrate operculum zontal. Annulus separating by fragments. Oper- (conic in Brachytheciastrum), etc. We consider culum rostrate. Peristome xerocastique, perfect. that such differentiation warrants taxonomic Spores small, 17-23 mm. Calyptra naked. Xerome- recognition, and therefor establish the new ge- sophytic or mesophytic. nus Eurhynchiastrum. Distribution: Holarctic and Central America. (b) In the present treatment Eurhynchiella (a) The nested position of Eurhynchium pul- (South Africa and probably also South Ameri- 262 MICHAEL S. IGNATOV & SANNA HUTTUNEN ca; not included in our molecular studies), was longly, broadly decurrent, rarely slightly so; tentatively put in Rhynchostegielloideae (Igna- margin serrate above, serrulate below; costa end- tov & al., 1999). However this genus is similar ing in acumen, sometimes subpercurrent, with or with Eurhynchiastrum in appearance, size, and without spine; laminal cells elongate, moderate- smooth seta (so it can not be excluded, that in ly thick-walled, sometimes forming tooth-like future studies its position will be reconsidered). projections on dorsal surface; subalar cells iso- Species included: diametric, relatively large and thin-walled, form- E. pulchellum (Hedw.) comb. nov. – Hyp- ing indistinctly delimited group. Branch leaves num pulchellum Hedw., Sp. Musc. Frond. 265. smaller, narrower, acuminate, with more strongly 68: f. 1-4. 1801. serrate margin and back of costa and more toothed dorsal lamina, than in stem leaves. Dio- Species of Eurhynchiastrum, which status has icous. Perichaetial leaves with reflexed acumina. to be confirmed: Seta rough. Capsule inclined to horizontal. An- Eurhynchium yezoanum Okam. (~ E. pulch- nulus separating by fragments. Operculum ros- ellum). trate. Peristome xerocastique, perfect. Spores small SUBFAM. BRACHYTHECIOIDEAE or more rarely large. Calyptra naked. Mesophyt- Type species: Brachythecium B. S. G. ic to hygromesophytic. Plants rigid to soft in texture, rich green, Distribution: Holarctic, Africa and South yellowish to very pale stramineous. Leaf mar- America. gin toothed to base or not; costa ending in spine (a) The species of this genus have typically or without spine; laminal cells ovate, hexago- broadly decurrent leaves. There are, however, two nal, or elongate to linear, thin- or thick-walled, exceptions: Kindbergia altaica and K. squarri- often strongly tooth-like prorate on upper dor- folia, both known from a single collection well sal lamina (in Bryhnia and Kindbergia); alar outside the range on K. praelonga, in Altai and cells small, subquadrate, relatively thick-walled, Japan respectively. or large, inflated, thin-walled, in subquadrate or (b) Kindbergia has many similarities with elongate group. Autoicous, dioicous, rarely phyl- Bryhnia and Eurhynchiadelphus. These genera lodioicous. Seta rough or smooth. Operculum have sometimes strongly prorate laminal cells, conic to rostrate. Peristome perfect to modified forming dorsal tooth-like projections. Robinson (in Unclejackia). Hygromesophytic, mesophytic, (1962) considered Kindbergia species within or xeromesophytic. Bryhnia, before he separated them in their own Genera included: Kindbergia, Eurhynchia- genus (Robinson, 1967). delphus, Bryhnia, Brachythecium, Sciurohypnum, (c) Stem on Kindbergia has longitudinal Unclejackia. ridges, which we call paraphyllia. They are low- Genera included with questionmark: Myuro- er that in other genera, where they are taller clada, Scleropodium, Brachytheciella. and therefore look “foliose”. However other- 4. Kindbergia Ochyra, Lindbergia 8: 53. 1982. wise they seems to be similar, at least to what Type: Hypnum stokesii Turn. = K. praelonga we call paraphyllia in Palustriella, Leskea and (Hedw.) Ochyra. similar groups. Stokesiella H. Robinson, Bryologist 70: 39. Species included: 1967, nom. illeg., later homonym. K. africanua (Herz.) Ochyra Plants medium-sized to robust, green to K. altaica (Ignatov) comb. nov. – Eurhynch- brownish-green. Stem prostrate, ascending to erect, ium altaicum Ignatov, Arctoa 7: 141. 37-38. 1998. with central strand, usually very loosely foliate, K. arbuscula (Broth.) Ochyra quite regularly pinnately branched, branches K. dumosa (Mitt.) comb. nov. – Hypnum du- straight, densely or moderately densely, julaceous- mosum Mitt., J. Linn. Soc. Bot. Suppl. 1: 80. 1859. ly foliate. Axillary hairs 4-5-celled. Pseudopara- K. kenyae (Dix. ex Tosco & Piovano) O’Shea phyllia acuminate. Paraphyllia sometimes present et Ochyra (see Fig. 4 and discussion below). Stem leaves K. oedogonium (C. Müll.) Ochyra erecto-patent to squarrose, broadly triangular and K. oregana (Sull.) Ochyra tapered to lanceolate acumen, at base cordate and K. praelonga (Hedw.) Ochyra Brachytheciaceae (Bryophyta) – a family of sibling genera 263

K. squarrifolia (Broth. ex Iishiba) comb. nov. In Central China plants referred to this – Eurhynchium squarrifolium Broth. ex Iishi- genus have narrower and more distant leaves ba, Bot. Mag. (Tokyo) 49: 602. 1935. (cf. Ignatov & al., 2003), but since these pheno- types occur also in North China and Japan, 5. Bryhnia Kaur., Bot. Not. 1892: 61. 1892. forming a continuous transition to the typical Type: Bryhnia scabrida (Lindb.) Kaur. (= Hyp- B. novae-angliae phenotype, we considered them num scabridum Lindb.). conspecific with the latter (Koponen & al, 2000). Plants small to rather robust, green, yellow- The present investigation showed that the Chi- ish to brownish-green. Stem prostrate, with cen- nese plants are very different from European tral strand, unevenly foliate, alternating cochle- and Siberian material. Is then American B. no- ariform parts with moderately loose foliage or vae-angliae closer to European material (de- moderately dense with very loose, irregularly to scribed as B. scabrida) or to Chinese popula- regularly pinnately branched, branches straight, tion? In the latter case, our data suggest that moderately densely, subcomplanately foliate. American “Bryhnia novae-angliae” might be not Axillary hairs 3-4-celled. Pseudoparaphyllia acute. even congeneric with the European B. scabrida. Stem leaves imbricate-appressed, erect to patent, Very preliminary comparisons of our Bryhnia suborbicular or broadly triangular and broadly trnL-F sequences and American one available acute to ovate-lanceolate and then tapered to in Genebank (sequence AF143029/AF161122) long acumen, decurrent; margin serrate above, show that this is not unlikely. Further studies serrulate below; costa ending in upper part of are needed. However, the above generic diagno- leaf, often with spine; laminal cells elongate- sis is retained here for Bryhnia in its tradition- flexuose, moderately thick-walled, sometimes al sense; Siberian and European populations, forming tooth-like projections on dorsal surface; used in the present study, were described and alar cells (or sometimes only subalar cells) iso- illustrated by Ignatov (1998) and Popov & al. diametric, rather large and thin-walled, forming (2000). The Siberian material used for molecu- indistinctly delimited group. Branch leaves nar- lar studies was chosen to be as similar to Amer- rower and smaller than stem leaves (sometimes ican specimens as possible, with rather numer- strongly and abruptly so), acute to acuminate ous branches and strongly prorate laminal cells. (sharper than stem leaves), with margin and back of costa more strongly serrate and dorsal lamina 6. Eurhynchiadelphus Ignatov, Huttunen et more toothed. Dioicous. Perichaetial leaves with T. Kop., gen. nov. Type: E. eustegia (Besch.) reflexed acumina. Seta rough. Capsule inclined Ignatov et Huttunen (= Brachythecium euste- to horizontal. Annulus separating by fragments. gium Besch.). Operculum high-conic, tapered to broad beak. Plantae mediocres vel robustae. Folia vari- Peristome xerocastique, perfect. Spores small or abiles, oblongo-lanceolatae, ovato-triangulatae vel more rarely large. Calyptra naked to hairy. Me- suborbiculatae; cellulae folii medianae breves, sophytic to hygromesophytic. cellulae alares subquadrates. Phyllodioica vel Distribution: Holarctic. dioica. Seta laevissima. Capsula horizontelis. Peris- Species included: tomium perfectum. Operculum lato rostratum. Bryhnia scabrida (Lindb.) Kaurin. Plants rather robust, yellowish-green, not or slightly glossy. Stem creeping, with central The circumscription of this genus needs a com- strand, rather densely foliate, foliage of proxi- plete re-evaluation with DNA markers. The mor- mal and distal parts of shoots sometimes dif- phology seems to be especially misleading in this ferent; remotely pinnate-branched. Axillary hairs genus. In Japan and its adjacent areas, Takaki 3–4-celled, 1–2 basal cells colored. Pseudopara- (1956) first split the genus into 15 species, but phyllia triangular. Stem leaves erect to spread- only 6 of them survived the revision by Noguchi ing, slightly to strongly concave, broadly ob- (1991), all others were synonymized, mostly with long-lanceolate to ovate, acuminate or acute, B. novae-angliae. Of the remaining ones, three spe- twisted at apex, broadly and shortly decur- cies show transitions to B. novae-angliae s. l., where- rent; margin serrulate to serrate throughout; as the two last ones are quite distinct. However, costa ending above mid-leaf, with spine; lami- their position in this genus is not obvious. nal cells irregularly elongate-polygonal, mod- 264 MICHAEL S. IGNATOV & SANNA HUTTUNEN erately thick-walled, sometimes forming tooth- lar. Axillary hairs 2-4-celled. Pseudoparaphyl- like projections on dorsal surface of upper leaf, lia orbicular-triangular. Stem leaves tightly towards base shorter and wider; alar cells sub- imbricate, orbicular or broadly ovate, very broad- quadrate to rectangular, with incrassate cell walls, ly acute and shortly apiculate, shortly decur- forming a moderately delimited opaque group; rent, strongly concave; margin serrate except subalar cells sometimes enlarged and thin- near base; costa ending in upper half of leaf walled. Branch leaves slightly smaller and rel- without spine; laminal cells short rhombic to atively narrower than stem leaves, otherwise rhombic, moderately thick-walled, towards base undifferentiated. Proximal (fairly developed) wider and longer; alar cells subquadrate, form- branch leaves similar to distal or sometimes ing indistinctly delimited area extending up differentiated and then short triangular, round- along margin; subalar cells rather large and ed above, very concave, forming cochleariform thin-walled, forming indistinctly delimited zones in both male and female shoots. Phyllo- group. Branch leaves similar to stem leaves. dioicous or dioicous. Perichaetial leaves with Dioicous. Perichaetial leaves with reflexed long, reflexed acumina. Seta long, smooth. Cap- acumina. Seta smooth. Capsule inclined to hor- sules inclined to horizontal. Annulus separat- izontal. Annulus separating by fragments. Oper- ing by fragments. Operculum conic with broad, culum high-conic, gradually tapered to broad distinctly or indistinctly delimited beak. Peris- beak. Peristome xerocastique, perfect. Spores tome xerocastique, perfect. Spores small. Calyp- small. Calyptra naked. Mesophytic. tra naked. Mesophytic. Distribution: East Asia and North America Distribution: East Asia. (introduced in Europe). (a) This East Asian species was transferred (a) Bescherelle (1893) considered this spe- many times from Brachythecium (due to game- cies to be close to Scleropodium, due to its co- tophytic characters) to Eurhynchium (due to chleariform foliage, but Robinson (1962) found beaked operculum) and back. Molecular data it to be close to Bryhnia. Myuroclada differs nicely confirmed the conclusion of a close rela- from Bryhnia in having smooth seta, and in- tionship between this genus and Bryhnia, which variably suborbicular leaves. Another genus, was originally suggested based on morpholo- closely related to Bryhnia and also having gy (Ignatov & al., 2003). Especially important smooth seta is Eurhynchiadelphus. The latter, for the latter conclusion was the finding of enor- however, strongly differs from Myuroclada in mous variation in leaf shape along individual leaf characters and branching pattern. shoots in some Chinese populations. (b) Analysis of chloroplast sequence data (b) For discussion and illustration see Taka- alone (Huttunen & Ignatov, 2003) places ki (1956) and Ignatov & al. (2003). Myuroclada in the same clade with Bryhnia Species included: and Eurhynchiadelphus. However, the inclu- Eurhynchiadelphus eustegia (Besch.) comb. sion of nuclear ITS2 data and morphological nov. — Brachythecium eustegium Besch., Ann. data resolve Myuroclada quite far from this Sci. Nat., sér. 7, 17: 375. 1893. position (Figs. 1, 2). The situation needs fur- ther investigation. Temporarily we keep 7. Myuroclada Besch., Ann. Sci. Nat., Bot., Myuroclada close to Bryhnia and Eurhynchia- ser. 7, 17: 379. 1893. Type: Myuroclada concin- delphus. na (Wils.) Besch. (= Hypnum concinnum Wils.) Species included: = M. maximowiczii (Borszcz.) Steere et Schof. M. maximowiczii (Borszcz.) Steere et Schof. Achrolepis (Lindb.) Card., Beih. Bot. Cen- tralbl. 17: 34. 1904, nom. illeg., type of earlier Species excluded: M. rotundifolia (H. Arnell) name included. – Myurella subgen. Achrolepis A. L. Abramova et I. I. Abramov – ~ Myrinia. Lindb., Act. Soc. Sci. Fenn. 275, 276. 1872. Type: 8. Scleropodium B. S. G., Bryol. Eur. 6: 27. Hypnum concinnum Wils. 1953. Type: Scleropodium illecebrum B. S. G. Plants rather robust, green to brownish-green. = Scleropodium touretii (Brid.) L. F. Koch. Stem prostrate to ascending, especially in dense Plants large to medium-sized, rigid, green to growth, with central strand, julaceous, irregu- yellowish or brownish. Stem prostrate, with cen- larly branched, branches straight, foliage simi- Brachytheciaceae (Bryophyta) – a family of sibling genera 265 tral strand, densely foliate to cochleariform, ir- masinii Sendtn. ex Boulay), selected here. Com- regularly and mostly sympodially branched. ment: four species were included in the original Axillary hairs 2-4-celled. Pseudoparaphyllia tri- description: Eurhynchium flotowianum (sub ve- angular. Stem leaves ovate to ovate-lanceolate, lutinoides), E. tommasinii, E. cirrosum, E. crassin- obtuse, acute to shortly acuminate, strongly con- ervium. No one of them has obvious advantages cave; margin serrulate; costa ending in upper for to be selected as the generitype. We choose half of leaf, laminal cells unistratose, flexuose- one that has a higher probability (very low, how- linear, thick-walled; basal cells very thick- ever) for separation of a genus of its own. walled, making leaf difficult to detach, alar cells Chamberlainia Grout, Moss. Fl. N. Am. 3: 27. isodiametric, larger, thin- to thick-walled. Branch 1928. Lectotype: C. acuminata (Hedw.) Grout leaves similar to stem leaves. Dioicous. Pericha- (= Leskea acuminata Hedw.), selected by Rob- etial leaves with reflexed acumina. Seta strongly inson (1962). rough, rarely almost smooth. Capsule inclined Plants small to large, green to yellowish green to horizontal. Annulus separating by fragments. and often pale stramineous with age. Stem pros- Operculum conic. Peristome xerocastique, per- trate to ascending, with central strand, densely, fect. Spores small. Calyptra naked. Hygrophyt- rarely more loosely, julaceously to rarely com- ic to xeromesophytic. planately foliate, irregularly or at a certain dis- Distribution: Holarctic and maybe Australia. tance regularly pinnately branched, branches (a) The present analysis (Figs. 2) supports a usually straight, densely or loosely, julaceous or close relationship of Scleropodium obtusifoli- rarely complanately foliate. Axillary hairs 2-3- um to the Brachythecioideae and Homalothe- celled, uppermost cell relatively long and often cioideae, but more detailed studies are needed gradually tapered to distal end. Pseudopara- to confirm this hypothesis. phyllia acute to acuminate. Stem leaves erect, Species included: imbricate, patent or falcate-secund, lanceolate to S. obtusifolium (Mitt.) Kindb. broadly ovate, acuminate or acute, deeply longi- S. touretii (Brid.) L. F. Koch tudinally plicate to smooth; margin serrate to subentire, costa ending above mid-leaf to percur- Species referred to the genus by Hedenäs (1996): rent, without spine or sometimes with; laminal S. australe Heden s ä cells elongate to linear, thin to moderately thick- Species referred to the genus by Lawton (1967): walled; alar cells subquadrate to shortly rectan- S. californicum (Lesq.) Kindb., S. cespitans gular, large thin-walled or small thick-walled, (Wils. ex C. Müll.) L. F. Koch, S. julaceum Lawt. distinctly or indistinctly delimited, alar group These four latter species have never been quadrate or extending up along margin or some- placed outside the genus and morphologically times only subalar cells differentiated. Branch are similar to the type species. However surpris- leaves narrower, with margin and back of costa es from Platyhypnidium (see below) call for more strongly serrate, usually smaller than stem more detailed studies using molecular data (one leaves. Autoicous, dioicous, more rarely polyga- or two species are subaquatic; seta strongly mous. Perichaetial leaves with reflexed acumi- rough to smooth, or in some species unknown). na. Seta long, rough or smooth. Capsule inclined Problematic species: to horizontal, more rarely erect. Annulus separat- S. coreense Card. and S. brachyphyllum Card. ing by fragments, not separating, or totally in- – We were unable to locate types or any material discernible. Operculum conic, very rarely rostrate. of two latter species in PC, BM, H, NICH, KYO. Peristome xerocastique, perfect, very rarely hy- 9. Brachythecium B. S. G., Bryol. Eur. 6: 5. grocastique. Spores small. Calyptra naked, very 1853. Type: B. rivulare B. S. G. (selected by rarely hairy. Hygrophytic to xeromesophytic. Robinson, 1962). Distribution: subcosmopolite. Paramyurium (Limpr.) Warnst., Krypt. Fl. (a) General classification of Brachythecium: Brandenburg 2: 763. 1905. – Eurhynchium sub- The genus Brachythecium, in a traditional sense, gen. Paramyurium Limpr. 1896, Laubm. Deutsch. was split only once by Robinson (1962), who 3: 173. 1896. Lectotype: Eurhynchium tomassi- sorted the species around two core taxa: B. riv- nii (Sendtn. ex Boulay) Mol. (= Hypnum tom- ulare (alar cells large, seta rough, leaves straight) 266 MICHAEL S. IGNATOV & SANNA HUTTUNEN and B. salebrosum (alar cells small, seta smooth, taxon concept, the nested position of Uncle- leaves often falcate). The latter group was seg- jackia in Brachythecium requires that either regated as the genus Chamberlainia. This ge- Unclejackia is synonymized with Brachyth- nus was first established for C. acuminata and ecium, or that Brachythecium is divided into two more species, which were found conspecific several smaller units. The second alternative with it, but later much expanded by Robinson seems inappropriate, because in this case dis- (1962). The distinction between the two groups, tinct phenotypes of what we now call Brachyth- however, appeared fairly unclear when all the ecium salebrosum would be in different gen- species are taken into consideration, hence later era. The inclusion of Unclejackia in Brachy- Robinson (1987) refrained from this idea. Al- thecium makes the morphological definition of though, as we believe, the separation of Sciuro- the latter genus, however, extremely broad. Un- hypnum and Brachytheciastrum makes clejackia is a large golden-lustrous moss with Brachythecium more natural, it still remains strongly concave leaves, a very flexuose leaf acu- quite heterogeneous. men, with very short (0.1-0.2 of leaf length) to Since the sequence variation is not very high absent costa, and a strongly modified peristome; within the Brachythecium, many of the clades it occurs in the subalpine savannah in New have no support (Fig. 2). The topology based Guinean mountains (at about 3500 m elev.), on POY analyses (Fig. 1) is better resolved where it grows always on trunks of the tree fern than strict consensus from Nona analyses. But Cyathaea. The placement of Unclejackia in the since a similar topology to Fig. 1 was obtained Brachytheciaceae has been disputed both in per- from POY analysis despite of the gap cost, we sonal correspondence and by Tan (2000) who believe, that the principal grouping of species believed that Unclejackia is better to be placed in Fig. 1 is more or less reliable. with Chaetomitrium in a separate family. Brachythecium rivulare and related species Based on morphological, molecular and phy- with rough seta are basal in the Brachythecium togeographic data, the origin of Unclejackia is clade. The more advanced species are more B. more or less clear. Its closest relative is probably salebrosum-like, mostly with smooth seta, but Brachythecium lamprocarpum, which is similar including also several species with the rough to Unclejackjia in some rare character states, like, seta, which precludes the segregation of Cham- for example, the peculiar “border” of pellucid berlainia (B. cirrosum, B. complanatum, B. ge- cells at the leaf base, etc. (Ignatov & al., 1999). heebii, B. tommasinii). The most apical portion However, this species has a smooth seta, while it of the clade includes southern and tropical spe- is rough is Unclejackia. Brachythecium lampro- cies, such as B. buchananii (East Asia), B. rud- carpum is the only widespread similar species erale [well known as B. stereopoma in tropical of Brachythecium of that area. In our analyses America and as B. implicatum in tropical Afri- (cf. Figs. 1-2) Unclejackia is resolved as closely ca], and B. lamprocarpum (Malesia, Oceania). related to B. lamprocarpum, though with only This group was segregated into a separate sec- low support. Thus, it is more or less clear that tion by McFarland (1994), but since he obvi- Unclejackia is close to the group of southern ously did not validate it, we will call it the B. Brachythecium taxa, and has most likely evolved stereopoma-group sensu McFarland. Brachyth- rapidly from them in a limited area and under ecium laetum also belongs to this group, as it is an unusual environment, which lead to a differ- similar to the tropical species of this group in entiation of several morphological characters the absence of an annulus. This clade includes compared to its sister taxa. Concerning the tax- also B. acuminatum, the type of the genus Cham- onomy, however, we see no better choice than to berlainia. And finally this clade also includes retain Unclejackia in a genus of its own, at least Unclejackia, the curious moss with almost ecos- until the evolution within the Brachythecium- tate leaves, recently transferred from Chaeto- subgroup is resolved in greater detail. It seems mitrium (Symphyodontaceae) to Brachytheci- like the problem with the rank of Unclajackia is aceae (Ignatov & al., 1999). similar to the acceptance of birds as a class of (b) The position of Unclejackia and Brachy- vertebrates, which makes the class of reptiles theciella: In accordance with a monophyletic paraphyletic (cf. for example, Forey, 2002). Brachytheciaceae (Bryophyta) – a family of sibling genera 267

Almost the same discussion can be repeated in Brachythecium. In Asia, however, this combi- for Brachytheciella, another New Guinean en- nation is known in B. coreanum, B. coarctum, and demic genus, for which molecular data were not some phenotypes of B. auriculatum. The latter available in this study. species was incorrectly synonymized with Palam- (c) Basal species with rough seta. The sister ocladium leskeoides by Hofmann (1998), who clade to the rest of the Brachythecium clade oversaw the rough seta in this species. includes Brachythecium rivulare, B. rutabulum, (f) This new circumscription of Brachyth- B. frigidum + South Chinese Bryhnia (cf. Fig. ecium opens an opportunity for the placement 1). The curious position of the latter taxa calls of one peculiar Chinese endemic, B. (Eurhynch- for further extensive studies of this group. This ium) coarctum. This species has medium-sized position of the South Chinese Bryhnia can hardly to large plants, long plicate leaves with a non- be considered an analytical artifact, as despite Homalothecium base, laminal cells with of excluding all the possibilities for contami- length:width ratio ca. 5-8:1, rough seta and a nation, this specimen was never resolved close rostrate operculum. Based on this character state to Bryhnia, but always close to Brachythecium combination we believe that it is close to B. frigidum (in analyses of chloroplast sequence tommasinii. data set alone, and chloroplast plus nuclear set, Species included: (+ – not listed in Crosby and combined molecular and morphological & al., 1999) sets with different gap costs, etc.). B. acuminatum (Hedw.) Aust. (d) Brachythecium tommasinii differs from B. albicans (Hedw.) B. S. G. the rest of Brachythecium in its rostrate oper- B. afroglareosum (Broth.) Par. culum. This offers important new insight of the +B. auriculatum Jaeg. value of this character. Though it is rather sta- B. austroglareosum (C. Müll.) Kindb. ble within two other subfamilies (invariably B. austrosalebrosum (C. Müll.) Kindb. rostrate in Rhynchostegielloideae; almost always B. bellii (Mitt. ex Broth.) Par. rostrate in Rhynchostegioideae, with the only B. buchananii (Hook.) Jaeg. exception of Pseudoscleropodium), it is rather B. campestre (C. Müll.) B. S. G. freely variable in the Brachythecioideae. The +B. coarctum (C. Müll.) comb. nov. – Eu- situation parallel to B. tomassinii was found rhynchium coarctum C. Müll., Nuov. Giorn. Bot. also in Sciurohypnum, where S. flotowianum Ital., n. ser. 5: 198. 1898. has been found close to S. populeum (see be- B. complanatum Broth. low). Both B. tommasinii and Sciurohypnum B. coreanum Card. flotowianum are characterized by a rostrate B. coruscum I. Hag. operculum and short laminal cells. Such com- B. digastrum C. Müll. ex Kindb. bination can also be recognized in the basal B. erythrorrhizon B. S. G. part of the Brachythecium-clade, if Bryhnia and B. frigidum (C. Müll.) Besch. Kindbergia are considered as derivatives from B. garovaglioides C. Müll. a Brachythecium rivulare-like ancestor. B. geheebii Milde (e) Wigh (1973) suggested to transfer B. glareosum (Bruch ex Spruce) B. S. G. Brachythecium geheebii, a rare Western Pale- B. helminthocladum Broth. et Par. arctic species, to Homalothecium, based partly B. kamounense (Harvey) Jaeg. on chromosome numbers. B. geheebii has n=10, a B. kuroishicum Besch. number known (that time!) in Brachythecium +B. laetum (Brid.) B. S. G. only in morphologically very distinct species B. lamprocarpum (C. Müll.) Jaeg. with smooth seta. However, n=10 or 11 is known B. longicuspidatum (Mitt.) Jaeg. in B. tommasinii and the East Asian B. (Camp- B. mildeanum (Schimp.) Schimp. in Milde tothecium) auriculatum and B. coreanum; all +B. papuense Ignatov three species have rough seta (see also Appendix B. pinnirameum C. Müll. 1). Brachythecium geheebii has very deeply pli- B. poadelphus C. Müll. cate leaves in combination with rough seta, which B. procumbens (Mitt.) Jaeg. in Europe is known only in Homalothecium, never B. rivulare B. S. G. 268 MICHAEL S. IGNATOV & SANNA HUTTUNEN

B. rotaeanum De Not. long, reflexed acumina. Seta strongly rough. B. ruderale (Brid.) Buck [= B. stereopoma, Capsule erect to slightly inclined, cylindric. B. implicatum] Annulus not differentiated. Operculum high- B. rutabulum (Hedw.) B. S. G. conic. Peristome xerocastique, modified; exos- B. salebrosum (Web. et Mohr) B. S. G. tome teeth red-brown, gradually tapered above, B. subplicatum (Hampe) Jaeg. papillose above, cross-striolate below; endostome B. spectabile Broth. with basal membrane about 1/4 of endostome +B. tommasinii (Sendtn. ex Boulay) comb. height, segments narrow, narrowly perforated nov.– Hypnum tommasinii Sendtn. ex Boulay, along keel, cilia very short. Spores large. Calyp- Fl. Crypt. Est, Muscin. 225. 1872. tra naked. Mesophytic (on trunks of fern-trees B. turgidum (C. Hartm.) Kindb. in subalpine savannah). B. udum I. Hag. Distribution: New Guinea. B. vellereum (Mitt.) Jaeg. (a) See discussion under Brachythecium, also Ignatov & al. (1999). Species referred to the genus based on lit- Species included: erature descriptions: U. crispifolia (E. B. Bartram) Ignatov, T. Kop. McFarland, 1994a: Brachythecium chocayae et D. Norris Herzog, B. conostomum (Taylor) Jaeg., B. poly- U. longisetula (E. B. Bartram) Ignatov, T. oicum (Thér.) McFarland, B. zanonii Buck. Kop. et D. Norris. Hedenäs, 2002: B. latinervium Hedenäs Buck, 1993: B. gloriosum (C. Müll.) Par. 11. Brachytheciella Ignatov in Ignatov, T. Kop. (3) Species excluded from the genus (see et D. Norris, Acta Bot. Fennica 165: 35. 1999. also all species under Brachytheciastrum and Type: B. stolonifera Ignatov. Sciurohypnum [there are no heterotypic species Plants medium-sized to moderately robust, epithet]): very fragile, golden-green to rich golden. Stem B. koponenii Ignatov – Cirriphyllum creeping, with central strand, julaceously foli- B. weinmannii (Nees) Par. = Callicladium ate, irregularly branched, branches straight, fo- haldanianum (type in LE!). liage similar. Axillary hairs 2-3-celled, upper cells hyaline. Pseudoparaphyllia longly acuminate. 10. Unclejackia Ignatov, T. Kop. et D. Norris, Stem leaves erect-appressed, from ovate, con- Acta Bot. Fennica 165: 37. 1999. Type: Uncle- cave basal part gradually contracted into fili- jackia longisetula (E. B. Bartram) Ignatov, T. form acumen, not decurrent; margin subentire; Kop. et D. Norris (=Chaetomitrium longisetu- costa absent; laminal cells linear, moderately lum E. B. Bartram). thick-walled, towards leaf base wider; alar cells Plants robust, golden-green to rich golden. larger in few rows along margin and this group Stem creeping, with central strand, julaceously extending shortly up along margin. Branch foliate, irregularly branched, branches straight, leaves similar to stem leaves. Dioicous? Per- branch foliage similar to that in stem. Axillary ichaetial leaves erect, acuminate. Seta slightly hairs 2-3-celled, upper cells hyaline. Pseudopara- rough below capsule, otherwise smooth. Capsule phyllia longly acuminate. Stem leaves loosely inclined, ovate-cylindrical, constricted below imbricate, from ovate or elliptic, very concave mouth. Annulus, peristome and calyptra un- basal part abruptly contracted into filiform, known. Operculum high-conic. Spores small (16- moderately to strongly flexuose (in both dry 22 mm). Mesophytic (on trunks of fern-trees in and wet condition) acumen, not decurrent; subalpine savannah). margin serrulate to subentire; costa reaching Distribution: New Guinea. 0.1(0.2) of leaf length or totally absent, ending (a) This genus is similar in many respects smoothly; laminal cells linear, moderately thick- to Unclejackia (see discussion on this genus walled, towards leaf base wider; alar cells larg- under Brachythecium, and also Ignatov & al., er, in few rows along margin, extending shortly 1999). up along margin. Branch leaves smaller than Species included: stem leaves, otherwise similar. Autoicous (or B. stolonifera Ignatov. sometimes dioicous?). Perichaetial leaves with Brachytheciaceae (Bryophyta) – a family of sibling genera 269

12. Sciurohypnum (Hampe) Hampe, Linnaea resented in that list by two species of Palam- 38: 220. 1874. – Hypnum subg. Sciurohypnum ocladium. Hence it is considered as a nomen nu- Hampe, Flora 50: 76. 1867, ‘Sciuro-hypnum’. Lec- dum and not used further. Moreover its concept totype: Hypnum plumosum Hedw. = S. plumo- differs greatly from the subsequent one. The first sum (Hedw.) Ignatov et Huttunen, selected here. publication with a description (also as a subdivi- Chionobryum Glow., Oesterr. Bot. Z. 63: 279. sion of Hypnum, Hampe, 1867), included two spe- 1913, syn. nov. Type: C. venturii (De Not.) Glow. cies, Hypnum plumosum and H. velutinum, which (= Bryum venturii De Not.) = Scuirohypnum were taken as examples, whereas the circumscrip- glaciale (B. S. G.) Ignatov et Huttunen, comb. tion of the group was extended to include all the nov.(fide Loeske in Mönkemeyer, 1927, as Brachythecium and Camptothecium sensu “Bry- Brachythecium glaciale), syn. nov. ologia Europaea”. In subsequent publications Cratoneurella H. Robinson, Bryologist 65: Hampe (1871, 1873) included in this section H. 140. 1962, syn. nov. Type: C. uncinifolia (Broth. campestre, H. lutescens, H. populeum, H. rutabu- et Par.) H. Robinson, syn. nov. (= Brachytheci- lum, H. salebrosum, etc. Later Hampe (1874) de- um uncinifolium Broth. et Par.) scribed a new species from Madagascar, calling it Plants small to large, light to deep green, Sciuro-hypnum borgenii (the only member of becoming brownish or rarely whitish with age. Brachytheciaceae in his publication). Since that Stem prostrate, with central strand, loosely, rare- species was considered a relative of Hypnum al- ly more densely, julaceously or rarely complan- bicans, and also according to the species descrip- ately foliate, irregularly to at places regularly tion, it is clear that the concept of this new genus pinnately branched, branches straight or curved, was simply taken from the concept of subgenus. foliage somewhat denser than that of stem and According to Art. 33.2 of the Saint Louis Code the more often complanate. Axillary hairs 2-3(-5)- generic name must be treated as a transfer from celled, uppermost cell obtuse, rarely indistinctly Hypnum subgen. Sciuro-hypnum 1867. Thus, Sci- tapered to distal end. Pseudoparaphyllia acute. uro-hypnum borgenii Hampe (indicated in In- Stem leaves erecto-patent, patent, reflexed, im- dex Muscorum as the type) is not the correct type bricate, ovate to ovate-lanceolate, acuminate or of the generic name and the lectotype must be acute, indistinctly plicate to smooth; margin ser- chosen from the two elements definitely includ- rate to subentire; costa ending above mid-leaf to ed in the 1867 protologue, namely H. plumosum percurrent, without spine; laminal cells ovate to or H. velutinum. We prefer H. plumosum. The rea- linear, thin- to moderately thick-walled; alar cells son is just etymological. If H. velutinum would isodiametric, large and thin-walled or small and be chosen as a type, then the earliest generic name thick-walled, alar group isodiametric, usually for this group of species would be Chionobryum poorly delimited. Branch leaves narrower than (“snow-moss”). However, the species of the genus stem leaves, with more strongly serrate margins are common in boreal, temperate, and subtropical and costa more often ending in a spine. Auto- forests. In addition, most species of the Brachyth- icous, very rarely dioicous (but some autoicous ecium velutinum group (that should be called species rare with sporophytes). Perichaetial leaves Sciurohypnum in this case) have a rather loose with reflexed acumina. Seta rough or very rare- foliage and are not “sciureous” at all. ly smooth. Capsule inclined to horizontal, more (b) Chionobryum Glow. was described for rarely erect. Annulus separating by fragments, one species, C. venturii (De Not.) Glowacki = not separating, or totally indiscernible. Opercu- Bryum venturii De Not. When describing the lum conic, more rarely rostrate. Peristome xero- latter, De Notaris compared it with Bryum castique, perfect. Spores small. Calyptra naked. funkii (!), but Mönkemeyer (1927) stated that Hygromesophytic to xeromesophytic. according to Loeske the only collection of this Distribution: subcosmopolite. species belongs to Brachythecium glaciale. We Nomenclatural notes: were able to study an isotype: “Auctor cot C. (a) The name Sciurohypnum, as a subdivision Müll. ex auct. De Not., Bryum Venturii De Not., of Hypnum, first appeared in Hampe’s (1852) Epilogo p. 408. Hochalpe Seant in Ratti. August publication without description, with the simple 74. Venturi”. (H-SOL 516001), and confirm listing of species. The Brachytheciaceae were rep- Loeske's opinion. 270 MICHAEL S. IGNATOV & SANNA HUTTUNEN

(c) Cratoneurella was established by Rob- Brachythecium tommasinii (above). inson (1962) based on Brachythecium uncini- Species included: folium, a very rare species from Japan. Japanese S. brotheri (Par.) comb. nov. – Brachyth- authors placed this species next to B. reflexum ecium brotheri Par., Index Bryol. (ed. 2) 2: 139. (Takaki, 1955b; Noguchi, 1991), and Takaki (l. 1904. c.) commented that they are closely related. It S. flotowianum (Sendtn.) comb. nov. – Hyp- seems like the important reason for the separa- num flotowianum Sendtn., Denkschr. Bayer. Bot. tion of this genus was that Robinson consid- Ges. Regensburg 3: 146. 1841, ‘flotovianum’. ered the leaf falcateness as a very important S. glaciale (B. S. G.) comb. nov. – Brachyth- character separating Brachythecium s. str. and ecium glaciale B. S. G., Bryol. Eur. 6: 15. 542. Chamberlainia s. lat. No species of Brachytheci- 1853. um sensu Robinson had falcate leaves, so the S. hylotapetum (N. Higinbotham et B. Hig- inclusion of this species into Brachythecium inbotham) comb. nov. – Brachythecium hylot- would reduce the number of differential char- apetum N. Higinbotham et B. Higinbotham, acters between the two genera seriously (be- Bryologist 61: 339. 1958. cause they are few). Our observations on spe- S. latifolium (Kindb.) comb. nov. – Brachy- cies morphology confirm Takaki's opinion that thecium latifolium Kindb., Forh. Vidensk.-Sel- B. uncinifolium and B. reflexum are very close, sk. Kristiania 1888 (6): 8. 1888. if not conspecific. S. majusculum (Newton) comb. nov. – Systematic notes: Brachythecium majusculum Newton, Brit. Ant- (d) This group existed in most previous treat- arct. Surv. Bull. 39: 45. f. 1. 1974. ments of Brachythecium s. l. as sections Cirri- S. oedipodium (Mitt.) comb. nov. – Hyp- phyllopsis and Reflexa. Its relative homogene- num oedipodium Mitt., J. Proc. Linn. Soc., Bot. ity is easy to see by retrieving keys for the 8: 32. 1865. Brachythecium in different handbooks: contrast, S. ornellanum (Molendo) comb. nov. – Hyp- such as {plants small to medium-sized, leaves num ornellanum Mol., Ber. Naturhist. Vereins plus minus smooth} vs. {plants large, leaves usu- Augsburg 18: 185. 1865. ally plicate} is commonly found at the begin- S. plumosum (Hedw.) comb. nov. – Hyp- ning of long keys to Brachythecium sensu B. S. G. num plumosum Hedw., Sp. Musc. Frond. 257. Large-sized plants are known in Sciurohypnum 1801. only in polyploid S. oedipodium and putative S. populeum (Hedw.) comb. nov. – Hyp- polyploid S. hylotapetum (cf. Appendix 1). Our num populeum Hedw., Sp. Musc. Frond. 270. 70 analysis resolved Sciurohypnum with high jack- f. 1-6. 1801. knife and bremer support (Fig. 2), and actually S. reflexum (Starke) comb. nov. – Hypnum its species form one of the most stable clades in reflexum Starke, Bot. Taschenb. 306. 1807. various analyses of different sets with different S. starkii (Brid.) comb. nov. – Hypnum starkei parameter (Huttunen & Ignatov, 2003). [starkii] Brid., Muscol. Recent. 2(2): 107. 1801. (e) The most unexpected novelty in this Species belonging to Sciurohypnum as de- group is the close relationship of Sciurohyp- duced from McFarland's (1988) descriptions: num (Cirriphyllum, Eurhynchium) flo- Brachythecium filirepens Dusen, B. cirriphyl- towianum to S. (Brachythecium) populeum, loides McFarland. which also got high jackknife and bremer sup- Species of Sciurohypnum which status is to port (Fig. 2). These two species have similar be confirmed: appearances and leaf shapes, and a percurrent Brachythecium uncinifolium Broth. et Par. costa is not rare in S. flotowianum, at least in (~reflexum). the branch leaves. However S. flotowianum B. dovrense (Limpricht) Amann (~glaciale) differs from S. populeum, as well as from all B. nelsonii Grout (~latifolium) other species of the genus, in having rostrate B. uematsui Broth. (~plumosum) operculum and rather short laminal cells. It B. pulchellum Broth. et Par. (~plumosum) seems like these two characters often evolve B. spuriopopuleum (Broth.) Par. (~populeum) together in a parallel way; cf. the case of B. pseudouematsui Noguchi (~plumosum) Brachytheciaceae (Bryophyta) – a family of sibling genera 271

SUBFAM. RHYNCHOSTEGIELLOIDEAE ly concave, plicate or not; margin serrate to ser- SUBFAM. NOV. rulate; costa ending without spine; laminal cells Plantae minutae vel robustae. Folia lato ovata linear, elongate or rhombic-elongate, moderate- vel anguste lineares, acuta, acuminata vel ob- ly thick-walled; alar cells subquadrate, small tusa; costa crassa vel tenuis; cellulae angusto or large, forming isodiametric group or extend- lineares vel brevi ovales. Autoica, dioica vel poly- ing up along margin. Branch leaves rather dif- oica. Seta scabra vel laevissima. Capsula erecta ferent from stem leaves in being narrower, less vel horizontalis. Peristomium perfectum vel abrupt tapering or contracting to acumen, not imperfectum. Operculum rostratum. decurrent, more strongly serrate. Dioicous. Per- Type species: Rhynchostegiella (B. S. G.) Limpr. ichaetial leaves abruptly contracted to long, Plants minute to robust. Leaves broadly ovate reflexed acumina. Seta long, rough. Capsule in- to narrowly linear, acute, acuminate, or obtuse; clined to horizontal, ovate. Annulus separating costa very strong to thin; cells narrowly linear by fragments. Operculum longly rostrate. Peris- to shortly ovate. Autoicous, dioicous, or poly- tome xerocastique, perfect. Spores small. Calyp- oicous (never phyllodioicous). Seta rough or tra naked. Mesophytic. more rarely smooth, long to very short. Capsule Distribution: Holarctic and Malesia. inclined to erect. Peristome perfect to various- (a) Cirriphyllum is one of the most trouble- ly reduced. Operculum rostrate (never conic). some genera to describe based on morphology. This is the largest subfamily in term of num- A change in the cell length to width ratio seems ber of genera, 19 [with some more to be de- to have evolved in parallel with an elongation scribed]; the following groups are well-support- of the rostrum of the operculum in many groups ed within it (? – species attributed to group of Brachytheciaceae. The group close to C. pilifer- based on morphology only): um can be understood as a series from the long- (1) Cirriphyllum + Okamuraea; (2) Oxyr- celled C. piliferum via the moderately short- rhynchium + Donrichardsia + ?Nobregaea + celled C. crassinervium to the short-celled ?Eurhynchiella; (3) Rhynchostegiella (Euro- Okamuraea. Interestingly, the axillary hairs in pean and partly African taxa); (4) Clasmat- these two genera have usually many short cells odon + Helicodontium + ?Rhynchostegiella p. proximally. p. (see note “a” under that genus) + Homaloth- Species included: eciella + ?Flabellidium + ?Mandoniella + ?Ju- +C. crassinervium (Tayl.) Loeske et Fleisch. ratzkaeella + ?Schimperella; (5) Aerolindigia (listed Crosby & al., 1999 under Eurhynchi- + Remyella; (6) Meteoridium + Zelometeori- um). um + Squamidium. +Cirriphyllum koponenii (Ignatov) comb. nov. – Brachythecium koponenii Ignatov, Acta 13. Cirriphyllum Grout, Bull. Torrey Bot. Cl. Bot. Fenn. 165: 29. 2. 1999. 25: 222. 1898. Type: C. piliferum (Hedw.) Grout C. piliferum (Hedw.) Grout (= Hypnum piliferum Hedw.). Plants large to medium-sized, slender to Species transferred to other genera: moderately rigid, light green to golden or brown- C. cirrosum (Schwägr.) Grout – Brachythecium ish green. Stem prostrate to ascending, with cen- C. flotowianum (Sendtn.) Ochyra – Sci- tral strand, densely to moderately densely, jula- urohypnum ceously foliate, irregularly to rather regularly C. germanicum (Greb.) Loeske et Fleisch. pinnately branched, branch foliage similar to – not a Brachytheciaceae that in stem. Axillary hairs 3-5-celled, often with C. tommasinii (Sendtn. ex Boulay) Grout – 3 short lower cells. Pseudoparaphyllia orbicu- Brachythecium. lar-triangular to acuminate. Stem leaves loose- 14. Okamuraea Broth., Oefvers. Foerh. Fins- ly to closely imbricate to erecto-patent, from ka Vetensk.-Soc. 49: 2. 1906. Type: O. cristata ovate or elliptic base more or less abruptly con- Broth. = O. hakoniensis (Mitt.) Broth. tracted into long to short, filiform or lanceolate Plants medium-sized, rather rigid, yellowish acuminate, or gradually acuminate apex, longly green. Stem creeping, with central strand and or moderately decurrent, moderately to strong- 4-6-stratose sclerodermis, densely to moderate- 272 MICHAEL S. IGNATOV & SANNA HUTTUNEN ly densely foliate, irregularly pinnately branched, Noguchi's (1953, 1991) publications include branch foliage similar to that in stem, some- all known species of the genus. times branches attenuate. Axillary hairs 3-6- Species included: celled, blunt, composed of short cells through- O. brachydictyon (Card.) Noguchi out or at least in the lower part. Pseudopara- O. brevipes Broth. ex S. Okamura phyllia triangular. Stem leaves erect to erect O. hakoniensis (Mitt.) Broth. appressed when dry, spreading when wet, ovate O. plicata Card. to ovate-lanceolate, longly acuminate, not de- 15. Oxyrrhynchium (B. S. G.) Warnst., Krypt. current, more or less concave, plicate or not pli- Fl. Brandenburg 2: 781. 1905, nom. cons. prop. – cate; margin serrulate; costa ending above mid- Eurhynchium subgen. Oxyrrhynchium B. S. G., leaf, without spine; laminal cells rhombic-ovate, Bryol. Eur. 5: 224. 1854. Type: Eurhynchium hians very thick-walled; at leaf base cells shorter, alar (Hedw.) Sande Lac. (= Hypnum hians Hedw., cells subquadrate, moderately thick-walled, form- Oxyrrhynchium hians (Hedw.) Loeske), typ. cons. ing extensive opaque area extending up along prop. (Ignatov & Isoviita, 2003). margin. Branch leaves similar to stem leaves. Plants medium-sized, moderately rigid, green Dioicous. Perichaetial leaves straight, abruptly to brownish green. Stem creeping, with central longly acuminate, costate, costa reaching 1/2– strand, very loosely or at places more densely 1/4 of leaf length. Seta long, smooth. Capsule julaceously foliate, at places rather regularly suberect to inclined, shortly ovate to cylindri- branched, branch foliage julaceous, subcomplan- cal. Annulus not differentiated. Operculum ate to sometimes distinctly complanate. Axil- moderately to longly rostrate. Peristome hygro- lary hairs 2-3-celled, often 2-celled with upper castique, strongly modified; exostome red-brown, cell vesiculate inflated, hyaline or pale brown- teeth massive, with numerous dense trabeculae, ish. Pseudoparaphyllia triangular, acute. Stem on the outer surface with small papillae to the leaves erecto-patent to patent, ovate, longly or base, inner surface not papillose; endostome with shortly acuminate, slightly concave, smooth; basal membrane to about 1/3 of tooth length, margin serrate to serrulate; costa ending in but without segments and cilia. Spores large. spine; laminal cells elongate to linear, moder- Calyptra hairy. Brood branches in clusters in ately thick-walled; alar cells weakly differen- axils of upper leaves present in one species. Me- tiated (sometimes only subalar cells differen- sophytic to xeromesophytic. tiated), short rectangular, thin-walled, forming Distribution: East Asia. an indistinctly delimited group. Branch leaves (a) In the original description Okamuraea different from stem leaves in being more ellip- was compared with Forsstroemia, Cryphaeace- tic in overall shape, shortly acuminate to acute, ae. Later, in 1908, Brotherus (1905-1909) placed asymmetric at base, twisted in mid-leaf and it in the Brachytheciaceae, Fleischer (1923) and more strongly serrate and with costa sometimes Brotherus (1925) in the Rhytidiaceae, Noguchi toothed distally, ending in a more stout spine. (1953) in the Brachytheciaceae, and Iwatsuki Dioicous or more rarely synoicous or polyoicous. & Noguchi (1973) and Noguchi (1991) in the Perichaetial leaves abruptly contracted to long, Leskeaceae. In almost all publications of the reflexed acumina. Paraphyses often conspicuous- last decades of the XXth Century it was placed ly exserted. Seta long, rough, more rarely smooth. in the latter family. Based on molecular data, Capsule inclined to horizontal, ovate. Annulus Tsubota & al. (2002) placed it in the Brachyth- separating by fragments. Operculum longly ros- eciaceae. Studies of the pseudoparaphyllia sup- trate. Peristome xerocastique, perfect. Spores port this, as well as the results of our analysis small. Calyptra naked. Mesophytic. of combined molecular and morphological data. Distribution: subcosmopolite. (b) In our analysis, Okamuraea was found (a) This genus includes most of the species to be closely related to Cirriphyllum. Note, formerly placed in Eurhynchium. They have that in C. crassinervium the cells are consider- smooth leaves, often twisted in the middle, ably shorter than in C. piliferum, and it can branch leaves tending to be elliptic; axillary hence be interpreted as a transitional morpho- hairs short, 2-3-celled, and in many species up- type (Figs. 1-2). per cells much broadened to almost vesiculate. Brachytheciaceae (Bryophyta) – a family of sibling genera 273

Most species of the genus have a rough seta, but this exceptional morphology. An alternative O. vagans, widely distributed from Himalayas hypothesis is that the overall similarity of O. to Malesia, and several South Hemispheran speciosum and other Oxyrrhynchium species species have a smooth seta. The very loose foli- is superficial. This dilemma needs a more de- age with remote, subcomplanate leaves often tailed study. Note that chromosome number in allow immediate recognition of the genus. How- O. speciosum is n=15 (versus n=7, 8, 10 in other ever, almost all species become subdendroid in taxa, cf. Appendix 1), suggesting that it is of a dense growth, then lacking the complanate foli- polyploid (e.g., alloploid) origin. age as leaves tend to be more rigid and subcar- 16. Donrichardsia Crum et Anderson, Field- inate. This polymorphism causes problems with iana, Bot., n. s. 1: 7. 1979. Type: D. macroneuron the identification of species, because without spe- (Grout) Crum et Anderson (= Hygroamblyste- cial experience it is difficult to believe that these gium macroneuron Grout). contrasting phenotypes belong to the same spe- Plants robust, rigid, in dense mats, dark-green cies. The taxonomy in regions where 2-3 species to brownish-green. Stem prostrate, occasionally of Oxyrrhynchium occur (Europe: hians-schle- curved at end, central strand present, cortical icheri), China (hians-savatieri-vagans) is espe- cells in (1–)2 rows of small, thick-walled cells, cially troublesome. densely to moderately densely foliate, foliage (b) Oxyrrhynchium was found monophyletic julaceous to homomallous, irregularly loosely with moderate support in Nona analysis (Fig. 2), branched, branches prostrate to ascending, rath- while POY analysis resolved it as paraphyletic er densely imbricately foliate, subjulaceous to group with Donrichardsia nested within it (Fig. 1). homomallous. Axillary hairs 2-4-celled, colored Species included: throughout. Pseudoparaphyllia orbicular-trian- O. asperisetum (C. Müll.) Broth. gular. Stem leaves erect to appressed, suborbic- O. clinocarpum (Taylor) Broth. ular to ovate-lanceolate, broadly acute, some- O. hians (Hedw.) Loeske times abruptly cuspidate or shortly apiculate; Oxyrrhynchium laevisetum (Geheeb) comb. strongly concave, shallowly longitudinally pli- nov. – Eurhynchium laevisetum Geheeb, Rev. cate; margin serrulate to serrate throughout with Bryol. 3: 4. 1876. small acute or blunt teeth; costa strong, shortly O. protractum (C. Müll.) Broth. excurrent, percurrent, or ending at (0.7–)0.8– O. pumilum (Wils.) Loeske 0.95 of leaf length; laminal cells unistratose or O. remotifolium (Grev.) Broth. partly 2-4-stratose, elongate to linear, moder- O. rugosipes (Besch.) Broth. ately thick-walled; basal cells shorter, thick- O. savatieri (Schimp. ex Besch.) Broth. walled, alar cells not differentiated, but subalar O. schleicheri (Hedw. f.) Roell cells often enlarged and forming an indistinct Oxyrrhynchium trichocladoides (Ignatov) group. Branch leaves erect to patent, differing comb. nov. – E. trichocladoides Ignatov, Acta from stem leaves in often being slightly asym- Bot. Fennica 165: 58. 1999. metric, slightly longer and narrower, and in Oxyrrhynchium vagans (Jaeg.) comb. nov. having somewhat longer laminal cells. Dioicous? – Rhynchostegium vagans Jaeg., Ber. Thätigk. Perichaetial leaves with slightly reflexed acumi- St. Gallischen. Naturwiss. Ges. 1876-77: 369. 1878. na. Seta long, rough. Capsule inclined to hori- (Ad. 2: 435). zontal. Operculum rostrate. Peristome xerocas- Problematic species: tique, perfect. Spores and calyptra unknown. O. speciosum (Brid.) Warnst. – the overall Hydrophytic or hygrophytic. morphology of this species is rather similar to Distribution: desjunctive between East Asia other species of the genus, but it has exceptio- and Eastern North America. nally long (often 5-celled) and brownish axil- (a) Donrichardsia macroneuron, a curious lary hairs, quite different from those of other aquatic species was originally described in Hy- members of the genus. Since many subaquatic groamblystegium, but later placed in Eurhynch- species often have long axillary hairs, one might ium (considering its affinities to Platyhypnidi- hypothesize, that the growth in this environ- um, which was that time included in Eurhynch- ments, exceptional for the genus, is resulted in ium) and then segregated in a monotypic genus 274 MICHAEL S. IGNATOV & SANNA HUTTUNEN of Amblystegiaceae (Crum & Anderson, 1979). (d) Further studies are needed to under- Vanderpoorten & al. (2002) in their analysis of stand the correct position of the species known Ambystegiaceae s. lat. suggested to accommodate as “Platyhypnidium austrinum”. It seems to be Donrichardsia in Platyhypnidium, which is not an another derivative of the Oxyrrhynchium surprising, because their study included only two lineage. genera of the Brachytheciaceae. We appreciate Species included: their decision to put Donrichardsia in the D. macroneuron (Grout) Crum et Anderson. Brachytheciaceae, but according to the present Donrichardsia patulifolia (Card. et Thér.) results it belongs to another subfamily than comb. nov. – Rhynchostegium patulifolium Platyhypnidium. The morphology of Platyhyp- Card. et Thér. in Thér., Bull. Acad. Int. Géogr. nidium seems to be no more than a running- Bot. 21: 272. 1911. – Platyhypnidium patulifo- water habitat syndrome, shared by different lium (Card. et Thér.) Broth., Nat. Pfl. ed. 2, 11: aquatic Brachytheciaceae and Amblystegiaceae 347. 1925. s. lat. (see also comments on Platyhypnidium). 17. Nobregaea Hedenäs, Bryophyt. Biblioth. (b) In our study the closest relative of Donri- 44: 124. 1992. Type: N. latinervis Hedenäs. chardsia macroneuron was found among East Plants small. Stem prostrate, with central Asian species (cf. classical “Liriodendron” disjunc- strand, loosely foliate, irregularly branched. tion, known also in mosses, cf. Iwatsuki, 1958). Axillary hairs 2-4-celled. Pseudoparaphyllia tri- Platyhypnidium patulifolium is an interesting angular. Stem leaves patent, ovate-lanceolate to plant from China originally described from a very lanceolate, narrowly acute; margin serrate; cos- poor collection in the genus Rhynchiostegium, and ta very broad, indistinctly separated from lam- later accepted by some authors in Platyhypnidi- ina; laminal cells at places bistratose, elongate, um. A second collection by Koponen in China rather irregular in shape, thick-walled; alar cells allowed for a more thorough description (Igna- rectangular, forming indistinctly delimited elon- tov & al., 2003). Recent studies of specimens in gate group. Branch leaves similar to stem leaves PE allow to discover also sporophytes or smaller and narrower. Gametangia and sporo- (PE#062123; Yunnan, co. Gongsham, 28.IX.1935 phyte unknown. Hygro- or hydrophytic. coll. [?] # 6462(391)). The operculum is rostrate, Distribution: Madeira. peristome is complete and the seta is rough (sporo- (a) The only species of the genus is known phytes either immature, or old, with considerably from only one place in Madeira, where it grows on broken peristome). The essential characters of this rock near a waterfall. The broad costa, small size species include concave leaves and a costa that and habit suggest a relationship with Rhyncho- reaches almost to the apex. The axillary hairs are stegiella, which, however, never has an as distinct- wide, but relatively short for this group of aquat- ly serrate margin and axillary hairs as long as ic plants. In contrast to D. macroneuron it does those in Nobregaea. Besides Rhynchostegiella, the not have an especially broad costa and the lami- broad costa is known in Brachytheciaceae only na is never bistratose. These characters are, how- in Donrichardsia, which belongs to the same sub- ever, variable in some other genera of Brachythe- family. The latter genus is also similar to Nobre- ciaceae: the bi- or unistratose lamina is known in gaea in having a multistratous lamina. Platyhypnidium (Ochyra & Vanderpoorten, 1999) Species included: N. latinervis Hedenäs. and the costa varies from very narrow to 3/4 of leaf width in Rhynchostegiella. Taking into con- 18. Eurhynchiella Fleisch., Musci Buitenzorg sideration the very high support (Fig. 2), we sug- 4: 1566. 1923. Lectotype: E. zeycheri (C. Müll.) gest to transfer Platyhypnidium patulifolium to Fleisch. (= Hypnum zeycheri C. Müll.), select- Donrichardsia. ed by Ignatov & al. (1999). (c) The Donrichardsia is certainly closely Plants small, yellowish or sordid green, glossy. related to Oxyrrhynchium, being resolved as a Stem prostrate, with weak central strand, mod- sister group (Fig. 2) or sometimes as an ingroup erately densely, julaceously foliate, irregularly (Fig. 1). This affinity is in a nice agreement with or at places regularly pinnately branched, branch the character state of rough seta, a characteristic foliage similar to that in stem. Axillary hairs 3- of most of the Oxyrrhynchium species. celled, pellucid, upper cell pellucid, somewhat Brachytheciaceae (Bryophyta) – a family of sibling genera 275 broader than cells below. Pseudoparaphyllia exceptions are known: for example, seta is smooth orbicular-triangular. Stem leaves rigidly erect- in Oxyrrhynchium vagans and Rhynchostegiel- spreading, ovate-lanceolate, broadly to narrow- la tenella. ly acuminate or somewhat attenuate; margin Species included: serrate to serrulate throughout; costa ending E. zeyheri (C. Müll.) Fleisch. above mid-leaf in a stout spine; laminal cells Species included based on Brotherus (1925): linear, thick-walled, smooth, towards base short E. acanthophylla (Mont.) Fleisch., E. semitorta in about 2 rows, alar cells not differentiated. (Jaeg.) Broth., E. tenuinervis Herzog, E. toncolen- Autoicous. Perichaetial leaves erect, acuminate. sis (Broth.) Broth. (all from South America). Seta smooth, rather short. Capsule inclined, ob- Species excluded from Eurhynchiella: long, contracted below mouth. Annulus sepa- E. decurrens P. Varde – Kindbergia africana rating by fragments. Operculum obliquely ros- (fide Ochyra, 1997) trate. Peristome xerocastique, perfect; exostome red/brown, with incrassate dorsal trabeculae; 19. Rhynchostegiella (B. S. G.) Limpr., endostome complete, cilia 1-2, nodose. Spores Laubm. Deustsch. 3: 207. 1896, nom. cons. – small. Calyptra naked. Mesophytic (tree base in Rhynchostegium subgen. Rhynchostegiella shaded place). B. S. G., Bryol. Eur. 5: 201. 1852. Type: Rhyn- Distribution: South Africa +?South America. chostegium tenellum (Dicks.) B. S. G. (= Hyp- (a) The selection of the lectotype of the ge- num tenellum Dicks., Rhynchostegiella tenella nus was not commented on by Ignatov & al. (Dicks.) Limpr.). (1999). Our argument was that the genus is based (Rhynchostegiella is conserved against Re- on Rhynchostegiella sect. Leptorhynchostegium myella C. Müll., which, however is considered Broth., to which six South-African species were here as a distinct genus) referred (without description) by Carl Müller Plants small, soft to very rigid, yellowish to (1899). Later five of these six species were syn- deep green or brownish. Stem prostrate, with onymized by Dixon & Gepp (1923) and by weak and discontinuous lacking central strand, Sims (1926 [cited in Index Muscorum]) with loosely to moderately densely foliate, remotely the sixth species, Rhynchostegiella (Eurhynchiel- to rather regularly pinnately branched, branch la) zeyheri. Thus, the only one South African spe- foliage similar to that in stem. Axillary hairs 2- cies is a good candidate for lectotype. 3-celled. Pseudoparaphyllia acuminate or acute. (b) The genus was not included in our anal- Stem leaves erect to spreading, in dry state yses of molecular data and its affinities are un- slightly different or with somewhat incurved clear and need further studies. Morphology gives margins, lanceolate to narrowly lanceolate, controversial evidences. The plants are small, with acuminate, not or hardly concave in basal por- smooth setae reminiscent of Eurhynchiastrum tion; margin wavy-crenulate to subentire; cos- pulchellum. Actually, one of the synonyms of ta thin and ending below mid-leaf to broader Eurhynchiella zeycheri is called Rhynchostegium than half leaf and excurrent, without spine, lam- afro-strigosum C. Müll. (Eurhychium strigosum inal cells elongate to linear, firm-walled; to- is a well-known synonym of E. pulchellum). wards base somewhat shorter, alar cells not dif- However, the uppermost cell of the axillary hairs ferentiated or a few cells in leaf corner sub- is broad in Eurhynchiella, sometimes almost in- quadrate. Branch leaves similar to stem leaves flated, thus suggesting a relationship between or smaller and narrower. Autoicous. Pericha- this genus and Oxyrrhynchium. Somewhat less etial leaves acuminate, straight or with slightly direct evidence on the close position of Eu- reflexed acumina. Seta long, rough to smooth. rhynchiella relative to Oxyrrhynchium is the fact, Capsule inclined to horizontal, ovate. Annulus that leaves of E. zeycheri are sometimes attenu- separating by fragments. Operculum longly ate and similarly attenuate leaves are known from obliquely rostrate. Peristome xerocastique or South American and New Zealandian O. remo- hygrocastique, moderately modified; exostome tifolium, a species with a rough seta. Though teeth red-brown, papillose above, cross striolate most species of Oxyrrhynchium (and Rhyncho- below, strioles smooth to moderately papillose; stegielloideae as a whole) have a rough seta, few endostome with high basal membrane, segments 276 MICHAEL S. IGNATOV & SANNA HUTTUNEN moderately broad, non- or slightly perforated R. teneriffae (Mont.) Dirkse et Bouman along keel, cilia long. Spores small. Calyptra Species, probably belonging to Rhyncho- naked. Xeromesophytic to hygromesophytic. stegiella: Distribution: Western Palearctic + ?East R. holstii (Broth.) Broth. Africa. Species excluded to other genera: (a) Ignatov & al. (2003) discussed the spe- R. brachypodia Fleisch. – Remyella cies of the genus in East Asia and found that R. cucullata (Mitt.) Broth. ex Par. – see note most of them can be classified into two groups “a” (“R. sinensis”–group) in that area. The first group comprises delicate R. hawaica (C. Müll.) Broth. – Remyella plants, usually growing on twigs, that have spic- R. laeviseta Broth. – Rhynchostegium psi- ulose leaves when dry. This group is treated lopodium. here as Remyella (see below). The second group R. leptoneura Dix. et Thér. – ?Remyella (“Rhynchostegiella sinensis”-group) includes R. menadensis (Sande Lac.) E. B. Bartram – small, dull plants, with lanceolate to ovate-lan- ?Remyella ceolate leaves, short, thick-walled laminal cells, R. mindorensis (Broth.) Broth. – see note “a” often with a collapsed cytoplasm pattern, sub- (“R. sinensis”–group) quadrate alar cells extending up along the mar- R. muriculata (Hook. et Wils.) Broth. – see gin, a slightly rough seta, a weakly inclined note “a” (“R. sinensis”–group) short capsule, and small spores. This group in- R. opacifolia Dix. – see note “a” (“R. sinen- cludes R. sinensis Broth. et Par., R. santaiensis sis”–group) Broth. et Par., R. mindorensis (Broth.) Broth., R. papuensis E. B. Bartram – see note “a” R. opacifolia Dix., and R. papuensis E. B. Bar- (“R. sinensis”–group) tram. The Australian R. cucullata (Mitt.) Broth. R. ramicola (Broth.) Broth. – Remyella ex Par. also seems to fit this group very well. R. sakuraii Takaki – Brachythecium cf. plu- The latter species was placed in Scorpiurium mosum (see Ignatov & al., 2003). by Hedenäs (1996), due to the similarity in leaf R. santaiensis Broth. & Par. – see note “a” structure. However Scorpiurium species have a (“R. sinensis”–group) smooth seta. It seems like this group of South- R. santosii E. B. Bartram – Remyella East Asian, Malesian and Australian Rhyncho- R. sinensis Broth. et Par. – see note “a” (“R. stegiella is closer to South American Helicodon- sinensis”–group) tium or maybe Flabellidium. Although with some R. sumatrana Fleisch. – Remyella hesitation we add to this group one more Aus- R. tenuicaulis (Spruce) Karttunen – not tralian species, R. muriculata (Hook. et Wils.) Brachytheciaceae, see note “b” Broth.: it has unusually long laminal cells for R. vriesei (Dozy et Molk.) Broth. – Remyella this group, but is otherwise very similar. The R. zeyheri (Spreng. ex C. Müll.) Broth. – final decision has to be confirmed by molecu- Eurhynchiella lar studies. (b) We were able to study the isotype of 20. Helicodontium Schwägr., Sp. Musc. Frond., Cirriphyllum germanicum (H), which has been Suppl. 3(2): 293. 1830. Type: H. tenuirostre synonymized with Rhynchostegiella tenuicau- Schwägr. = H. capillare (Hedw.) Jaeg. lis by Karttunen (1990), and found that the ?Dubyella Schimp., Musci Europaei Novi, pseudoparaphyllia are not of the Brachytheci- fasc 3-4. 1866. Type: D. italica Schimp. acean pattern. Plants small, slender, sordid green to brown- (c) The revision of Dirkse & Bouman (1995) ish, dull. Stem creeping, with central strand, for Canary Islands covers most taxa of the ge- moderately densely foliate, irregularly branched, nus in strict sense. branches moderately densely foliate. Axillary Species included: hairs 2-3-celled, pellucid. Pseudoparaphyllia R. curviseta (Brid.) Limpr. orbicular-triangular. Stem leaves appressed R. litorea (De Not.) Limpr. when dry, more or less spreading when wet, ovate R. macilenta (Ren. et Card.) Card. to lanceolate, gradually tapering to blunt apex, R. tenella (Dicks.) Limpr. slightly concave; margin crenulate almost Brachytheciaceae (Bryophyta) – a family of sibling genera 277 throughout, narrowly recurved in lower half; codontium in all important points. costa ending at 0.6-0.7(0.8) of leaf length, with- 21. Clasmatodon Hook. et Wils., J. Bot. out spine; laminal cells rhombic to rhombic- (Hooker) 4: 421. 1842. Type: C. pusillus Hook. ovate, thick-walled, cell walls sometimes sinu- et Wils. = C. parvulus (Hampe) Sull. ous; alar cells oblate to subquadrate, extending ?Anisodon B. S. G., Bryol. Eur. 5: 75. 1852. up along margin. Branch leaves similar to stem Type: Anisodon perpusillus B. S. G. leaves. Autoicous. Perichaetial leaves erect, rather ?Phlogostomum Hampe, Flora 50: 74. 1867, broadly acuminate. Seta relatively long, slight- nom. illeg. ly rough to smooth. Capsule erect, ovate. Annu- Plants small, slender, sordid green to brown- lus separating by fragments. Operculum oblique- ish, dull. Stem creeping, with central strand, ly rostrate. Peristome xerocastique, moderately densely foliate, irregularly branched, branches modified; exostome teeth red-brown, cross-stri- densely foliate. Axillary hairs 2-celled, pellucid, olate below; endostome with moderately high upper cell sometimes vesiculate-inflated. basal membrane, segments moderately broad, Pseudoparaphyllia orbicular-triangular. Leaves perforated, cilia absent. Spores small. Calyptra closely imbricate when dry, more or less spread- naked. Xeromesophytic. ing when wet, ovate to suborbicular, shortly Distribution: South and Central America acuminate or acute, slightly to strongly con- and Mexico + ?. cave; margin serrulate to subentire; costa end- (a) This genus was for a long time placed ing in mid-leaf without spine; laminal cells in the Fabroniaceae and then in the Myriniace- rhombic to rhombic-ovate, thick-walled, lumina ae, until molecular analysis resolved it as a with round ends; alar cells oblate to subquad- member of Brachytheciaceae (Buck & al., 2000). rate, extending up along margin. Branch leaves This placement is in agreement with the pseudo- similar to stem leaves. Autoicous. Perichaetial paraphyllia pattern in the genus. leaves erect, rather broadly acuminate. Seta rel- (b) The close relationship between Helico- atively long, smooth. Capsule erect, ovate. An- dontium and Clasmatodon and Homalotheciella nulus of 3 rows of cells, separating by fragments. was strongly supported by the present analysis Operculum obliquely rostrate. Peristome strong- (Figs. 1-2), and the same topology has also been ly modified; exostome teeth pale, very short, found by Buck & al. (2000). However each of hardly exceeding mouth level; endostome with these three genera has unique morphological very low basal membrane, filiform, much spaced characteristics, so we prefer to retain them dis- segments, cilia absent. Spores small. Calyptra tinct at the generic level, until the delimitation naked. Xeromesophytic. of Helicodontium from the large group of Rhyn- Distribution: temperate Eastern North America. chostegiella species centered around R. sinen- (a) Similarly to Helicodontium, this genus was sis has been critically addressed (Ignatov & for a long time placed in the Fabroniaceae and al., 2003, see also discussion under Rhynchoste- then in the Myriniaceae, until molecular analysis giella above). suggested close relationship with the Brachy- (c) Other relatives of this genus have to be theciaceae (Buck in Crosby & al., 1999, Buck & searched among “short-celled Rhynchoste- al., 2000). This placement is in agreement with giella” of East Asia, Malesia, and neighbouring the pseudoparaphyllia pattern in the genus. areas (group 2 in discussion under Rhyncho- (b) Morphologically Clasmatodon is one of stegiella). the most unusual representatives of Brachy- Species included: theciaceae due to the strong reduction of the H. capillare (Hedw.) Jaeg. exostome. Only thin, cilia-like segments can Buck (1980a) synonymized a number of spe- slightly affect the spore dissemination. The plants cies described from South America with H. cap- are also very small and the leaf laminal cells illare, the only species on that continent. Spe- are shortly ovate or rhombic. Phylogenetic in- cies from other regions need to be revised. Heli- ferences from sequence data suggest close af- codontium italicum (Schimp.) Fleisch., the only finities to other New World taxa, namely species from Europe, is known only from the Homalotheciella and Helicodontium capillare. type collection. Its description agrees with Heli- 278 MICHAEL S. IGNATOV & SANNA HUTTUNEN

(c) Anisodon perpusillus B. S. G., the type erus' (1925) view that it is closely related to of Anisodon, was described from a single col- Scorpiurium. The two genera differ by the fron- lection from Germany. The description in “Bry- dose branching and thick-walled but not opaque ologia Europaea” referred, to our mind, to Habro- alar cells of Flabellidium. These characters don perpusillus. We have no explanation why would also be congruent with a relationship Schimper (1860) especially underlined that his with Helicodontium that is widespread in the Anisodon is not the same as Habrodon, and ac- South America. The size of plants of Helicodon- cepted Sullivant's opinion (cited in Schimper, tium is the same as of Flabellidium, while Medi- l. c. ), that Anisodon is the same as the North terranean Scorpiurium species are usually larger. American Clasmatodon. Surprisingly, the de- Helicodontium capillare, the only species of the scription and illustration of Clasmadon in Lim- genus retained for Latin America by Buck pricht (1896) are correct. We however suspect (1980a), differs from Flabellidium in having a that they were based on non-European materi- somewhat rounded leaf, blunt marginal teeth, al: Duell (1994) reported no proofed materi- and a costa that is not or only very slightly als from Germany. Hence, Anisodon is probably projecting. A similar morphology is characteris- a earlier synonym of Habrodon. However, a fi- tic for some East Asian species traditionally re- nal decision can only be made once the expand- ferred to Rhynchostegiella (see discussion un- ed search of the original collections of Anisod- der that genus). on would give no results. Species included: (d) The name Phlogostomum was suggest- Flabellidium spinosum Herz. ed by Hampe to replace Anisodon, since he 23. Homalotheciella (Card.) Broth., Nat. Pfl. thought that this name was already used in the 1(3): 1133. 1908. – Homalothecium sect. Homa- Solanaceae. He obviously confused it with Ani- lotheciella Card., Bull. Herb. Boissier 7: 374. 1899. sodus (Solanaceae). Type: Homalothecium subcapillatum (Hedw.) Species included: Sull. (= Pterigynandrum subcapillatum Hedw.) Clasmatodon parvulus (Hampe) Sull. = Homalotheciella subcapillata (Hedw.) Broth. 22. Flabellidium Herz., Biblioth. Bot. 87: 163. Plants small, soft, pale yellowish-green. Stem 1916. Type: F. spinosum Herz. prostrate to ascending, with weak and at places Plants small, rather rigid, sordid to yellow- lacking central strand, densely foliate, densely, ish green. Pseudoparaphyllia triangular. Stem but irregularly pinnately branched, branch foli- creeping to ascending, with central strand and age similar to that in stem. Axillary hairs 2-celled. 2-3-stratose sclerodermis, densely foliate, irreg- Pseudoparaphyllia orbicular-triangular, blunt. ularly pinnately branched, branches slightly Stem leaves erect, from narrowly elliptic or obo- arcuate, foliage same as that of stem. Stem leaves vate base acuminate, concave; margin serrate to erect to imbricate, ovate, gradually tapered serrulate to alar group; costa thin, ending short- above, acute, more or less concave, not plicate; ly above mid-leaf without spine; laminal cells margin serrate throughout; costa ending above elongate to linear, thick-walled, flexuose; towards mid-leaf in sharp spine, distally toothed; lami- base cells shorter to short rectangular in 2-3 rows, nal cells rhombic-ovate to shortly elongate-flex- alar cells subquadrate, in rather well delimited uose, thick-walled; in alar area subquadrate, group, well extending up along margin. Branch forming extensive opaque area extending up leaves similar to stem leaves. Autoicous. Pericha- along margin. Branch leaves similar to stem etial leaves acuminate, more or less straight. Seta leaves. Dioicous. Perichaetial leaves straight, long, rough. Capsule erect to slightly inclined, abruptly broadly acuminate, costa reaching ovate to oblong. Annulus separating by fragments. above mid-leaf. Seta smooth. Calyptra naked. Operculum moderately longly rostrate. Peristome Mesophytic? (on tree roots in forest). Other hygrocastique, strongly modified; exostome teeth sporophytic characters unknown. pale, papillose almost to base; endostome with Distribution: Bolivia. low basal membrane, rather strongly papillose (a) The genus is only known from the holo- segments, segments free or adherent to exostome type collected in Bolivia. Enroth (1995), who teeth, cilia absent. Spores large. Calyptra hairy. monographed Flabellidium, agreed with Broth- Mesophytic to xeromesophytic. Brachytheciaceae (Bryophyta) – a family of sibling genera 279

Distribution: temperate Eastern North America. fracting; basal cells shorter, irregular in shape, (a) This genus has a strongly modified peris- more strongly thick-walled in few rows across tome with the endostome mostly adherent to base or only in alar area. Branch leaves smaller the exostome. In the Brachytheciaceae a similar than stem leaves and acuminate to acute. Auto- case is found only in Homalothecium philip- icous. Perichaetial leaves gradually tapered to peanum and H. laevisetum, which explains why reflexed acumina. Seta long, smooth. Capsule Homalotheciella subcapillata was first placed erect, ovate. Annulus of 1-2 rows of large cells, in Homalothecium, where it was accommodat- hardly separated. Operculum obliquely rostrate. ed in a distinct section. In all published sys- Peristome hygrocastique, modified; exostome tematic arrangements Homalotheciella has been teeth red-brown, not spaced at mouth level, on considered close to Homalothecium. outer surface papillose above and cross striolate (b) The phylogenetic inferences made from below; endostome with low basal membrane, our data are congruent with those of Buck & narrow segments, split along keel, and very short al. (2000) in that Homalotheciella appears only or absent cilia. Spores small. Mesophytic or xe- distantly related to Homalothecium, but with romesophytic (on tree at high elevation). close affinities to Helicodontium capillare and Distribution: Bolivia. Clasmatodon. The latter are also epiphytes in (a) When describing this species, Herzog (Bibl. the New World, and share with Homalotheciella Bot. 87: 165. 1916) provided a rather detailed strongly modified peristomes (though in dif- illustration of it. The shape of the operculum is ferent manners), and subquadrate alar cells described by Williams (1910), we did not see extending up the margin in an elongate group, any. The genus is known by only one collection. but differ by their shorter laminal cells, small- (b) Gametophytically this species is some- er and blunt leaves and subentire to bluntly what similar to Brachythecium acuminatum or crenulate margins. Another possible affinity of Homalotheciella, but the costa is excurrent and Homalotheciella is with three genera that were the seta is smooth. A position near the latter not included in our phylogenetic analyses, genus is most probable. Schimperella, Mandoniella, and Juratzkaeella. Species included: Species included: M. spicatinervia (R. S. Williams) Herz. H. subcapillata (Hedw.) Broth. 25. Schimperella Thér., Recueil Publ. Soc. Problematic species: Havraise Études Diverses 1925: 26. 1926. Type: H. sinensis Card. et Thér. S. rhynchostegioides Thér. H. tenerrima (C. Müll.) E. B. Bartram Rhynchocarpidium P. Varde et Leroy, Bull. Jard. Bot. État 18: 188. 1947. Type: R. katalense 24. Mandoniella Herzog, Biblioth. Bot. 87: P. Varde et Leroy. 165. 1916. Type: M. spicatinervia (R. S. Will- Plants small to medium-sized, golden-green. iams) Herzog (= Helicodontium spicatinervi- Stem creeping, with central strand, moderately um R. S. Williams). densely, terete foliate, irregularly branched, branch Plants moderately small, soft, yellowish. Stem and stem foliage similar. Stem leaves erecto- creeping, with central strand, loosely to moder- patent, ovate, acute to broadly acuminate, twist- ately densely foliate, remotely branched, branch- ed just below apex, not or slightly concave, not es moderately densely to densely foliate. Axil- plicate; margin sharply serrate above, serrulate lary hairs on branches 2-celled (not found on below; costa ending above mid-leaf in small spine, stem), upper cells blunt. Pseudoparaphyllia tri- sometimes with additional spine a little below; angular. Stem leaves erecto-patent, ovate-lan- laminal cells long-hexagonal or elongate to lin- ceolate, acuminate, not or hardly concave, some- ear, thin-walled, alar cells short-rectangular, thick- times with 1-2 sharp, but shallow plicae; mar- walled across base and extending up along gin sharply serrate above, serrate to serrulate to margin. Branch leaves narrower and with long- base; costa ending above mid-leaf in long spine, er costa, otherwise similar to stem leaves. Auto- sometimes with additional spine a little below; icous. Perichaetial leaves erect, with recurved laminal cells elongate to linear, with narrowly apices. Seta long, smooth, becoming very dark acute ends, moderately thick-walled, highly re- with age. Capsule erect or suberect. Annulus of 280 MICHAEL S. IGNATOV & SANNA HUTTUNEN

1-2 rows of large separating cells. Operculum Species included: longly, obliquely rostrate. Peristome modified; J. sinensis (Fleisch. ex Broth.) Buck exostome teeth on outer surface papillose above 27. Aerolindigia Menzel, Nova Hedwigia 52: and cross-striolate below; endostome with low 321. 1991. Type: A. capillacea (Hornsch.) Men- basal membrane, narrow segments, split along zel (= Pilotrichum capilaceum Hornsch.). keel, and very short or no cilia. Spores small. Plants small, soft, pale green to yellowish. Calyptra naked. Mesophytic. Stem mostly creeping, sometimes pendent, with Distribution: East Africa and Madagascar. weak central strand, loosely to moderately dense- (a) We did not see any material of this ge- ly foliate, remotely branched, branch foliage sim- nus. The above description is compiled from ilar to that in stem. Axillary hairs 2-celled. Buck (1985, 1993) and Thériot (1926). Pseudoparaphyllia longly acuminate. Stem Species included by Buck (1985, 1993): leaves erecto-patent to slightly reflexed, ovate- S. bello-intricata (Broth.) Buck lanceolate, acuminate, not or hardly concave; S. rhynchostegioides Thér. margin serrate to serrulate; costa ending in mid- 26. Juratzkaeella Buck, Rev. Bryol. Lichénol. leaf or often shortly below, without spine; lam- 43: 312. 1977. Type: J. sinensis (Fleisch. ex inal cells elongate to rhombic-elongate, firm- Broth.) Buck (= Juratzkaea sinensis Fleisch. walled; towards base considerably wider, alar ex Broth.). cells not differentiated. Branch leaves similar Plants moderately small, soft, yellowish. Stem to stem leaves. Autoicous. Perichaetial leaves creeping, thin, with central strand, 2-3-stratose straight. Seta relatively short, 2-5 times longer sclerodermis and only 1-2-stratose medullary zone, than urn, rough. Capsule straight, ovate. Annu- densely foliate, subpinnately branched, branches lus separating by fragments. Operculum short- densely foliate. Axillary hairs 2-3-celled, pellucid. ly rostrate. Peristome xerocastique, moderately Pseudoparaphyllia triangular. Stem leaves erect- modified; exostome teeth narrow, cross-striolate appressed to slightly homomallous, lanceolate, below; endostome with high basal membrane, acuminate, plane to slightly concave; margin ser- moderately broad segments, and no cilia. Spores rulate above, bluntly serrulate below; costa nar- large. Calyptra naked. Xeromesophytic, tolerant row, ending above mid-leaf, without spine; lami- to desiccation. nal cells elongate-rhomboidal, with narrowly acute Distribution: tropical America and Africa. ends, thick-walled, highly refracting; alar cells (a) The genus was revised and illustrated subquadrate, often reaching costa, and extending by Menzel (1991). Ignatov & al. (1999) dis- far up along margin. Branch leaves similar to cussed the possible relationship of Aerolindi- stem leaves. Autoicous. Perichaetial leaves erect, gia to the group of East Asian, Malesian and longly acuminate. Seta long, smooth, strongly twist- Oceanian species, which were uncomfortably ed. Capsule erect or suberect, short-cylindrical. placed in Rhynchostegiella. They argued that Annulus of one row of large cells, hardly separat- based on morphology the latter seem more sim- ing. Operculum obliquely rostrate. Peristome xe- ilar to Aerolindigia than to the European Rhyn- rocastique, slightly modified; exostome teeth red- chostegiella species. The results of our phyloge- dish, not spaced at mouth level, on outer surface netic analyses based on morphological and papillose above and cross-striolate below; endos- molecular data (Figs. 1-2) support the close re- tome with high basal membrane (about 1/3 of lationship between Aerolindigia and at least endostome height), moderately broad segments, one Rhynchostegiella species of this group, the split along keel, and without cilia. Spores small. Malesian R. brachypodia. At the same time, some Habitat unknown, but probably not epiphytic important morphological differences (long and (stem covered with sand). very strongly rough seta, large spores, golden Distribution: China. color) support keeping Aerolindigia as a sepa- (a) The genus is described, illustrated and rate genus. Fortunately this group of Malesian discussed by Buck (1977). It is similar to Homa- and Oceanian Rhynchostegiella has an exist- lotheciella in its gametophytic characters, but ing generic name, Remyella (see below). has a smooth seta and rather completely de- Species included: veloped peristome. A. capillacea (Hornsch.) Menzel Brachytheciaceae (Bryophyta) – a family of sibling genera 281

28. Remyella C. Müll., Flora 82: 477. 1896. Rhynchostegiella vriesei Dozy et Molk., Ann. Type: R. hawaica C. Müll. Sci. Not. Bot., sér. 3, 2: 309. 1844, [‘vriesii’]. Plants small, soft to somewhat rigid, yellow- Species probably belonging to Remyella: ish to rich golden. Stem mostly creeping, some- Rhynchostegiella menadensis (Sande Lac.) times pendent, with weak central strand, loosely E. B. Bartram, R. leptoneura Dix. et Thér., and to densely foliate, remotely to rather regularly Rhynchostegium nano-pinnatum (Broth.) pinnately branched, branch foliage more dense. Kindb. probably relate to this group, but the Axillary hairs 2-celled. Pseudoparaphyllia long- seta is smooth in these species, the spores are ly acuminate. Stem leaves spreading to reflexed, small, and the areolation is somewhat narrower, in dry state with similar orientation, but in some than in large-spored species of the genus. Hence species becoming tubulose-spiculose due to in- the further studies are needed before nomencla- curved to inrolled margins, ovate-lanceolate, tural transfers. acuminate, not or hardly concave in basal por- tion; margin serrate to serrulate; costa ending in 29. Meteoridium (C. Müll.) Manuel, Lind- mid-leaf or below, without spine, laminal cells bergia 4: 47. 1977. – Neckera subsect. Meteor- elongate to rhombic-elongate, firm-walled; to- idium C. Müll., Syn. Musc. Frond. 2: 672. 1851. wards base considerably wider, alar cells not dif- Type: Neckera remotifolia (C. Müll.) Hornsch. ferentiated. Branch leaves smaller and narrower, ex C. Müll. (= Leskea remotifolia C. Müll., Me- otherwise similar to stem leaves. Autoicous. Per- teoridium remotifolium (C. Müll.) Manuel). ichaetial leaves with reflexed acumina. Seta long, Plants large, soft, green to yellowish. Stem very rough. Capsule erect, cylindrical. Annulus creeping to pendent, with weak and at places not separating. Operculum with long beak. Peris- absent central strand, loosely foliate in creep- tome xerocastique, moderately modified; exos- ing part, more densely foliate in pendent part, at tome teeth red-brown, strongly papillose above, places rather regularly remotely branched, cross-striolate below, strioles papillose, inner sur- branch foliage same as that of pendent part of face incrassate; endostome with high or rather stem. Axillary hairs 4-5-celled. Pseudoparaphyl- low basal membrane, segments moderately broad, lia triangular. Stem leaves widely spreading, non- or slightly perforated along keel, cilia short broadly ovate to ovate-lanceolate, longly acumi- to slightly above half of segment length. Spores nate to attenuate, not or hardly concave; mar- large. Calyptra naked. Xeromesophytic, tolerant gin serrulate; costa ending in mid-leaf or just to desiccation. above, without spine, laminal cells linear, thick- Distribution: East Asia, Malesia, Oceania, and walled; alar cells scarcely differentiated, but a ?Australia. few subalar cells somewhat enlarged, shortly (a) This group was briefly commented un- rectangular, rather thick-walled. Branch leaves der Rhynchostegiella (see above) and also by similar to stem leaves. Dioicous. Perichaetial Ignatov & al. (1999, 2003). leaves with +erect to slightly reflexed acumi- Species included: na. Seta 1.5-2 times longer that urn, smooth. R. brachypodia (Fleisch.) comb. nov. – Rhyn- Capsule emergent from perichaetium, straight. chostegiella brachypodia Fleisch., Musci Buiten- Annulus separating by fragments. Operculum zorg 4: 1564. 246. 1923. shortly rostrate. Peristome xerocastique, moder- R. hawaica C. Müll. ately modified; exostome teeth narrow, cross- R. ramicola (Broth.) comb. nov. – Rhyn- striolate below; endostome with very low basal chostegium ramicola Broth., Bull. Soc. Roy. Bot. membrane, narrow filiform segments and no Belgique 41(1): 91. 1905. cilia. Spores large. Calyptra naked. Mesophytic, R. santosii (E. B. Bartram) comb. nov. – tolerant to desiccation. Rhynchostegiella santosii E. B. Bartram, Phil- Distribution: tropical America. ipp. J. Sci. 87: 279. 1958. (a) This genus was segregated from Meteo- R. sumatrana (Fleisch.) comb. nov. – Rhyn- riopsis by Manuel (1977a), who recognized it chostegiella sumatrana Fleisch., Musci Buiten- for a single species. Originally the name was zorg 4: 1566. 1923. given to a monotypic section of Neckera. R. vriesei (Dozy et Molk.) comb. nov. – (b) In our analysis Meteoridium was always 282 MICHAEL S. IGNATOV & SANNA HUTTUNEN sister to Squamidium plus Zelometeorium (Figs. very homogenous. At least in the type section 1-2). Meteoridium differs from two latter gen- one species has small and thick-walled alar cells era in its xerocastique, relatively complete peris- (incl. S. brasiliense, used in the present analy- tome and naked calyptra (we did not see any, sis), whereas the other four species have inflat- information about calyptra is from Crum, 1994). ed alar cells. This group of three pendent tropical, mainly (b) Our analysis showed a very high sup- American genera form a rather isolated clade, port for a close relationship between Zelome- sister to the remainder of the subfamily Rhyn- teorium and Squamidium. This allows to hy- chostegielloideae. pothesize that an analysis of all species of Squ- Species included: amidium and Zelometeorium might show more M. remotifolium (C. Müll.) Manuel complex interrelations among their species. Species included: Species included by Buck (1999): S. brasiliense (Hornsch.) Broth. M. tenuissimum (Hook. et Wils.) M. A. Lewis Species included by Allen & Crosby (1986): 30. Squamidium (C. Müll.) Broth., Nat. Pfl. S. diversicoma (Hampe) Broth., S. isocladum 1(3): 807. 1906. – Meteorium sect. Squamidi- (Ren. et Card.) Broth., S. leucotrichum (Tayl.) um C. Müll., Linnaea 42: 420. 1879. Type: Me- Broth., S. livens (Schwägr.) Broth., S. macro- teorium lorentzii C. Müll. [‘i’] (= Squamidium carpum (Spruce ex Mitt.) Broth., S. nigricans lorentzii (C. Müll.) Broth.) = S. brasiliense (Hook.) Broth. (Hornsch.) Broth. Plants large, soft, green to pale yellow, old 31. Zelometeorium Manuel, J. Hattori Bot. parts tinged with black to totally black. Stem Lab. 43: 110. 1977 [1978]. Type: Z. patulum creeping to pendent, with weak and discontinu- (Hedw.) Manuel (= Hypnum patulum Hedw.). ous central strand, rather densely foliate. Axil- Plants large, soft, green to yellowish, old parts lary hairs 3-7-celled. Pseudoparaphyllia trian- tinged with black to totally black. Stem creep- gular. Stem leaves strongly concave in basal part, ing to pendent, with weak and at places absent resulting in terete foliage, broadly ovate to ovate, central strand, loosely foliate in creeping part, gradually or abruptly tapered into short or long more densely foliate in pendent part, at places and sometimes flexuose acumen; margin serru- rather regularly remotely branched, branch foli- late or subentire; costa ending in mid-leaf or age same as that of pendent part of stem. Axil- above without spine, laminal cells linear, thick- lary hairs 3-4-celled. Pseudoparaphyllia trian- walled; alar cells weakly differentiated, a few gular. Stem leaves spreading to recurved from subalar cells enlarged, shortly rectangular, thick- clasping base, broadly ovate and suborbicular to or thin-walled. Branch leaves similar to stem ovate-lanceolate, acute to acuminate, acumen leaves. Dioicous. Perichaetial leaves with +erect sometimes attenuate-flexuose, margin serrulate; acumina. Seta smooth to slightly rough. Capsule costa ending in mid-leaf or above without spine, immersed to emergent, straight. Annulus sepa- laminal cells linear, relatively thin-walled; alar rating by fragments. Operculum shortly rostrate. cells weakly differentiated, a few subalar cells Peristome hygrocastique, modified; exostome teeth enlarged, shortly rectangular, thick-walled. Branch narrow, papillose to base; endostome with rather leaves less longly acuminate. Dioicous. Pericha- high basal membrane, narrow segments, and ru- etial leaves with +erect acumina. Seta as long as dimentary cilia. Spores large. Calyptra hairy. urn, slightly rough to almost smooth. Capsule Mesophytic, tolerant to desiccation. immersed to emergent, straight. Annulus sepa- Distribution: tropical America and Africa. rating by fragments. Operculum shortly rostrate. (a) Squamidium was segregated as a sec- Peristome hygrocastique, modified; exostome teeth tion of Meteorium, and since that time its con- narrow, papillose to base; endostome with low cept remains almost the same. It has been re- basal membrane, narrow segments, and rudimen- vised recently by Allen & Crosby (1986), who tary cilia. Spores large. Calyptra hairy. Meso- accepted 7 species. Despite that the genus is phytic, tolerant to desiccation. not big, the species are grouped into two sec- Distribution: tropical America. tions, and even this does not make the groups (a) This genus was segregated from Meteo- Brachytheciaceae (Bryophyta) – a family of sibling genera 283 riopsis by Manuel (1977b), who included 5 indistinctly plicate; margin serrate, costa end- species in it. See also note under Squamidium. ing above mid-leaf and often at 0.8-0.9 of leaf Species included: length, without spine; laminal cells linear, mod- Z. patulum (Hedw.) Manuel erately thick-walled; alar (or sometimes only subalar) cells shortly rectangular, thin-walled, Species included by Manuel (1977b): Z. indistinctly delimited from neighbouring cells. allionii Manuel, Z. ambiguum (Hornsch.) Man- Branch leaves smaller than stem leaves, other- uel, Z. patens (Hook.) Manuel, Z. recurvifolium wise similar. Autoicous. Perichaetial leaves with (Hornsch.) Manuel slightly reflexed, often rather stiff acumina. Seta SUBFAM. RHYNCHOSTEGIOIDEAE smooth. Capsule inclined to horizontal. Annu- SUBFAM. NOV. lus separating by fragments. Operculum ros- Plantae mediocres vel robustae. Folia lato trate. Peristome xerocastique, perfect. Spores ovata vel anguste lanceolata; cellulae anguste small. Calyptra naked. Hygrophytic, occasion- lineares vel elongatae. Autoica, dioica vel phyl- ally hydrophytic. lodioica. Seta laevissima. Capsula horizontalis, Distribution: Subcosmopolite. raro erecta. Peristomium perfectum, raro imper- (a) Platyhypnidium riparioides was found fectum. Operculum rostratum, raro conicum. in our analysis (Fig. 1-2) to be not distinct from Type genus: Rhynchostegium B. S. G. Rhynchostegium. However, this species was rep- Plants commonly large, rich to pale green to resented in our analysis by cpDNA only, thus yellowish-brown, often characteristically whit- we hesitate to use obtained data for certain ish and with silky (? oily) gloss. Axillary hairs judgements on relationships between Rhyncho- frequently long and their upper cells more stegium and Platyhypnidium. However, their strongly colored than the basal cells. Autoicous, close relationship is very probable. dioicous, phyllodioicous. Seta always smooth. (b) Platyhypnidium was established for Operculum in most groups with sharply delim- aquatic plants and thus placed in the Amblys- ited beak (excepting only Pseudoscleropodi- tegiaceae by Fleischer (1923) and Brotherus um). Peristome perfect, very rarely modified (1925). Subsequent authors usually included its (Palamocladium, Eriodon). species in Rhynchostegium, or more rarely Eu- Genera included: Rhynchostegium, Platyhyp- rhynchium. The most recent revisions, however, nidium, Scorpiurium, Eurhynchium s. str. (= spe- accepted it as a genus of its own (McFarland, cies of E. striatum group), Plasteurhynchium, 1994; Ignatov & al., 1999; Buck, 1999). Hedenäs Palamocladium, Bryoandersonia, Pseudosclero- (2002) also recognized it and stated that the podium, Aerobryum. genus seems to be well circumscribed. Ironically, Genus included based in morphology only: the only species recognized by him in Australia, Eriodon. P. austrinum, has never been found in our anal- 32. Platyhypnidium Fleisch., Musci Buiten- ysis close to P. riparioides. All the analyses put P. zorg 4: 1536. 1923. Lectotype: Platyhypnidium austrinum into the Rhynchostegielloideae, not muelleri (Jaeg.) Fleisch. (= Rhynchostegium Rhynchostegioideae. This position is well in muelleri Jaeg.), selected by Ignatov & al. (1999). agreement with the fact, that P. austrinum is dif- Plants large, deep green, becoming brown- ferent from all other members of Platyhypnidi- ish with age. Stem prostrate, with central strand, um (and also from all other members of the moderately densely, julaceously foliate, old leaf- subfamily Rhynchostegioideae!) in having a less stems often remaining for a long time, ir- rough seta. Another difference between P. austri- regularly branched, branches straight, foliage num and other species of Platyhypnidium with similar to that of stem. Axillary hairs 4-7-celled, smooth setae is the rather short costa (0.6-0.8 vs. upper cell often brownish and verrucose. 0.7-0.9). Our analysis suggests the position of Pseudoparaphyllia triangular. Stem leaves erect Platyhypnidium austrinum sister to the Cirr- or gradually reflexing from an erect base, in iphyllum–clade or Rhynchostegiella–Cirriph- their middle often twisted, broadly ovate or sub- yllum–Homalotheciella clade (Figs. 1-2), but orbicular to ovate-lanceolate, ± broadly acute, more studies are needed to resolve its relation- with acute to blunt apex, smooth, more rarely ships and to define the taxonomic status. Al- 284 MICHAEL S. IGNATOV & SANNA HUTTUNEN though such polyphyly of Platyhypnidium moderately thick-walled; alar cells very weakly appeared rather unexpectedly, this is not the to strongly differentiated, rectangular to isodia- first case when taxa referred to this genus are metric, thin-walled, in distinctly or indistinctly found very distant. Loeske (1911), when segre- delimited group. Autoicous. Perichaetial leaves gating this group (under Platyhypnum), includ- with slightly reflexed acumina or straight, long- ed in it besides P. riparioides also Hygrohyp- ly acuminate. Seta smooth. Capsule inclined to num dilatatum, H. molle, and H. alpinum (now horizontal, often relatively small (comparative- not in Brachytheciaceae). The aquatic environ- ly with that of Brachythecium), strongly con- ment seems to affect morphology strongly, re- tracted below mouth. Annulus separating by frag- sulting in convergent similarities. ments. Operculum obliquely rostrate. Peristome (c) Chinese Platyhypnidium patulifolium xerocastique, perfect. Spores small. Calyptra na- was found very closely related with Donri- ked. Mesophytic. chardsia macroneuron, and thus is discussed Distribution: Subcosmopolite (though ab- under Donrichardsia. sent in most of boreal zone in Holarctic). Species included: (a) It is rather surprising, that this big and P. aquaticum (Jaeg.) Fleisch. relatively diverse genus, which was thought to P. muelleri (Jaeg.) Fleisch. be heterogeneous, appears monophyletic [with P. pringlei (Card.) Broth. the only exception of Platyhypnidium ripario- P. riparioides (Hedw.) Dix. ides, whose position remains ambiguous due to the incomplete data set for this species]. We at- Species probably belonging to Platyhypnidium: tempted to cover this genus both from the mor- Eurhynchium fuegianum Card. phological and geographical points of view and Species excluded from Platyhypnidium: included in the analysis both tropical species, P. patulifolium (Card. et Thér.) Broth. – “typical Steerecleus”, such as R. pallidifolium Donrichardsia and R. serratum (pale, large leaved, complanate), P. austrinum (Hook. et Wils.) Fleisch. – R. murale (imbricate, deeply green), R. confer- unclear generic position within the Rhynchost- tum (usually green, often with costa ending in a egielloideae, maybe close to Oxyrrhynchium. spine), R. psilopodium (green, narrow leaved 33. Rhynchostegium B. S. G., Bryol. Eur. 5: species, previously referred to Rhynchostegiel- 197. 1852. Type: R. confertum (Dicks.) B. S. G. la), and R. rotundifolium (with exceptionally (= Hypnum confertum Dicks.). lax areolation, dark green to brownish). Steerecleus H. Robinson, Mem. New York Bot. (b) According to the present results, the sta- Gard. 45: 480. 1987. Type: S. serrulatum (Hedw.) tus of Steerecleus as independent genus is not H. Robinson (= Hypnum serrulatum Hedw.). supported, though species of this segregate, namely Scleropodiopsis Ignatov, Arctoa 7: 149. 1998. R. serrulatus and R. pallidifolius, appear in the syn. nov. Type: S. laxiretis Ignatov =Rhyncho- present analysis close to each other (Figs. 1-2). stegium arcticum (I. Hag.) comb. nov., syn. nov. (c) Scleropodiopsis laxiretis was found to Plants large or medium-sized, light green, be- be identical with Rhynchostegium murale var. coming argentate-whitish with age. Stem pros- arcticum Hagen, which, however seems to merit trate, with central strand, loosely to densely, ju- a species rank. laceously or complanately foliate, irregularly (d) Being widespread and variable, some branched, branches straight, foliage similar. Axil- Rhynchostegium species exhibit characters, which lary hairs 3-6-celled, upper cell often brownish were considered as characteristic for Eurhynch- and verrucose. Pseudoparaphyllia triangular. ium s. l. and moreover diagnostic for this genus Stem leaves from erect to patent, more rarely versus Rhynchostegium. For example R. confer- imbricate, in middle often twisted, broadly ovate tum often has its costa ending in a spine, the or lanceolate, acuminate or more rarely acute, not foliage is not complanate at all in narrow-leaved plicate; margin serrate to serrulate, occasionally species, like R. psilopodium, etc. Thus it is not subentire, costa ending shortly above mid-leaf, surprising, that authors failed to find stable char- occasionally below mid-leaf, without or with acters to segregate these two genera and there- spine; laminal cells linear, moderately thin- to fore considered them congeneric. Brachytheciaceae (Bryophyta) – a family of sibling genera 285

(d) The genus Rhyncho-hypnum Hampe is 34. Eriodon Mont., Ann. Sci. Nat. Bot. ser. 3, 4: usually considered as established by Hampe 98. 1845. Type: E. conostomus Mont. based on his earlier (Hampe, 1852) subgenus of ?Helicodontiadelphus Dix., J. Bot. 74: 5. 1936. Hypnum subg. Rhynchohypnum. The latter in- Type: Helicontiadelphus australiensis Dix. = cluded two species, Hypnum confertum and H. Eriodon cylindritheca (Dix.) Dix. et Sainsb. murale (here in Rhynchostegium), that, however, (synonymized by Buck, 1981a). are not mentioned in the paper where Rhyncho- Plants medium-sized, soft, light green, whit- hypnum is validated as a genus. Seems, Hampe ish with age. Stem creeping to pendent, without never transferred neither Rhynchostegium con- central strand, loosely to moderately densely fertum, nor R. murale to Rhyncho-hypnum, where foliate, remotely branched, branch foliage similar he classified mostly species now referred to the to that in stem. Axillary hairs 2-3-celled, very Sematophyllaceae. We retain the problem of lec- short. Pseudoparaphyllia triangular. Stem leaves totypification of Rhynchohypnum for future. widely spreading, ovate-lanceolate, longly acumi- Maybe this generic name can be used in the Se- nate, not or hardly concave; margin sharply ser- matophyllaceae. rate above, serrulate to base; costa ending above Species included: mid-leaf without spine, laminal cells linear, mod- Rhynchostegium arcticum (I. Hag.) comb. nov. erately thick-walled; at base cells wider, espe- – R. murale var. arcticum I. Hag., Tromsø Mus. cially a few cells in submarginal area. Branch Aarsh. 21-22 (3): 305. 1904. – Scleropodiopsis leaves smaller than stem leaves, otherwise simi- laxiretis Ignatov, Arctoa 7: 150. 1998. syn. nov. lar. Autoicous. Perichaetial leaves straight, short- R. assumptionis Besch. ly acuminate. Seta very long, thin and smooth. R. beskeanum (C. Müll.) Jaeg. Capsule erect, longly cylindrical or slightly R. brachypterum (Hornsch.) Jaeg. curved (in Leskea manner). Annulus separating R. celebicum (Sande Lac.) Jaeg. by fragments. Operculum very long, straight, grad- R. comorae (C. Müll.) Jaeg. ually tapered to rostrum. Peristome hygrocas- R. complanum (Mitt.) Jaeg. tique, strongly modified; exostome teeth narrow, R. compridense (C. Müll. ex Broth.) Par. very evenly and longly tapered, pale yellowish, R. confertum (Dicks.) B. S. G. spaced at mouth level, on outer surface papillose R. contractum Card. to base, inner surface papillose to base; endos- R. distratum (Hampe) Jaeg. tome colored stronger than exostome, lightly R. duthiei C. Müll. ex Dix. orangish, with low basal membrane, narrow fili- R. herbaceum (Mitt.) Jaeg. form segments, rigidly standing and sometimes R. hookeri Jaeg. twisting distally (in Tortula manner), with long R. inclinatum (Mitt.) Jaeg. gaps along keel, cilia absent. Spores large. Calyp- R. javanicum (Bél.) Besch. tra naked. Mesophytic, tolerant to desiccation. R. laxatum (Mitt.) Par. Distribution: Chile and Argentina. R. megapolitanum (Bland. ex Web. et Mohr) (a) The above description is based on Eri- B. S. G. odon conostomus, the only species undoubtedly R. murale (Hedw.) B. S. G. referred to the genus. R. ovalifolium Okam. (b) The genus is unique in having the long- R. pallidifolium (Mitt.) Jaeg. est peristome in the family. Leaf texture and Rhynchostegium psilopodium nom. nov. – areolation show certain similarities with Rhyn- Rhynchostegiella laeviseta Broth., Symb. Sin. 4: chostegium, but differ from most of its species 109. 1929. in a very short hyaline axillary hairs. See also R. rotundifolium (Brid.) B. S. G. below discussion under E. cylindritheca. R. scariosum (Tayl.) Jaeg. Species included: R. sellowii (Hornsch.) Jaeg. Eriodon conostomus Mont. R. serrulatum (Hedw.) Jaeg. Problematic species: R. tenuifolium (Hedw.) Reichardt Eriodon cylindritheca (Dix.) Dix. et Sains- A number of names are synonymized by Ig- bury. Buck (1981b) retained this species in Eri- natov & al. (2003). 286 MICHAEL S. IGNATOV & SANNA HUTTUNEN odon, despite several differences in sporophytic membrane and narrow segments (much shorter characters. Hedenäs (2002) included it in Rhyn- than in E. conostomus). Hence, the position of this chostegium, where it was originally placed. He- species in Eriodon is unlikely. denäs stated that this species is gametophytical- 35. Aerobryum Dozy et Molk., Ned. Kruidk. ly very close to Rhynchostegium, as peristome Arch. 2(4): 279. 1851. Type: A. speciosum (Dozy specialization is known in many tropical groups. et Molk.) Dozy et Molk. (= Meteorium specio- However, to our mind, Eriodon conostomus is also sum Dozy et Molk.). derived from Rhynchostegium, and shares with Plants very large, light green to whitish and it several gametophytic characters [Brotherus' soon becoming dark brown to black. Stem creep- (1925) illustration and statement that laminal ing to pendent, with central strand, moderately cells are strongly prorate in this species is mis- densely, julaceously foliate, in creeping parts leading – we found, that cells are smooth or very sometimes subcomplanate, very sparsely branched, slightly prorate, not differing in this respect from branch foliage similar to that in stem. Axillary all other studied Rhynchostegium species]. Sim- hairs 4-6-celled, uppermost cells pale brown. ilarly to E. conostomus, E. cylindritheca grows on Pseudoparaphyllia triangular. Stem leaves spread- twigs and its sporophyte can be regarded as a ing from erect base, sometimes with clasping base grade toward the ultimate specialization of E. (due to U-shaped insertion on stem), very broadly conostomus. It has slightly curved capsules, grad- ovate, rather shortly acuminate, not plicate, not ually tapered long and broad beaks, a pale exos- concave, but somewhat conduplicate and twist- tome, and even an orangish endostome, which is, ed at mid-leaf; margin serrulate to subentire, costa however, much less modified from the basic hyp- ending shortly below mid-leaf without spine; noid type, than that of E. conostomus. Whether laminal cells elongate to linear in upper leaf, the South American E. conostomus and the Aus- firm-walled and often cell wall thickenings re- tralasian E. cylindritheca, arose via reduction from sulting in bone-shaped pattern; basal cells slight- distinct ancestors or a shared ancestor remains ly larger, more strongly porose, alar cells undif- to be critically tested. Whether Eriodon should ferentiated; a few subalar cells sometimes slightly be retained as a distinct genus eventhough Rhyn- enlargened, short-rectangular, thick-walled. chostegium would thereby be paraphyletic [if Branch leaves similar to stem leaves. Dioicous the above written hypothesis on its affinity is or phyllodioicous. Perichaetial leaves with re- true, of course], we can only repeat the same as for flexed acumina. Seta smooth, rather short, 2-4 Unklejackia (cf. discussion on p. 266). times longer than urn. Capsule inclined to hori- Eriodon radicalis Spruce ex Jaeg. This species zontal. Annulus separating by fragments. Oper- has been referred to Entodontopsis (Stereophyl- culum shortly rostrate. Peristome xerocastique, laceae) by Buck (1981b), who reported filamen- perfect. Spores small. Calyptra naked. Mesophytic. tose pseudoparaphyllia in this species (no speci- Distribution: East Asia and Malesia. mens cited, but the drawing of leaf and areola- (a) This genus was only recently transferred tion is called “from type, NY”). We were able to to the family (Buck & Goffinet, 2000). Thus, its study an isotype in H-BR (1401. Eriodon radica- relation to other genera of the Brachytheciaceae lis Spruce. Andes Quitoensis. Leg. R. Spruce) and was never a subject for discussion. According to found that it has Brachytheciaceaen pattern of the results of our phylogenetic analyses it obvi- pseudoparaphyllia. The leaf characters are similar ously belongs to the subfamily Rhynchostegio- to those of E. conostomus (except that basal cells ideae, but its position within the latter is not are larger and the costa ends in spine). The plants stable: Nona analysis resolve it more closely re- seem not to be pendent and are golden in color. lated to Rhynchostegium, while in POY topol- Axillary hairs were not observed. The capsule is ogies it is often resolved sister to Pseudosclero- cylindrical. The exostome teeth are dark red-brown, podium (cf. Figs. 1 & 2). Morphologically Aero- strongly loop-like curved at base (similarly to bryum is somewhat similar to the former genus Leskea polycarpa), and according to Brotherus in size, often whitish-argentate color, subcomplan- (we hesitated to destroy the last capsule with ate foliage in creeping shoots, smooth seta, etc. teeth) at their base cross- to obliquely-striolate. (we found no explanation of the statement of The endostome is dark-pellucid, with a low basal Norris & Koponen (1985), that the seta is rough Brachytheciaceae (Bryophyta) – a family of sibling genera 287 in New Guinean populations – we found it to the subfamily Rhynchostegioideae, but its posi- be smooth in all studied collections from that tion within the latter varies: it was found ei- area). Aerobryum has, however, many unique or ther in basal position within Rhynchostegio- rare character states in the Brachytheciaceae: the ideae clade, or sister to Aerobryum (cf. Figs. 1- seta is short, in combination with an inclined- 2). Pseudoscleropodium is unique in this sub- horizontal capsule; old parts of the plants are family in having a conic operculum. dark-brown to black; the growth is pendent in Species included: the main part of its geographical area; the lami- P. purum (Hedw.) Fleisch. nal cells have strongly incrassate walls, resulting 37. Bryoandersonia H. Robinson, Bryologist in some cases in a bone-shaped pattern [i. e. the 65: 137. 1962. Type: B. illecebra (Hedw.) H. longitudinal cell walls strongly incrassate and Robinson (= Hypnum illecebrum Hedw.). look like bones, due to two “pores” (=short thin- Plants large, rigid, green to golden-brownish. walled places) near each cell end. Stem prostrate or more commonly ascending to Species included: erect in dense tufts, with central strand, moder- A. speciosum (Dozy et Molk.) Dozy et Molk. ately densely, julaceously foliate, irregularly pin- 36. Pseudoscleropodium (Limpr.) Fleisch., nately branched or with sympodial shoots only, Musci Buitenzorg 4: 1542. 1923. – Scleropodium branch foliage similar to that in stem. Axillary sect. Pseudoscleropodium Limpr., Laubm. Deut- hairs 4-6-celled, uppermost cell obtuse, pale brown. sch. 3: 142. 1896. Type: Scleropodium purum Pseudoparaphyllia acute. Stem leaves rigidly and (Hedw.) Limpr. (= Hypnum purum Hedw., Pseu- loosely imbricate, broadly ovate-triangular to doscleropodium purum (Hedw.) Fleisch.). suborbicular, rather slightly plicate, strongly con- Plants large, pale sordid green to yellowish cave; margin serrate to serrulate, costa ending or whitish. Stem prostrate or ascending, with cen- above mid-leaf without spine; laminal cells elon- tral strand, densely, julaceously foliate, rather gate-flexuose, moderately thick-walled; basal cells regularly pinnately branched, branch foliage sim- with much incrassate cell walls, subalar cells ilar to that in stem. Axillary hairs 4-9-celled, subquadrate, larger than neighbouring cells, thick- uppermost cells pale brown. Pseudoparaphyllia walled, forming opaque, poorly delimited group. orbicular-triangular. Stem leaves imbricate, broad- Branch leaves similar to stem leaves. Dioicous. ly elliptic, broadly obtuse to rounded and api- Perichaetial leaves with reflexed acumina. Seta culate, slightly plicate, strongly concave; margin smooth. Capsule inclined to horizontal. Annulus subentire, costa ending in or shortly above mid- separating by fragments. Operculum rostrate. leaf without spine; laminal cells linear-flexuose, Peristome xerocastique, perfect. Spores small. Ca- moderately thick-walled; basal cells wider, much lyptra naked. Mesophytic to xeromesophytic. incrassate, alar cells subquadrate, more or less Distribution: temperate Eastern North America. inflated, thin- or thick-walled, forming poorly (a) This genus was segregated from Cirri- delimited group. Branch leaves smaller and nar- phyllum by Robinson (1962), without indica- rower than stem leaves. Dioicous. Perichaetial tion of its possible affinity. In Nona analysis it leaves with reflexed acumina. Seta smooth. Cap- was resolved in a very basal position in Rhyncho- sule inclined to horizontal. Annulus separating stegioideae, close to Pseudoscleropodium with a by fragments. Operculum conic. Peristome xero- high support, while POY analysis resolved it to castique, perfect. Spores small. Calyptra naked. the basal position in Rhynchostegielloideae (cf. Mesophytic to xeromesophytic. Figs. 1-2). It’s position seems to be highly sensi- Distribution: subcosmopolite (in some areas tive to small differences in sequence alignment. introduced?). Species included: (a) Buck (1980b) revised this monospecif- Bryoandersonia illecebra (Hedw.) H. Robinson ic genus, but did not discussed its affinity within 38. Eurhynchium B. S. G., Bryol. Eur. 5: 217. the family. Originally it was segregated from 1854. Type: E. longirostre B. S. G. = E. striatum Scleropodium. The position of the latter genus (Hedw.) Schimp. is fairly unresolved, but these two genera never Pancovia Neck. ex J.J. Kickx, Flore Cryptog- appeared closely related. In our analyses Pseu- amique des Flandres 1: 75, 91. 1867. nom. illeg. doscleropodium always appears as a member of 288 MICHAEL S. IGNATOV & SANNA HUTTUNEN

[originally this genus included species from E. fuegianum Card. – ?(probably Platyhyp- many genera of Brachytheciaceae and Isotheci- nidium) um; Lindberg (1879) used it as a subdivision E. hians (Hedw.) Sande Lac. – Oxyrrhynchium of Hypnum in the sense close to Eurhynchium E. laevisetum Geheeb – Oxyrrhynchium? of “Bryologia Europaea”]. E. meridionale (B. S. G.) De Not. – Plasteu- Plants large, rigid, light green to somewhat rhynchium whitish or brownish. Stem prostrate, with cen- E. oreganum (Sull.) Jaeg. – Kindbergia tral strand, moderately densely, julaceously foli- E. praelongum (Hedw.) B. S. G. – Kindbergia ate, irregularly pinnately branched and with E. pumilum (Wils.) Schimp. – Oxyrrhynchium numerous erect and arching sympodial shoots, E. remotifolium (Grev.) Jaeg. – Oxyrrhynch- branch foliage similar to that in stem. Axillary ium? hairs 2-4-celled, uppermost cell obtuse, hyaline E. savatieri Schimp. ex Besch. – Oxyr- to pale brown. Pseudoparaphyllia acute. Stem rhynchium leaves rigidly erect-spreading to spreading, broad- E. schleicheri (Hedw. f.) Milde – Oxyr- ly ovate-triangular to ovate-lanceolate, acute to rhynchium acuminate, plicate, concave in basal part; margin E. serricuspis C. Müll. – Kindbergia serrate to serrulate, costa ending far above mid- E. speciosum (Brid.) Jur. – ?(see discussion leaf, at 0.7-0.8(0.9) of leaf length, with spine; under Oxyrrhynchium) laminal cells elongate to linear, thin-walled; to- E. squarrifolium Broth. ex Iishiba – Kindbergia wards base shorter; alar cells differentiated or E. striatulum (Spruce) B. S. G. – Plasteu- only subalar cells differentiated, large, thin-walled, rhynchium forming indistinctly delimited group. Branch E. trichocladoides Ignatov – Oxyrrhynchium leaves somewhat narrower and smaller than stem 39. Plasteurhynchium Fleisch. ex Broth., Nat. leaves, otherwise similar. Dioicous or phyllodio- Pfl. ed. 2, 11: 212. 1925. Lectotype: P. striatulum icous. Perichaetial leaves with reflexed acumina. (Spruce) Fleisch. (= Hypnum striatulum Seta smooth. Capsule inclined to horizontal. Spruce), selected here. Annulus separating by fragments. Operculum Plants large, rigid, deep green to brownish- rostrate. Peristome xerocastique, perfect. Spores or olive-green. Stem prostrate, with central strand, small. Calyptra naked. Mesophytic. moderately densely, julaceously foliate, irregu- Distribution: Palearctic. larly pinnately branched and with arching sym- (a) The new definition of the genus makes podial shoots, branch foliage similar to that in it very homogeneous, and the two species in- stem. Axillary hairs 3-5-celled, uppermost cell cluded in it are sometimes treated as subspe- obtuse, pale brown. Pseudoparaphyllia acute. Stem cies. The closest relatives of this genus are Plast- leaves rigidly erect-spreading to erect-appressed, eurhynchium and Palamocladium (see com- ovate-triangular to ovate, broadly acuminate, more ments under the latter genus). or less plicate, concave in basal part; margin ser- Species included: rate to serrulate, costa ending in or above mid- E. angustirete (Broth.) T. Kop. leaf in spine; laminal cells elongate, moderately E. striatum (Hedw.) Schimp. thick-walled; basal cells much shorter and with Species transferred to other genera: incrassate walls, forming opaque area across leaf E. africanum Herz. – Kindbergia base or alar group of short cells delimited. Branch E. altaicum Ignatov – Kindbergia leaves somewhat narrower and smaller than stem E. arbuscula Broth. – Kindbergia leaves, otherwise similar. Dioicous. Perichaetial E. asperisetum (C. Müll.) E. B. Bartram – leaves with reflexed acumina. Seta smooth. Cap- Oxyrrhynchium sule inclined to horizontal. Annulus separating E. clinocarpum (Tayl.) Par. – Oxyrrhynchium by fragments. Operculum rostrate. Peristome xe- E. crassinervium (Tayl.) B. S. G. – Cirriphyllum rocastique, perfect. Spores small. Calyptra naked. E. dumosum (Mitt.) Jaeg. – Kindbergia Mesophytic to xeromesophytic. E. eustegium (Besch.) Dix. – Eurhynchiadelphus Distribution: Western Palearctic. E. flotowianum (Sendtn.) Karttunen – Sci- (a) Plasteurhynchium was originally placed urohypnum in the Lembophyllaceae to accommodate spe- Brachytheciaceae (Bryophyta) – a family of sibling genera 289 cies superficially rather similar to Isothecium wise similar. Dioicous or phyllodioicous. Per- (Brotherus, 1925). In recent publications it was ichaetial leaves with reflexed acumina. Seta included in the Brachytheciaceae as a genus smooth. Capsule inclined to erect. Annulus sep- (Ochyra & al., 1992), or more frequently its arating by fragments. Operculum rostrate. species were classified within Eurhynchium Peristome hygrocastique, slightly to moderately (Smith, 1978; Hedenäs, 1992, etc.). Molecular modified; exostome teeth cross-striolate below, and morphological data confirmed this latter red-brown or pale yellow; endostome with basal placement within the Brachytheciaceae and re- membrane 1/2–1/4 of its length. Spores small solved the genus as a close relative of Eurhynch- or large. Calyptra naked. Mesophytic to xer- ium s. str. and Palamocladium. Plasteurhynchi- omesophytic. um is somewhat similar to Eurhynchium stria- Distribution: pantropical, expanding to East- tum in its general appearance: robust plants ern North America and East Asia. with curved branches, plicate leaves, and a com- (a) Palamocladium was considered as a rel- plete peristome. The morphological similarity ative of Homalothecium since Lindberg (1879), with Palamocladium lays in the opaque basal who united them in Hypnum subgen. Pleuropus. areolation. However Plasteurhynchium differs This opinion was supported by Brotherus (1925), from both these genera in having a prorately Robinson (1962), Hofmann (1997) and almost toothed dorsal lamina, and a more pronounced all other authors. Our analysis does not confirm differentiation into primary and secondary this. Palamocladium has been found in all anal- stems. A phyllodioicous condition was never yses within the subfam. Rhynchostegioideae. The found in Plasteurhynchium, unlike in both Eu- most closely related genera are probably Eu- rhynchium and Palamocladium. rhynchium s. str. and Plasteurhynchium (Figs. 1, Species included: 2), however a support for this has not been ob- P. meridionale (B. S. G.) Fleisch. tained. Palamocladium differs from the latter P. striatulum (Spruce) Fleisch. genera by its narrow leaves and hygrocastique peristome that is more or less modified. Palam- 40. Palamocladium C. Müll., Flora 82: 465. ocladium is similar to Eurhynchium in its phyl- 1896. Type: Pleuropus fenestratus Griff. = lodioicous sexual condition and to Plasteu- Palamocladium leskeoides (Hook.) Britt. rhynchium in its opaque basal areolation. Pleuropus Griff., Calcutta J. Nat. Hist. 2: 473. (b) The genus was monographed by Hof- 1842. nom. illeg., later homonym. mann (1997). She accepted the pantropical P. Plants large, rigid, deep green to brownish- leskeoides in a very broad sense. Based on or olive-green. Stem prostrate to ascending, with morphology, it would probably be impossible weak central strand, moderately to very densely, to consider this very variable species in anoth- julaceously foliate, irregularly pinnately er way. However, a confirmation of its genetic branched, branch foliage similar to that in stem. unity based on molecular data is desirable. Axillary hairs 3-6-celled, uppermost cell ob- Species included: tuse, pale brown. Pseudoparaphyllia acute. Stem P. euchloron (Bruch ex C. Müll.) Wijk et leaves rigidly erect, lanceolate-triangular to Marg. ovate-lanceolate or lanceolate, gradually and P. leskeoides (Hook.) Britt. broadly acuminate, at base slightly auriculate, P. wilkesianum (Sull.) C. Müll. deeply plicate or in slender plants smooth; margin sharply serrate, with teeth partly re- 41. Scorpiurium Schimp., Syn. Musc. Europ. curved, costa ending above mid-leaf or almost 2: 855. 1876. Type: S. rivulare Schimp. = S. de- reaching apex, without spine; laminal cells flexifolium (Solms) Fleisch. et Loeske. elongate, relatively short (in comparison with Plants medium-sized to moderately small, large size of leaf), moderately thick-walled; bas- somewhat rigid, green to yellowish or brownish al cells much shorter and wider, with incras- green. Stem prostrate to ascending, with central sate cell walls; alar cells numerous, subquad- strand, densely foliate, irregularly pinnately rate, forming extensive opaque area, sometimes branched, branch curved, their foliage similar to throughout auricules. Branch leaves somewhat that in stem, sometimes branches attenuate. Ax- narrower and smaller than stem leaves, other- illary hairs 4-6-celled, hyaline or brownish. 290 MICHAEL S. IGNATOV & SANNA HUTTUNEN

Pseudoparaphyllia triangular. Stem leaves ap- EPILOGUE pressed when dry, erect-spreading when moist, The current grouping of genera of the broadly ovate to ovate-lanceolate, acute or very Brachytheciaceae differs quite a lot from previ- broadly acuminate, more or less concave, not pli- ous ones. The greatest changes are suggested cate; margin bluntly serrate to serrulate; costa for the genera Eurhynchium, in its traditional ending above mid-leaf with spine, and often with sense, Cirriphyllum, Platyhypnidium, and may- several teeth near its end (especially in branch be also Bryhnia. All bryologists who dealt with leaves); laminal cells rhombic-ovate to rhom- Eurhynchium in the recent past included in it bic/elongate, thick-walled; at base cells shorter, also Oxyrrhynchium, sometimes specifically un- forming opaque area across base, alar area sub- derlining that they are not distinct (Takaki, quadrate to oblate, forming extensive opaque area 1956; Buck, 1988; Koponen, 1987; Robinson, extending up along margin. Branch leaves smaller 1962; Hedenäs, 1992; Ignatov, 1998; Matteri & than stem leaves, otherwise similar. Dioicous. Ochyra, 1989; Crum & Anderson, 1981). Our Perichaetial leaves with reflexed, relatively short inferences based on molecular and morpho- acumina. Seta long, smooth. Capsule suberect to logical data strongly suggest that these two inclined, shortly ovate to cylindrical. Annulus genera are not closely related, and following separating by fragments. Operculum rostrate. our systematic concept, they belong to differ- Peristome xerocastique, perfect. Spores small. ent subfamilies. Platyhypnidium was re-estab- Calyptra naked. Mesophytic to xeromesophytic. lished recently for the small group of aquatic Distribution: Western Palearctic. species. Its homogeneity was underlined by Ig- (a) This genus was united with Eurhynchi- natov & al. (1999), and Hedenäs (2002). Phy- um s. l. by some authors (Mönkemeyer, 1927, etc.), logenetic inferrences suggest that Platyhypnid- whereas Brotherus (1925) placed Scorpiurium ium is polyphyletic with species scattered be- and Eurhynchium at opposite ends in his ac- tween different genera of different subfamilies. count of genera of the Brachytheciaceae. Broth- The situation with Bryhnia novae-angliae is even erus seems to have considered it especially iso- more contrasting with the traditional views. Dif- lated, though he did not express this explicitly. ferent populations of this “species” were not Our studies support this idea. Scorpiurium (with resolved as a monophyletic entity, and may Bryoandersonia) appeared in the analyses in belong to more than one genus. basal position in the Rhynchostegioideae (Fig. 1, Seeing these sometimes confusing groupings, 2), the most basal clade of the family. should we trust these results based mainly on (b) A key for all three European species molecular data? After careful evaluation of the was published by Matracci (2001). results, one can notice that at least some of the Species included: newly revealed groups quite nicely correspond S. circinatum (Brid.) Fleisch. et Loeske to some morphological characters, which were S. deflexifolium (Solms) Fleisch. et Loeske probably under-evaluated by earlier authors. S. sendtneri (Schimp.) Fleisch. For example, Oxyrrhynchium species often grow Species not confirmed in the genus: in loose mats (but in dense growth they become S. cucullatum (Hampe) Hedenäs. Many mor- subdendroid), with subcomplanate branches in phological character states of this Australian loose growth, the branch leaves have a tendency endemic species are similar to those of three to be elliptic, and in loosely foliate parts of the Mediterranean species of Scorpiurium, but they branches they are often twisted from the mid- also fit Flabellidium, Helicodontium, Rhyncho- dle. Such characters are difficult to use due to stegiella (see note “a” under that genus), and their rather inexact definition, and thus they we expect it must be in one of these genera. were not taken into consideration or involved Especially, the rough seta in S. cucullatum is in earlier classifications. according to our phylogenetic analysis in con- Although species of Platyhypnidium resemble flict with a placement in Scorpiurium, since no each other in leaf characters, the genus may be one species of Rhynchostegioideae has this char- polyphyletic: our analysis suggests that P. austri- acter state. Thus, we doubt its generic place- num may belong to the Rhynchostegielloideae ment within this subfamily. whereas P. riparioides seems nested within the Brachytheciaceae (Bryophyta) – a family of sibling genera 291

Rhynchostegioideae. Note that P. austrinum has a tional classification to a certain extend. Then, rough seta, a state never present in the Rhyncho- after finding a new and presumable better stegioideae. This forces us to consider that their grouping, we are coming to updated the circum- position in different subfamilies is not an artifact scription of subfamilies and genera. However, of the analysis. The aquatic environment obvious- do these new circumscriptions help to attribute ly has a strong effect on morphology (especially the problematic species in certain group of the leaf morphology), and for this reason the taxo- Brachytheciaceae? We found it is more or less nomic position of aquatic taxa must be evaluated easy for three big genera – in Rhynchostegium, with the extreme caution. It is worth to remember Homalothecium, and Brachythecium, if we ac- that Loeske (1911) included in the genus Platy- cept the latter in a relatively broad sense, as hypnum (later transformed to Platyhypnidium) suggested here. Note, however, that attributing also Hygrohypnum dilatatum, H. molle, and H. al- a species to one of four main infrageneric units pinum. These latter taxa were traditionally segre- of Brachythecium, groups {around Brachytheci- gated in Hygrohypnum, Amblystegiaceae, but ac- um rivulare}, {B. tommasinii-geheebii-complana- cording to Vanderpoorten & al. (2002), Hygrohyp- tum-?cirrosum}, {around B. salebrosum}, and {B. num is still more severely polyphyletic than Platy- buchananii-ruderale-laetum} is not always easy. hypnidium, as its species are suggested to belong Sometimes we hesitate to refer species to to at least three different families. Brachythecium vs. Sciurohypnum based on the Another morphological character induced morphological description only. In other gen- apparently by aquatic habitat is a bistratose era the problems are much worse. Cirriphyllum lamina. It was considered as unique character, seems to totally lack morphological synapomor- having high taxonomic value (Ochyra, 1985). phies. Genera with unusual combinations of However later Ochyra & Vanderpoorten (1999) character states, like Mandoniella, Juratzkaeel- found it had independently evolved within la, and Schimperella are difficult to link with Platyhypnidium, and Stech & Frahm (2001) certainty to one or another subfamily. Plants of expanded this list further. the group of Helicodontium, Scorpiurium, Fla- Before this study, a rostrate operculum was bellidium, and the group of Rhynchostegiella not known to occur in Brachythecium s. str., in species with short and thick-walled cells are fact, it was considered incompatible with the very similar – but note, that they are in two concept of the genus. The nested position of subfamilies! The enumeration of cases where “Cirriphyllum tommasinii” in Brachythecium morphology is misleading in Brachytheciaceae and that of “Cirriphyllum flotowianum” in can be continued further. But we believe that Sciurohypnum (earlier included in Brachyth- the reader who knows this family is convinced ecium), the latter with a high support, overturn already, that it is not always possible to refer a this belief. When the value of this character is species with certainty to any specific genus. In reconsidered, one might notice the overall simi- this situation we can probably speak about “sib- larity between Sciurohypnum flotowianum and ling genera” using a parallel from “sibling spe- S. populeum in appearance and in some leaf cies”. Actually, a similar situation started to characters, which appear to be more important become common at the genus level after molec- for delimiting this group than the shape of the ular studies became a widely used tool of sys- operculum. In both cases trends towards a ros- tematics. The situation at the level of family is trate operculum were associated with a consid- even more dramatic: cf. Vanderpoorten & al. erably decreasing of length:width ratio of leaf (2002), who suggested a new classification of laminal cells. If we consider that these two char- Amblystegiaceae with “morphological synapo- acter states often correlate (cooccur) with each morphies unknown” for the family, some sub- other (especially in epiphytic groups), then the families and tribes. new placements of these former Cirriphyllum So, are we far from Bridel, who circumscribed species do not look so odd. taxa, giving no key to identify them? After that, This and similar minute parallels convince the authors of “Bryologia Europaea” did their us, that the obtained results based largely on best in selecting “recognizable” characters which molecular data are able to improve the tradi- could be used for identification of taxa. Some 292 MICHAEL S. IGNATOV & SANNA HUTTUNEN groups have been delimited, so that we are able trachypodium n=10. [Brachythecium appleyardi- to use them in practical work now, but for ex- ae n=11] ample in the case of the rostrate operculum, this Eurhynchiastrum pulchellum n=10, 11 (more key character “helped” to place in the subge- rarely 7, 8+X/Y, 12, 20). nus Oxyrrhynchium six species, which accord- Brachythecioideae ing to the present data belong to five genera of Kindbergia arbuscula n=10, 11, oregana n=8, three subfamilies. Note also that Bridel put praelonga n=9 (more rarely 7, 8, 9+1 acc., 9+2 Hypnum velutinum (cf. our Brachytheciastrum), acc., 9+4 acc., 10, 10+m, 11). H. reflexum, and H. starkei (cf. our Sciurohyp- Bryhnia hultenii, noesica, novae-angliae, ten- num) in a separate section, different from the errima, trichomitria n=11 (more rarely 10), de- rest of Brachythecium. Bruch & al. (1851-55) curvans, tokubuchii, n=10; tenerrima n=8. combined them with the rest of Brachytheci- Eurhynchiadelphus eustegium n=11. um based on the key character state of conic Myuroclada maximowiczii n=10, 11. operculum, and now we are returning back to Brachythecium rivulare-group: frigidum Bridel And we are almost sure, that further n=6; rivulare n=6 (much more rarely 11, 12, 16), authors will find no less drastic errors in the rutabulum n=12 (much more rarely 5, 6, 10, 10+m, present paper too. 13, 20, 22, 24). How can we then classify a species with an Brachythecium [basal species with rough unusual combination of characters, if it is known seta]: auriculatum n=11; coreanum n=11; ge- only by old and imperfect collections? It seems heebii n=10; tommasinii n=10, 11; like the best (if not the only) way is to se- Brachythecium salebrosum-group: albicans quence the living beings with the maximally n=7 (more rarely 6, 9), austroglareosum n=13, similar phenotypes from the maximally close austrosalebrosum n=10, 11, 13, 20; campestre n=16 geographical area and then re-evaluate the (more rarely 17, 11, 12), glareosum n=9 (more morphological characters, testing their signifi- rarely 6, 10), mildeanum n=13, 14; salebrosum cance in this particular group, and only then n=13 (much more rarely 10+m, 12+m, 13+m); make taxonomic decisions. At least in the turgidum n=14. Brachytheciaceae this might help. Brachythecium stereopoma-group (sensu APPENDIX 1. McFarland, 1988): buchananii n=10 (also 9+m, 10+m, 11, 14, 20+2m), garovaglioides n=10, ka- CHROMOSOME NUMBERS IN mounense n=10, 10+m, kuroishicum n=10. BRACHYTHECIACEAE Sciurohypnum flotowianum n=11; glaciale The increasing of information on the chro- n=8, oedipodium n=20 (much more rarely 7, 10, mosome numbers lead to slightly controversial 11), populeum n=10 (more rarely 9, 10+m, 11), results, making the whole amount of data diffi- plumosum n=10 (more rarely 9+m, 10+m, 11, cult to discuss. However, if species with vari- 11+m), reflexum n=10 (more rarely 11, 12, 20), able counts from few specimens are omitted, the starkei n=10 (more rarely 20). catalogue of Fritsch (1991) can be generalized Rhynchostegielloideae as follow [with addition of data of Uniyal, Scleropodium obtusifolium n=10+m, 22; 1997]: touretii n=11. Homalothecioideae Rhynchostegiella curviseta n=11; tenella Homalothecium aureum n=10 (more rarely n=11, 22; R. tenerrifae n=11. 10+2m); fulgescens n=10+m, 16; laevisetum Cirriphyllum crassinervium n=11; pilifer- n=11; lutescens n=10 (more rarely 8, 10+m, 10+B, um n=11. 10+1 acc., 11, 12, 14, 24); philippeanum n=8, Clasmatodon parvulus n=11, 10+m. 8+2m, 10, 16; sericeum n=8, 10 (more rarely Oxyrrhynchium hians n=7 (very rare 7+2 8+X/Y, 9, 10+m, 10+B, 10+1acc., 10+3 acc., 11, acc., 11), pumilum n=10, 10+m; savatieri n=8; 11+m, 11+2m). schleicheri n=11; speciosum n=15; vagans n=10. Brachytheciastrum collinum n=10+m; par- Okamuraea brachydiction n=11, hakkonien- adoxum n=22; velutinum n=10 (much more sis n=11. rarely 10+m, 10+1, 10+2 acc., 11, 12, 12+m, 18), Homalotheciella subcapillata 10+m. Brachytheciaceae (Bryophyta) – a family of sibling genera 293

Rhynchostegioideae chromosomes. It can be hypothesized that S. Pseudoscleropodium purum n=11 (Europe), hylotapetum, the largest species of the genus, more rarely 7 (Japan). is also a polyploid. Chromosome numbers in Plasteurhynchium meridionale n=11, stiatu- Brachythecium are more or less corresponding lum n=11. to the main groups of species: (1) the B. rivu- Palamocladium leskeoides n=11. lare-group has n=6 and 12; (2) the B. stereo- Eurhynchium angustirete n=6, 11 (more rare- poma-group sensu McFarland has n=10 [how- ly 6, 12, 12+1 acc.), striatum n=11 (more rarely ever, many counts are reported so far for only 6, 12). one species, B. buchananii], and (3) B. salebro- Rhynchostegium cataractile n=10, 22; con- sum group is very variable (n= 7, 10, 13, 14, fertum n=10, 11 (more rarely 10+m, 13+m, 12); 16); note however, that the number is more or duthiei n=10, 10+m; herbaceum n=10+m, 11; less stable within the individual species; (4) inclinatum n=22; laxatum n=20+m, 22; mega- only two species have n=10, B. tommasinii and politanum 10, 11 (more rarely 11+1m); murale B. geheebii (both with a rough seta and in n=10, 11 (more rarely 10+m, 11+m); pallidifoli- some analyses (cf. Fig. 1), they are closely re- um n=11; rotundifolium n=10+m, 11; serrula- lated, though without support); B. auricula- tum n=9, 9+m, 11; tenuifolium n=22. tum and B. coreanum, which we consider close Platyhypnidium riparioides n=11 (more to the latter group, have n=11 (they also have rarely 7, 8, 10, 12, 20); muelleri n=9. rough seta). Bryoandersonia illecebra n=12. Hence some peculiar groups in Brachytheci- Scorpiurium circinatum n=11. aceae also have deviation in chromosome counts (i. e. others than n=10-11). Thus, the most common numbers in the Rhyn- ACKNOWLEDGEMENTS chostegielloideae and Rhynchostegioideae are We are greatly indebted to Jaakko Hyvönen n=10 and 11. The only genus that is strongly for help in various ways, especially in provid- different is Oxyrrhynchium, where n=7, 8, and ing laboratory facilities, to Lars Hedenäs for 15 are stable and found in many counts for stimulating criticism on the msc. and many cor- more than the half species. Note that 7 is known rections, to Bernard Goffinet for linguistic cor- as the most common number in only one more rections as well as critical comments, to Ryszard species, Brachythecium albicans, while 15 is Ochyra for important nomenclatural and tax- unique among the Brachytheciaceae. onomical correction, to Pekka Isoviita for con- In the Homalothecioideae n=10 and 11 are sultation on puzzling nomenclatural problems, also common, but some species have n=8 and Elena Ignatova for a lot of technical work with more rarely 9. the manuscript, to Timo Koponen for arrange- The Brachythecioideae includes two large ment of visits of the former author to Helsinki genera, which differ in chromosome numbers. Herbarium and for the initiating of this work, Sciurohypnum has mostly n=10; only two spe- to Curators of BM, COLO, H, MHA, NY, S kindly cies have n=8 and 11, both based on a single send us some specimens for sequencing. The work count, and also S. oedipodium has mostly n=20. has been partly supported by grant 200A/17 The polyploid nature of the latter species can of the Finnish Academy of Science and by grants be supported from morphology, as it is much 02-04-48358 and 02-04-50035 from the Russian larger than other related species studied for Foundation for Basic Research.

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