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Brachytheciaceae (Bryophyta) – a Family of Sibling Genera Brachytheciaceae (Bryophyta) – Семейство Родов–Бли

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Brachytheciaceae (Bryophyta) – a Family of Sibling Genera Brachytheciaceae (Bryophyta) – Семейство Родов–Бли Arctoa (2002) 11: 245-296 BRACHYTHECIACEAE (BRYOPHYTA) A FAMILY OF SIBLING GENERA BRACHYTHECIACEAE (BRYOPHYTA) ÑÅÌÅÉÑÒÂÎ ÐÎÄÎÂÁËÈÇÍÅÖΠMICHAEL 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
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