Cephaloziaceae (Marchantiophyta) of the Sino-Himalayan region

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1 Nova Hedwigia Vol. 93 issue 1 2, Stuttgart, August 2011 Article Cephaloziaceae (Marchantiophyta) of the Sino-Himalayan region JiÍR VáÁa 1 and David G. Long 2* 1 Department of Botany, Charles University, Benátská 2, Praha 2, Czech Republic 2 Royal Botanic Garden, Edinburgh EH3 5LR, UK Váňa, J. & D.G. Long (2011): Cephaloziaceae (Marchantiophyta) of the Sino-Himalayan region. Nova Hedwigia 93: Abstract: 10 genera and 29 species of Cephaloziaceae are reported from the Sino-Himalayan region. Several published species of Cephalozia are considered to be of doubtful distinctness and require further study. Sphenolobus langkyrdumii Ajit P.Singh et V.Nath is relegated to the synonymy of Metahygrobiella albula (Steph.) Grolle. Key words: Cephaloziaceae, Alobiellopsis, Cephalozia, Cladopodiella, Hygrobiella, Iwatsukia, Metahygrobiella, Nowellia, Odontoschisma, Pleurocladula, Schiffneria. Introduction This paper is the third in a series of checklists of selected families of Sino-Himalayan liverworts, initiated under the European Community s SYNTHESYS programme. It follows the first paper which covered the Jungermanniaceae (Váňa & Long 2009) and the second one dealing with Gymnomitriaceae (Váňa & Long 2010). In contrast to those two families, however, there is little controversy surrounding the delimitation of Cephaloziaceae as a family and indeed of its component genera, although its position in the overall liverwort phylogenetic tree has changed in recent times (Crandall- Stotler et al. 2009, Frey & Stech 2009); it is now placed in order Jungermanniales suborder Cephaloziineae, along with the families Adelanthaceae, Cephaloziellaceae and Scapaniaceae s.l. (including Lophoziaceae and Anastrophyllaceae) (Roo et al. 2007, Söderström et al. 2010). However, in contrast to the rather stable generic picture, species concepts within the largest genus of the family, Cephalozia, remain in a state of considerable uncertainty. This is considered in some detail below and an interim solution proposed along with the suggestion that a combined molecular and *Corresponding author d.long@rbge.ac.uk 2011 J. Cramer in Gebr. Borntraeger Verlagsbuchhandlung, Stuttgart, Germany DOI: / /2011/ /2011/ $

2 morphological study of the genus is essential for progress to achieve a stable and well-supported species concept. The most problematic groups within Cephalozia are treated under the aggregate names 'Cephalozia bicuspidata complex' and the 'Cephalozia lunulifolia complex'. The checklist The checklist is in part a compilation of published data from literature available to the authors. Almost all of the voucher specimens in European herbaria have been checked by the senior author. The study-area (Váňa & Long 2009) is the biogeographic unit informally known as the 'Sino-Himalayan region', comprising the Pakistan Himalaya, Indian Himalaya (Jammu & Kashmir, Himachal Pradesh, Uttaranchal, Sikkim, Darjeeling District of West Bengal, Assam, Meghalaya, Manipur and Arunachal Pradesh), Nepal, Bhutan and western China (Yunnan, Sichuan and Xizang (Tibet)). A number of corrections to the geography of published reports have been made, for example all records from 'Uttar Pradesh' have been reassigned to the modern state of Uttaranchal. Many old records from 'Sikkim' are ambiguous as Sikkim in the past was often used to include the Darjeeling District of West Bengal e.g. by Mitten ( ). Such records, e.g. for Kurseong, Senchal etc. have been reassigned to West Bengal, though others, e.g. for the Singalela Ridge, remain ambiguous. Records from the Hengduan Shan in China (Liu 2000) Cephalozia lunulifolia (Dumort.) Dumort. and C. macounii (Austin) Austin are ambiguous and are not listed as they may refer to Yunnan, Sichuan or Xizang Provinces. Inadequately-localised reports, e.g., for 'Himalaya', 'East India' and 'East Indies' have not been included. Author abbreviations follow Brummitt & Powell (1992) and IPNI (International Plant Names Index, for more recent authors. ALOBIELLOPSIS R.M.Schust. Enumeration of Cephaloziaceae taxa TYPE SPECIES: Alobiellopsis acroscypha (Spruce) R.M.Schust. Alobiellopsis parvifolia (Steph.) R.M.Schust. CHINA, SICHUAN (Gao 2003) CEPHALOZIA (Dumort.) Dumort. TYPE SPECIES: Cephalozia bicuspidata (L.) Dumort. A. The 'Cephalozia bicuspidata complex' The section Cephalozia of the genus Cephalozia is the most difficult and taxonomically problematic group of species in the genus; fortunately the type species is the most widespread and familiar member of the genus. Some species placed in 154

3 this section were first described from sterile or otherwise inadequate material which has hampered unequivocal interpretation of the names; in addition all the species of this complex exhibit phenotypic plasticity under different ecological conditions, at least in morphological characters of the gametophyte. Those morphological characters used by some authors for the separation of sterile taxa (stem anatomy, remoteness of leaves, leaf shape, shape of leaf lobes and their apices, cell size) show great variability under different ecological conditions. Chromosome numbers, which have sometimes been used (in conjunction with cell size) to distinguish some critical pairs of holarctic taxa (C. bicuspidata (L.) Dumort./ C. lammersiana (Huebener) Spruce and C. hamatiloba Steph./C. otaruensis Steph.) do not fully correlate with cell size, and the chromosome races C. lammersiana (Europe and North America) and C. otaruensis (East Asia) have for that reason been variously reduced to subspecies, varieties or even regarded as synonyms in different treatments. Váňa (1988) accepted only two characters for reliable separation of the species of this complex: sexuality (autoicous vs. dioicous) and dentition of the perianth mouth (crenulate vs. ciliate). These characters differentiate C. bicuspidata (along with C. ambigua C.Massal., sometimes treated as a separate species on the basis of small cells, characteristic ecology, etc.), from other accepted species in this complex. Cephalozia bicuspidata is autoicous, with the perianth mouth crenulate or sometimes slightly and irregularly dentate (with teeth of 1 2 superposed cells). By this strict definition of C. bicuspidata, the closely related taxa C. cavifolia S.W.Arnell (type from Uganda), C. grandifolia Steph. (type from Bolivia), C. randii S.W.Arnell (type from Marion I.), C. robusta Steph. (type from South Africa), C. skottsbergii Steph. (type from South Georgia), C. vallis-gratiae Gottsche ex Steph. (type from South Africa) and C. vulcanicola Steph. (type from Rwanda) and probably also C. austrigena R.M.Schust. et J.J.Engel (type from New Zealand) should all be treated as synonymous. The populations from tropical South and Central America with narrow leaf lobes (3 4 cells wide at the base) ending in 2 3( 4) superposed cells, which are autoicous and with perianth mouth mostly ciliate (C. bischlerae Fulford, C. dussii Fulford, C. patentiloba Steph. and C. venezuelana R.M.Schust.), have been placed under C. crossii Spruce. Finally, the populations described as dioicous from South-east Asia with the perianth mouth ciliate have been treated as C. hamatiloba, and those with the perianth mouth crenulate or irregularly dentate have been treated under the name C. darjeelingensis Udar et D.Kumar. Additional detailed comparative studies by the first author based on numerous collections, have shown that even the dentition of the perianth mouth may not be a consistently reliable character. The cilia of the perianth mouth in South and Central American populations (which seem to be consistently well-defined by the leaf and lobe form), can be very long (7 8 cells) as in C. crossii, somewhat shorter (4 5 cells long), as in C. venezuelana, short-ciliate, as in C. bischlerae and "irregularly crenulate and setulose", as described in C. dussii (the type specimen of C. patentiloba is sterile). Because of this range of variation, Schuster (2002) did not accept Váňa s 155

4 (1988) concept of one broad species for this complex and Schuster (l.c.) retained "with considerable reservations, some of the narrower species limits advocated by Fulford [(1968)]". A comparable problem exists in South-east Asia for populations described as dioicous, where the reliability of perianth dentition is again called into question. East Asiatic populations treated as C. hamatiloba (= C. otaruensis) by Váňa (1988) have the perianth mouth shortly ciliate, but in C. siamensis N.Kitag. the perianth mouth is "laciniate-ciliate" (Kitagawa 1969). In C. darjeelingensis Udar et D.Kumar the perianth mouth was described as "irregularly lobate, margin more or less smooth" (Udar & Kumar 1976), and this statement was the basis for keeping C. darjeelingensis as a separate species in contrast to C. hamatiloba by Váňa (1988), although it agrees well in nearly all other characteristics with C. hamatiloba and with C. siamensis (treated as separate species, subspecies or synonym of C. hamatiloba by various authors). Schuster (1974), who collected dioicous plants of this complex in North and South Carolina, also considered that these plants were identical to Japanese ones, and therefore treated these populations as C. bicuspidata subsp. otaruensis (Steph.) S.Hatt. (= C. hamatiloba). Three additional facts relevant to these species concepts should be noted: 1. Some species were described (and mostly remain known) only in the sterile condition. This fact is a major problem for several taxa, e.g., C. patentiloba (described from Ecuador), the type specimen of C. neesiana Steph. (described from Java), C. skottsbergii (described from South Georgia), the type specimen of C. vallisgratiae (described from South Africa) and especially the type specimens of C. pandei Udar et D.Kumar and C. kashyapii Udar (both from India). 2. Perianths used for descriptions by various authors are sometimes immature or poorly-developed. In such perianths the cilia of the mouth are much shorter, sometimes barely differing from other cells of the mouth which then appear to be only dentate or crenulate. This was already pointed out, e.g. by Váňa (1988), for the plants named as C. robusta. 3. Sexuality (or sexuality as described in the literature by some authors) of plants is another major problem. For example, C. lammersiana has on occasion been wrongly described as dioicous by some authors because of the fact "that the gynoecial, postical branches are often so elongated that any connection to the leading, usually androecial shoots is difficult to ascertain" (Schuster 1974). Similarly, C. dussii was originally described as dioicous by Fulford (1968), probably in error, as stated Váňa (1988). The same concerns apply to C. patagonica Fulford (described as 'dioicous' in the same study but under a different section); however Engel (1978) already corrected this error in the description under the clearly autoicous C. pleniceps (Austin) Lindb. with which C. patagonica is now synonymised. Furthermore, the same erroneous description as 'dioicous' has been applied to C. cavifolia (because of one shoot with "female organ apical, only young observed"), C. vulcanicola (the type material contains only sterile fragments and androecial shoots) and also material named as 156

5 C. vallis-gratiae described by Arnell (1963) but already doubted by Schuster (1974). All the above species must now be treated as synonymous with the autoicous C. bicuspidata. Finally, C. robusta, described originally as monoicous, was incorrectly described by Arnell (1963) as dioicous (examination of the specimens seen by Arnell shows they are clearly autoicous and belonging to C. bicuspidata, cf. Váňa (1988)). On occasion some of the above mentioned species were described as heteroicous for the same reasons. In fact, the single sporadically heteroicous (some shoots being paroicous) species of Cephalozia, C. chilensis J.J.Engel et R.M.Schust. (= C. heteroica R.M.Schust. et J.J.Engel, nom. illeg.), should perhaps be placed in Metahygrobiella R.M.Schust. (Schuster 1974). In summary, a critical taxonomic study of this group incorporating molecular methods is highly desirable. It could even be the case that only one species, C. bicuspidata, exists, albeit perhaps with some geographically-defined subspecies. In the enumeration below we have listed the Indian species separately and append our comments relating to individual taxa. However, it must be noted that up to now Indian authors have mostly disagreed with uniting these taxa and have consistently retained all such species individually; in our opinion they have not given due consideration to the often inadequate material and the often obscure morphological differences by which they were characterized. Cephalozia bicuspidata (L.) Dumort. INDIA, SIKKIM (Bapna & Kachroo 2000); CHINA, XIZANG (TIBET) (Gao & Zhang 1981, Piippo 1990, Gao 2003), YUNNAN (Gao & Cao 2000, Gao 2003). NOTE: the earlier report of Mitten ( ) represents C. hamatiloba. Cephalozia darjeelingensis Udar et D.Kumar INDIA, WEST BENGAL: Darjeeling (Udar & Kumar 1976, Váňa 1988, Parihar et al. 1994, Bapna & Kachroo 2000, Singh & Nath 2007), MEGHALAYA: East Khasia Hills, West Khasia Hills, Jaintia Hills (Singh & Nath 2007); BHUTAN (Long & Grolle 1990, Bapna & Kachroo 2000, Singh & Singh 2007). NOTE: Accepted as a distinct species by Váňa (1988) on the basis of different perianth mouth than in the East Asiatic C. hamatiloba (the perianth is more similar to C. bicuspidata, which is autoicous). It may be possible that this species (and many specimens of the genus Cephalozia collected in India, too) is conspecific with the concept of C. hamatiloba. If this opinion is confirmed, C. hamatiloba will be the only species of the sect. Cephalozia in South-east Asia, following the concept of Váňa (1988). Cephalozia hamatiloba Steph. INDIA, UTTARANCHAL: Kumaon, Kabir Khan, det. Stephani as C. bicuspidata (herb. Levier, BM; here as new record from the area), WEST BENGAL: Kurseong, coll. Decoly & Schaul, det. Schiffner as C. blepharostoma in sched., det. Váňa (herb. Levier 649, BM; here as new record from the area), MEGHALAYA: Khasia Hills (JDH et TT - cf. Mitten as C. bicuspidata); BHUTAN (Váňa 1993, Bapna & Kachroo 2000, Long 2005, Singh & Singh 2007); NEPAL (Váňa 1993, Bapna & Kachroo 2000, Long 2005, Singh & Singh 2007); CHINA, YUNNAN (Piippo et al. 1998, Gao & Cao 2000 as C. otaruensis, Gao 2003 as C. otaruensis). 157

6 Cephalozia hamatiloba subsp. siamensis (N.Kitag.) Váňa SYN: Cephalozia siamensis N.Kitag. BHUTAN (Long & Grolle 1990, Bapna & Kachroo 2000). NOTE: Acording to Kitagawa (1969) C. hamatiloba (as C. otaruensis) differs from C. siamensis in the nearly orbicular or broadly ovate leaves as well as the shortly ciliate perianth mouth, and also in the lack of gemmiparous shoots which are a frequent occurrence in the latter. In fact, as most plants are sterile, the frequent occurrence of gemmiparous, microphyllous shoots is almost always the only diagnostic feature of the subspecies; perianths are very rare. Cephalozia kashyapii Udar [? = C. darjeelingensis Udar et D.Kumar] SYN.: C. laxifolia Udar et D.Kumar, nom. illeg., non C. laxifolia (Hook.) Lindb. [= Hygrobiella laxifolia (Hook.) Spruce] INDIA, W BENGAL: Darjeeling (Udar & Kumar 1976, Udar 1978, Váňa 1988, Parihar et al. 1994, Bapna & Kachroo 2000, Singh & Nath 2007), MEGHALAYA: East Khasia Hills (Singh & Nath 2007). NOTE: According to Váňa (1988) C. kashyapii is identical with C. darjeelingensis. The type specimen of C. kashyapii is completely sterile, and almost certainly represents a juvenile phase, not fully developed C. darjeelingensis. However, Singh & Singh (2007) advocated the separation of these taxa and repeated the differences given in the original description (more cells in the stem t.s., more closely placed leaves and presence of gemmae in C. darjeelingensis, as well as different ecology). Cephalozia pandei Udar et D.Kumar [? = C. darjeelingensis Udar et D.Kumar] INDIA, W BENGAL: Darjeeling (Udar & Kumar 1976, Váňa 1988, Parihar et al. 1994, Bapna & Kachroo 2000, Singh & Nath 2007), MEGHALAYA: East Khasia Hills (Singh et al. 2003, Singh & Nath 2007). NOTE: According to Váňa (1988) this taxon is probably a modification of C. darjeelingensis. Plants (type specimen) are fully sterile, and probably represent a poorly developed form of C. darjeelingensis. Singh & Nath (2003) collected in the East Khasia Hills fertile plants of a Cephalozia which they determined as C. pandei (following examination of the type of C. pandei). They summarized the features of their specimen in comparison with the original description of C. darjeelingensis and concluded that both taxa are different, an opinion repeated by Singh & Singh (2007). Cephalozia udarii D.Kumar [? = C. bicuspidata (L.) Dumort.] INDIA, UTTARANCHAL: Valley of Flowers (Kumar 1987, Parihar et al. 1994, Bapna & Kachroo 2000). NOTE: Kumar (1987) considered this species to be closely related to C. gollanii Steph. According to the description and drawings of C. udarii, it resembles the holarctic C. bicuspidata, particularly in being described as monoicous. However, C. bicuspidata has been disregarded by nearly all Indian authors (see above) who invariably used only species described from India for comparison. 158

7 B. Cephalozia lunulifolia complex Cephalozia affinis Lindb. ex Steph. BHUTAN (Váňa 1988, Long & Grolle 1990); NEPAL (Grolle 1966 as C. cf. media, Váňa 1988, Long & Grolle 1990, Bapna & Kachroo 2000). NOTE: This species is separated from C. lunulifolia only on the basis of sexuality (autoicous plants); the problems in the taxonomy of this species were summarized by Schuster (1974). Bearing in mind the situation described above in the C. bicuspidata complex, only one species of this complex, C. lunulifolia, should perhaps be recognised in the study-area. Cephalozia lunulifolia (Dumort.) Dumort. CHINA, YUNNAN (Nicholson 1930, Piippo 1990, Piippo et al. 1998, Gao & Cao 2000, Gao 2003). Cephalozia schusteri Sushil K.Singh et D.K.Singh [? = C. lunulifolia (Dumort.) Dumort.] INDIA, HIMACHAL PRADESH: Kullu (Singh & Singh 2007) NOTE: Very similar to C. lunulifolia of which it is probably only a modification. The characters used by the authors to distinguish the two species (narrower shoots, slightly smaller leaves, angulate or pyriform, rarely stellate gemmae and crenulate-dentate perianth mouth in C. lunulifolia) are based on the comparison of descriptions of both taxa and are characters prone to environmental modification. C. Other species Cephalozia catenulata (Huebener) Lindb. Cephalozia connivens (Dicks.) Lindb. BHUTAN (Long & Grolle 1990, Bapna & Kachroo 2000); Cephalozia gollanii Steph. INDIA, UTTARANCHAL: Garhwal (= Pauri) (Stephani , Kashyap & Chopra 1932, Chopra 1943, Parihar , Kitagawa 1969, Kachroo 1970, Long & Grolle 1990, Parihar et al. 1994, Long 2005), SIKKIM (Mainamcha, 10,000 ft., J.D.Hooker et T.Thomson 1586 cf. Mitten as C. connivens), WEST BENGAL: Darjeeling (Stephani , Chopra 1943, Parihar , Kitagawa 1969, Kachroo 1970, Udar et al. 1970, Udar & Nath 1973, Udar & Kumar 1976, Long & Grolle 1990, Parihar et al. 1994, Bapna & Kachroo 2000, Long 2005, Gao 2003); prope Kurseong, Mahaldaram Gov. Forest, 6800 ft., Nov 23, 1899 Decoly & Schaul, det. Schiffner (Herb. Levier 648) as C. himalayana Schiffn. in sched. (Parihar as C. himalayana Schiffn. et Pandé, nom. nud., Bapna & Kachroo 2000 as C. himalayana Schiffn., nom. nud.); BHUTAN (Long & Grolle 1990, Bapna & Kachroo 2000, Long 2005, Singh & Singh 2007); NEPAL (Long 2005, Singh & Singh 2007); NOTE: Cephalozia himalayana Schiffn. et Pandé was never validly published. Udar & Kumar (1976: p. 35) are the only authors who commented on this species as "is 159

8 only a manuscript species and would await critical investigation for finally settling its status". Cephalozia indica Udar et D.Kumar INDIA, W BENGAL: Darjeeling (Udar & Kumar 1976, Parihar et al. 1994, Bapna & Kachroo 2000). NOTE: This species was commented on by Váňa (1988) that it shows some similarities to members of the subgenus Macrocephalozia R.M.Schust.; however, this subgenus was known at that time only from South and Central America. It is also possible it represents young, poorly-developed plants of another Indian species. The variable shape of the leaves, shallow sinus and the lobes mostly only formed by two cells are unusual and atypical for Cephalozia; they resembles a juvenile Lophocolea species, although the authors state that underleaves are absent which would rule out Lophocolea. Cephalozia lacinulata (J.B.Jack ex Gottsche et Rabenh.) Spruce Cephalozia leucantha Spruce Cephalozia macounii (Austin) Austin Cephalozia pleniceps (Austin) Lindb. BHUTAN (Váňa 1988, Long & Grolle 1990, Bapna & Kachroo 2000); NEPAL (Miehe 1990); CHINA, YUNNAN (Gao & Cao 2000, Gao 2003). CLADOPODIELLA H.Buch TYPE SPECIES: Cladopodiella francisci (Hook.) Jörg. Cladopodiella francisci (Hook.) H.Buch CHINA, XIZANG (TIBET) (Gao 2003), YUNNAN (Gao & Cao 2000, Gao 2003). HYGROBIELLA Spruce TYPE SPECIES: Hygrobiella laxifolia (Hook.) Spruce Hygrobiella laxifolia (Hook.) Spruce IWATSUKIA N.Kitag. TYPE SPECIES: Iwatsukia exigua N.Kitag. Iwatsukia jishibae (Steph.) N.Kitag. NEPAL (Grolle 1966, Bapna & Kachroo 2000). 160

9 METAHYGROBIELLA R.M. Schust. TYPE SPECIES: Metahygrobiella acuminata (Herzog) R.M.Schust. Metahygrobiella albula (Steph.) Grolle SYN.: Cephalozia terminalis S.Hatt.; Sphenolobus langkyrdumii Ajit P.Singh et V.Nath, Hep. Khasi & Jaintia Hills 93, 2007, syn. nov. INDIA, SIKKIM (Stephani as Pleuroclada albula Steph.; Kachroo 1970, Parihar et al. 1994), ASSAM (Kachroo 1970, Parihar et al. 1994, Bapna & Kachroo 2000), MEGHALAYA: Khasia Hills (J.D.Hooker et T.Thomson 1339b [Churra, ft.] cf. Mitten , Stephani as Pleuroclada albula; Kachroo 1970, Long & Grolle 1990, Parihar et al. 1994, Bapna & Kachroo 2000, Singh & Nath 2007 as Sphenolobus langkyrdumii); BHUTAN (Grolle 1963, Kachroo 1970, Long & Grolle 1990, Parihar et al. 1994, Bapna & Kachroo 2000); NEPAL (Hattori 1975 as Cephalozia terminalis S.Hatt.; Parihar et al as Cephalozia terminalis, Bapna & Kachroo 2000 as Cephalozia terminalis). NOWELLIA Mitt. TYPE SPECIES: Nowellia curvifolia (Dicks.) Mitt. Nowellia aciliata (P.C.Chen et P.C.Wu) Mizut. CHINA,YUNNAN (Gao & Cao 2000, Gao 2003). Nowellia curvifolia (Dicks.) Mitt. ODONTOSCHISMA (Dumort.) Dumort. TYPE SPECIES: Odontoschisma sphagni (Dicks.) Dumort. Odontoschisma denudatum (Nees in Mart.) Dumort. INDIA, SIKKIM (Long 2005), MEGHALAYA: East Khasia Hills (Singh & Nath 2007); BHUTAN (Hattori 1971, Long & Grolle 1990, Bapna & Kachroo 2000, Long 2005); NEPAL (Grolle 1966, Long & Grolle 1990, Long 2005, Parihar et al. 1994, Bapna & Kachroo 2000, Singh & Nath 2007); CHINA, YUNNAN (Gao & Cao 2000, Gao 2003) NOTE: Earlier reports from 'Sikkim' and 'Himalaya' are probably all based on J.D.Hooker 1609 (BM, single stem). [Fir-woods, Kambachen, 1100 ft.] cf. Mitten as Sphagnocetis communis, Chopra 1943, Parihar , Kachroo 1970, Parihar et al. 1994, Singh & Nath 2007). The locality is in East Nepal, and the specimen is Bazzania sp. det. Váňa. Odontoschisma grosseverrucosum Steph. Odontoschisma sphagni (Dicks.) Dumort. INDIA, SIKKIM (Chopra 1938, 1943, Parihar , Kachroo 1970, Parihar et al. 1994, Bapna & Kachroo 2000); 161

10 PLEUROCLADULA Grolle TYPE SPECIES: Pleurocladula albescens (Hook.) Grolle Pleurocladula albescens (Hook.) Grolle SCHIFFNERIA Steph. TYPE SPECIES: Schiffneria hyalina Steph. Schiffneria hyalina Steph. INDIA, SIKKIM (Stephani , Chopra 1938, 1943, Kachroo 1970 as S. viridis Steph., Long & Grolle 1990, Parihar et al as S. levieri Schiffn. ex K.I.Goebel and S. viridis; Bapna & Kachroo 2000, Long 2005), WEST BENGAL: Darjeeling (Kitagawa 1973, Long & Grolle 1990, Long 2005); BHUTAN (Long & Grolle 1990, Long 2005); NEPAL (Long 2005); CHINA, YUNNAN (Gao & Cao 2000, Gao 2003), [?SICHUAN] (Piippo 1990 as S. szechuanensis P.C.Chen, nom.nud.). Schiffneria yunnanensis C.H.Gao et W.Li CHINA, YUNNAN (Li et al. 2006). Acknowledgements The senior author acknowledges support for part of this work from the EC SYNTHESYS exchange programme and of the project MŠMT ČR References ARNELL, S.W. (1963): Hepaticae of South Africa. Kungl. Boktryckeriet P.A. Norstedt & Söner, Stockholm. BAPNA K.R. & P. KACHROO (2000): Hepaticology in India - I. Himanshu Publ., Delhi. BRUMMITT, R.K. & C.E. POWELL (1992): Authors of plant names. Royal Bot. Gard., Kew. CHOPRA, R.S. (1938): Notes on Indian Hepatics. II. Sikkim Himalayas and Bengal. Proc. Indian Acad. Sci. 8: CHOPRA, R.S. (1943): A census of Indian Hepatics. J. Indian Bot. Soc. 22: CRANDALL-STOTLER, B.J., R.E. STOTLER, & D.G. LONG (2009): Morphology and classification of the Marchantiophyta. In: GOFFINET, B. & A.J. SHAW (eds.): Bryophyte Biology, 2nd ed.: Cambridge Univ. Press, Cambridge. ENGEL, J.J. (1978): A taxonomic and phytogeographic study of Brunswick Peninsula (Strait of Magellan) Hepaticae and Anthocerotae Fieldiana, Bot. 41: FREY, W. & M. STECH (2009): Division Marchantiophyta Stotler & Crand.-Stotl. (Hepaticae, Liverworts). In: FREY, W. (ed.): Syllabus of Plant Families, 13th ed., Part 3 (Bryophytes and seedless Vascular Plants): Gebr. Borntraeger Verlagsbuchhandlung, Berlin, Stuttgart. FULFORD, M. (1968): Manual of the leafy Hepaticae of Latin America Part III. Mem. Now York Bot. Gard. 11(3): GAO, C. (2003): Cephaloziaceae, pp In: GAO, C. (ed.) (2003): Flora Bryophytorum Sinicorum. Vol. 9. Takakiales Calobryales Jungermanniales. Science Press, Beijing. 162

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