Journal of Bryology ISSN: 0373-6687 (Print) 1743-2820 (Online) Journal homepage: http://www.tandfonline.com/loi/yjbr20 Bryophytes of Kédougou (Eastern Senegal), with a key to the Fissidens of Senegal Djibril Diop, Doudou Diop, Maria Alida Bruggeman-Nannenga, Mame Samba Mbaye, Kandioura Noba, Lars Hedenäs, S. Robbert Gradstein, Catherine Reeb & Alain Vanderpoorten To cite this article: Djibril Diop, Doudou Diop, Maria Alida Bruggeman-Nannenga, Mame Samba Mbaye, Kandioura Noba, Lars Hedenäs, S. Robbert Gradstein, Catherine Reeb & Alain Vanderpoorten (2018): Bryophytes of Kédougou (Eastern Senegal), with a key to the Fissidens of Senegal, Journal of Bryology, DOI: 10.1080/03736687.2017.1415662 To link to this article: https://doi.org/10.1080/03736687.2017.1415662 Published online: 05 Feb 2018. Submit your article to this journal Article views: 2 View related articles View Crossmark data Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=yjbr20 Bryophytes of Kédougou (Eastern Senegal), with a key to the Fissidens of Senegal Djibril Diop 1*, Doudou Diop2*, Maria Alida Bruggeman-Nannenga3*, Mame Samba Mbaye1, Kandioura Noba1, Lars Hedenäs 4, S. Robbert Gradstein5, Catherine Reeb 6, Alain Vanderpoorten7 1 Département de Biologie Végétale, Faculté des Sciences et Techniques, UCAD, Dakar, Sénégal, 2Institut Fondamental d’Afrique Noire Cheikh Anta DIOP, UCAD, Dakar, Sénégal, 3Zeist, Netherlands, 4Department of Botany, Swedish Museum of Natural History, Stockholm, Sweden, 5Cryptogamie, Muséum National d’Histoire Naturelle, Paris, France, 6Institut Systématique et Évolution, Paris, France, 7Institute of Botany, University of Liege, Liege, Belgium The bryophyte flora of the Sahelian region of sub-Saharan Africa is characterized by a low species richness, but is also extremely poorly documented. We present here the results of a floristic survey in Kedougou (eastern Senegal), which yielded 22 species. Two liverworts and 15 moss species are new to the country, so that the bryophyte flora of Senegal now includes a total of 4 liverwort and 34 moss species. An identification key is provided for Fissidens, which, with eight species, is the richest moss genus of the country. Keywords: Liverworts, Mosses, Senegal, Sub-Saharan Africa Introduction Sub-Saharan Africa is characterised by a poor angiosperm (Linder, 2014) and bryophyte (Geffert et al., 2013) flora as compared with other tropical regions. It is widely acknowledged that this general pattern results from the impact of extinctions caused by historical climate shifts towards increasing drought (Parmentier et al., 2007). Evaluating the actual species richness of the sub-Saharan bryophyte flora is, however, hampered by a severe lack of floristic knowledge of many areas, as is evidenced by the striking difference in species numbers reported for countries in the most recent checklists for African mosses (O’Shea, 2006) and liverworts (Wigginton, 2009), and in recent floristic work. This lack of floristic knowledge is highlighted, for example, by the results of a brief (nine-day) bryological investigation of the Atewa Forest, Ghana (Hodgetts et al., 2016), which added 58 species to the 203 previously reported from the country. Although the species richness of this Sahelian region will be relatively low, a lack of floristic knowledge is especially obvious in dry, lowland countries such as Senegal, where only two liverwort and twenty moss species had been reported (O’Shea, 2006; Wigginton, *These authors contributed equally to this work. Correspondence to: Alain Vanderpoorten, Institute of Botany, University of Liege, B22 Sart Tilman, 4000 Liege, Belgium. Email: a.vanderpoorten@ uliege.be © British Bryological Society 2018 DOI 10.1080/03736687.2017.1415662 Published online 05 Feb 2018 2009), and where the only floristic reports consist of single observations made in Casamance during the beginning of the 20th century (see below). In this context, three of us (D. Diop, D. Diop and M.S. Mbaye) initiated a large-scale programme aiming at promoting bryological research and teaching in Senegal. As part of this project, preliminary fieldwork was undertaken in 2015–2016 in Kédougou, which is the easternmost of the 14 regions of Senegal. This area of 16,816 km2 is the most mountainous of the country, ascending to 648 m on the border with Guinea. It is characterised by a Sudano-Guinean climate, with an average total rainfall of 1200 mm per year and a seven-month dry season between October and April. Temperatures are globally high, with 32.9°C in April and a minimum monthly average of 21°C in January. The vegetation is dominated by woody savannas, with gallery forests along the permanent or temporary watercourses (Paterne & Mathieu, 2017). Methods Nomenclature follows O’Shea (2006) for mosses and Söderström et al. (2016) for liverworts. For each species, we provide information on the locality (Table 1), habitat, voucher specimen, literature and references to illustrations, and herbarium where the collection is kept, including the Swedish Museum of Journal of Bryology 2018 1 Diop et al. Bryophytes of Kédougou, Senegal Table 1. List of the localities. Locality Latitude–longitude (altitude) Afia Dande Bandafassi-Kedougou Ségou Lombel, waterfall of Dimboli Djourndé Fongolembi Dindéfélo 12.365°N, 12.363°N, 12.532°N, 12.370°N, 12.447°N, 12.370°N, 12.423°N, 12.365°N, 12.303°W (390 m) 12.328°W (425 m) 12.314°W (250 m) 12.325°W (305 m) 11.996°W (310 m) 12.329°W (295 m) 11.997°W (383 m) 12.325°W (270 m) Natural History (S), the University of Liege (LG) and the Institut Fondamental d’Afrique Noire (INFA) (abbreviations follow Index Herbariorum, http:// sweetgum.nybg.org/science/ih/). A more specific account on Fissidens, which, with eight species, is the richest moss genus of the country, is provided, including African distributions following O’Shea (2006) with some updates and amendments, and based on Hollis & Brummitt’s (1992) geographical units, and a key. Results We recorded 22 species from our study area, of which 17 (15 mosses and 2 liverworts), marked with an *, are new to Senegal. In addition to the species reported below, we found a species of Ectropothecium that is similar to E. perrotii Renauld & Cardot. [Dindéfélo, damp rock in the waterfall (Diop et al. DD9; IFAN, S)]. However, Ectropothecium in Africa includes several similar taxa (Hedenäs, 2005) and the genus is in great need of revision. We therefore refrain from putting a definitive name on this specimen. Liverworts *Riccardia longispica (Steph.) Pearson. Dindéfélo, rocks in the spray zone of a waterfall (Diop et al. DD15, DD16b, DD21a, DD26a, DFL5b; IFAN, LG). Documentation: Wigginton (2004). *Spruceanthus abbreviatus (Mitt.) X.Q.Shi, R.L.Zhu & Gradst. Dindéfélo, damp rocks by the waterfall with Riccardia, Philonotis hastata and Vesicularia oreadelphus (Diop et al. DD26c; IFAN, LG). Documentation: Wigginton (2004), under Archilejeunea abbreviata (Mitt.) Vanden Berghen. Mosses Bryum coronatum Schwägr. Bandafassi-Kedougou, Dande, Fongolembi, rocks along track and manmade walls in rural and urban areas (Diop et al. BDF6, DAN1, FOG1a; IFAN, LG). Documentation: Sharp et al. (1994). *Callicostella subemarginatula Broth. & P.de la Varde. Fongolembi, on Lecaniodiscus cupanioides Planch. ex Benth. roots near the stream (Diop et al. DA1, S, IFAN). Documentation: Demaret & Potier de la Varde (1952). 2 Journal of Bryology 2018 *Entodontopsis rhynchostegioides (Broth. & Paris) W.R.Buck & Ireland. Dande, roots of Ficus cordata Thunb. in gallery forest (Diop et al. DAN2; IFAN, S). Documentation: Buck & Ireland (1985). *Fissidens crispulus Brid. Dindéfélo, Dande, Ségou, ground and rocks in gallery forest and spray zone of a waterfall (Diop et al. DAN7, DD26e, DFL5a, SEG15a; IFAN, priv. herb. Bruggeman-Nannenga). This species is characterised by (1) large axillary nodules, (2) mammillose laminal cells and (3) narrow, up to 2.2 mm long, elimbate leaves (vaginant laminae of perichaetial leaves weakly limbate). Specimens with the above features, but with leaves longer than 2.2 mm, may belong to F. ovatus Brid. Mostly growing on soil or rocks, rarely on wood. Illustration: Bruggeman-Nannenga & Arts (2010), Figure 6. Distribution: Common and widespread in the paleotropics. Also reported from Queensland, northern Australia (Streimann & Klazenga, 2002). In Africa, known from west tropical Africa (Ghana (Bruggeman-Nannenga, 2009), Ivory Coast, Nigeria, Sierra Leone, Togo); west-central tropical Africa (Cameroon, Central African Republic, Democratic Republic of the Congo, Equatorial Guinea (Bioko, Rio Muni), Gabon, São Tomé (Shevock et al., 2013)); north-east tropical Africa (Chad, Eritrea, Ethiopia and Sudan); east tropical Africa (Kenya, Rwanda, Tanzania); south tropical Africa (Angola (Müller, 2015), Malawi, Zambia, Zimbabwe); and western Indian Ocean (Comore islands, La Réunion, Madagascar, Mauritius, Rodriguez, Seychelles). F. dasyphus Welw. & Duby in Duby (not observed in the course of the present study). F. glauculus var. circinicaulis (Cardot) P.de la Varde, syn. nov. Bruggeman-Nannenga & Pursell (1996) and Bruggeman-Nannenga (2009) reduced F. dasyphus and F. glauculus var. circinicaulis into synonymy with F. weirii. It subsequently appeared that the Neotropical F. weirii is not conspecific with the African F. dasyphus (unpublished, but mentioned by Hodgetts et al. (2016); a revision of limbate papillose African Fissidens by Bruggeman-Nannenga is in preparation). Consequently, F. glauculus Müll.Hal. ex Dusén and F. glauculus var. circinicaulis (Cardot) P.de la Varde are synonyms of F. dasyphus rather than of F. weirii. Fissidens dasyphus is characterised by pluripapillose laminal cells and leaves with a limbidium on all laminae (although the extension of the limbidium on the dorsal lamina is variable, as limbidia may be reduced to a few cells in the mid-part of the lamina or be fully developed along almost the complete length). The similar African F. glaucissimus, which also exhibits limbate laminae and pluripapillose cells, can be distinguished by its narrower, linear leaves Diop et al. Bryophytes of Kédougou, Senegal that are always limbate on all lamina. On termite mounds, rock, soil and wood. Illustration: Bruggeman-Nannenga (2006b), Figure 24 f–k (as F. weirii); Chuah-Petiot (2003), Figure 36 (as F. weirii); Ngaba Zogo (1983), p. 243, Figure 2B (as F. glauculus). Distribution: Endemic to Africa, where it is known from west tropical Africa (Burkina Faso, Ghana, Guinea, Ivory Coast, Liberia, Nigeria, Senegal, Sierra Leone, Togo); west-central tropical Africa (Cameroon, Central African Republic, Democratic Republic of the Congo, Equatorial Guinea (Bioko, Rio Muni), Gabon); east tropical Africa (Kenya, Uganda) and south tropical Africa (Angola). In Senegal, there is no report apart from the type collection in Casamance (Cardot, 1908). Fissidens gardneri Mitt. Dande, damp dead wood along the stream with Leucobryum cucullatum and Octoblepharum albidum Hedw. (Diop et al. DA13b; IFAN, priv. herb. Bruggeman-Nannenga). This tiny, elimbate species is characterised by (1) open to almost open vagina lamina, pluripapillose cells, (2) obtuse leaf apices, (3) costae ending far below leaf apex, often covered by laminal cells distally, (4) limbidia restricted to the vaginant laminae of the upper leaves of perichaetial plants and (5) short, undivided to irregularly divided, straight, papillose peristome teeth. Lignicolous (on bark and wood). Illustration: Pursell (2007), Figure 76A–L. Distribution: Pantropical (Pursell et al., 1992). Known from Australia (New Territories, Queensland (Streimann & Klazenga, 2002)), Asia, the Neotropics (Pursell, 2007) and Africa. In Africa, it occurs in west tropical Africa (Guinea, Ivory Costa, Nigeria, Senegal, Togo); west-central tropical Africa (Central African Republic); east tropical Africa (Tanzania) and south tropical Africa (Malawi). In Senegal, the species was reported once on tree bark in Casamance (Chevalier, 1920). Fissidens intramarginatus (Hampe) A.Jaeger. Afia, Dande, on termite mounds (Diop et al. AFI12b, DAN9; IFAN, priv. herb. Bruggeman-Nannenga). This species is recognised by (1) vaginant laminae of all well-developed leaves limbate, limbidia reaching the apex of the vaginant laminae and pluripapillose cells. In Africa, it represents a species-complex rather than a species (see below). Mostly on soil or rock, also decaying wood, termite mounds, roots and tree-ferns. Illustrations: Bruggeman-Nannenga & Arts (2010), Figure 24; Bruggeman-Nannenga (2006a), Figure 11; Chuah-Petiot (2003), Figure 31. Distribution: Known from the Neotropics (Pursell, 2007) and Africa. In Africa, it is known from west tropical Africa (Benin, Gambia (Bruggeman-Nannenga, 1997), Ghana, Guinea, Ivory Coast, Liberia, Nigeria, Senegal, Sierra Leone, Togo); west-central tropical Africa (Central African Republic, Cameroon, Democratic Republic of the Congo, Equatorial Guinea (Bioko), Gabon, São Tomé (Shevock et al., 2013)); north-east tropical Africa (Ethiopia (Hylander et al., 2017), Sudan); east tropical Africa (Kenya, Rwanda, Tanzania, Uganda); south tropical Africa (Angola (Müller, 2015), Malawi, Mozambique (Hedderson et al., 2015)); southern Africa (South Africa) and western Indian Ocean 2009), La (Comores (Bruggeman-Nannenga, Réunion, Madagascar). In Senegal, the species was reported from Casamance (Cardot, 1908). Note. As in F. flaccidus Mitt. (BruggemanNannenga, 2017), the African and Neotropical concepts differ. The circumscription of African specimens is broad and calls for further research. Fissidens marthae Cardot. (not observed in the course of the present study). This species is characterised by (1) short, marginal to intramarginal limbidia on the basal part of vaginant laminae of all well-developed leaves, (2) pluripapillose laminal cells and (3) costae ending two to three cells below leaf apex. Mostly lignicolous (bark, roots, twigs, rotten wood), occasionally found on rock or soil. Illustrations: Bruggeman-Nannenga (2006a), Figure 13f–j. Distribution: Endemic to Africa. Known from west tropical Africa (Benin, Ghana, Ivory Coast, Nigeria, Senegal, Sierra Leone); west-central tropical Africa (Central African Republic, Cameroon, Democratic Republic of the Congo); east tropical Africa (Tanzania, Uganda) and western Indian Ocean (Seychelles). In Senegal, the species was reported once from Casamance (Cardot, 1908). Fissidens parkii Mitt. (not observed in the course of the present study). Fissidens parkii is best characterised by its unique sporophyte. Fortunately, capsules are usually abundantly present. The thecae have distinctive large, oblong exothecial cells (clearly visible at a magnification of ×10), lowly conical opercula with long rostra, and short, undivided, erect peristome teeth with an ornamentation of numerous, horizontal riblets on the exterior side (BruggemanNannenga, 2009, Figure 5). The laminal cells are smooth to weakly mammillose. Fissidens parkii can be confused with F. scleromitrius, which, however, lacks a peristome. Mostly collected on termite nests or soil, more rarely saxicolous. Illustrations: Bruggeman-Nannenga (2009), Figure 5 (peristome); Suzuki (2014), Figure 1. Distribution: A paleotropic species known in Asia from Thailand (Ellis et al., 2014) and Laos (Suzuki, 2014). In Africa, the species was recorded from west Journal of Bryology 2018 3 Diop et al. Bryophytes of Kédougou, Senegal tropical Africa (Senegal, Gambia (BruggemanNannenga, 2009), Guinea, Ivory Coast, Nigeria, Mali); north-east tropical Africa (Chad); westcentral tropical Africa (Central African Republic, Gabon) and east tropical Africa (Tanzania). In Senegal, the species was first collected in Casamance and reported as Moenkemeyera macrocarpa Broth. & Par. (a synonym of F. parkii) (Paris, 1911). *Fissidens ramulosus Mitt. Dindéfélo, rocks along the stream (Diop et al. DD38; IFAN, priv. herb. Bruggeman-Nannenga). This species is characterised by mammillose laminal cells and elimbate leaves, except for the upper and mid leaves of perichaetial stems. It is found on soil or rocks, less often on termite mounds or lignicolous (dead wood and bark). Illustrations: Bruggeman-Nannenga & Arts (2010), Figure 33; Bruggeman-Nannenga (2006b), Figure 20f–k. Distribution: African endemic known from west tropical Africa (Ghana (Bruggeman-Nannenga, 2009), Ivory Coast); west-central Africa (Cameroon, Central African Republic, Democratic Republic of the Congo, Gabon, São Tomé (Shevock et al., 2013)); north-east tropical Africa (Ethiopia (Hylander et al., 2017)); east tropical Africa (Kenya, Tanzania, Uganda); south tropical Africa (Angola, Malawi); southern Africa (Botswana (Bruggeman-Nannenga, 2009)) and west Indian Ocean (Comores, Madagascar, Rodriguez). Fissidens submarginatus Bruch ex C. Krauss (not observed in the course of the present study). This species is well characterised by limbate vaginant laminae on all leaves and highly mammillose laminal cells. Mostly on soil, also on decaying wood, stones and termite mounds. Illustrations: Bruggeman-Nannenga & Arts (2010) Figure 26; Bruggeman-Nannenga (2006b), Figure 24a–e. Distribution: Pantropical, occurring in the United States (Alabama and Louisiana), the Neotropics, (Pursell, 2007), Hawaii (Pursell, 2007; Staples et al., 2004) and Africa. In Africa known from west tropical Africa (Benin, Ghana, Guinea, Ivory Coast, Liberia, Nigeria, Senegal, Togo); west-central tropical Africa (Cameroon, Central African Republic, Democratic Republic of the Congo, Equatorial Guinea, São Tomé (Shevock et al., 2013)); north-east tropical Africa (Ethiopia (Hylander et al., 2017)); east tropical Africa (Kenya, Rwanda (Hedderson et al., 2014), Tanzania, Uganda); south tropical Africa (Angola (Müller, 2015), Malawi, Zambia, Zimbabwe); southern Africa (Namibia, South Africa, Swaziland) and western Indian Ocean (Madagascar). In Senegal, the species was reported once from Casamance (Cardot, 1908). 4 Journal of Bryology 2018 *Hyophila involuta (Hook.) A.Jaeger. Dande, Dindéfélo, Djourndé, Fongolembi, Dimboli, Ségou, rocks along stream in gallery forest and by waterfall, also on damp well walls in rural areas with Bryum coronatum (Diop et al. DAN4, DAN6, DD1, DRD1, DFL3, FOG1b, LOM1, SO11a; IFAN, LG). Documentation: Magill (1981). *Leucobryum cucullatum Broth. Dande, damp dead wood along a stream with Octoblepharum albidum (Diop et al. DA13a; IFAN, LG). Documentation: Potier de la Varde (1936). Octoblepharum albidum Hedw., Dande, damp dead wood along the stream with Leucobryum cucullatum and Fissidens gardneri (Diop et al. DA13b). Pelekium gratum (P.Beauv.) Touw. Afia, Dande, Ségou, rocks and dead wood in gallery forest and by waterfall, also on Terminalia catappa L. trunk in rural areas (Diop et al. AFI15, AFI30, DA6, DA17, DAN8, SEG21, SO11b; IFAN, LG, S). African and Asian specimens assigned to this species by Norhazrina et al. (2016) were not monophyletic and a revision of the genus is necessary. Documentation: Touw (1976). *Pelekium investe (Mitt.) Touw. Ségou, on damp rock along stream in gallery forest (Diop et al. SEG15b, SEG21; S, IFAN). African and Neotropical specimens assigned to this species by Norhazrina et al. (2016) were not monophyletic and a revision of the genus is necessary. Documentation: Touw (1976), under Thuidium byssoideum Besch. and T. tenuissimum Welv. & Duby cf. Phephu (2013). *Phyllodon truncatus (Welw. & Duby) W.R.Buck. Dindéfélo, on damp rocks in a waterfall (Diop et al. DD21c; IFAN, S). Documentation: Kis (2002). *Philonotis hastata (Duby) Wijk & Margad. Dindéfélo, Dimboli, waterfall (Diop et al. DD26d, LOM2; IFAN, LG). Documentation: Magill (1987). *Racopilum africanum Mitt. Dande, Ségou, on Treculia africana Decne. trunk in rural area, also on rocks with Stereophyllum radiculosum in gallery forest (Diop et al. DA15a, DA16b, SEG7; IFAN, S). Documentation: Potier de la Varde (1936). *Rhacopilopsis variegata (Welw. & Duby) M.C.Watling & O’Shea. Dande, on Diospyros mespiliformis Hochst. ex A.DC. trunk (Diop et al. DA11; IFAN, S). Documentation: Hedenäs & Watling (2005), Watling & O’Shea (2000). *Stereophyllum radiculosum (Hook.) Mitt. Dande, Djourndé, Ségou, rocks, dead wood and Carapa Aubl. trunk in gallery forest with Khaya senegalensis (Desr.) A.Juss. (Diop et al. DA16a, DRD2; S, IFAN, LG). Documentation: Buck (1998). *Taxithelium homalophyllum (Mitt.) Broth. Ségou, roots of Saba senegalensis (A.DC.) Pichon in gallery forest (Diop et al. SO7; IFAN, S). Documentation: Câmara (2011). Diop et al. Bryophytes of Kédougou, Senegal *Trachyphyllum dusenii (Müll.Hal. ex Broth.) Broth. Dande, on Treculia africana trunk in rural area with Racopilum africanum (Diop et al. DA15b; S, IFAN). Documentation: Buck (1979). *Vesicularia oreadelphus (Broth.) Broth. Dindéfélo, damp rocks by the waterfall with Riccardia (Diop et al. DD16a; IFAN, S). Documentation: Hedenäs & Watling (2005). Key to the Fissidens species of Senegal 1 Peristome undivided or irregularly divided . . . . 2 1 Basal part peristome undivided and distal part divided into two long filaments, or peristome lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2 Vaginant laminae subequal with the smaller lamella ending close to the margin; laminal cells weakly mammillose; peristome ornamentation of horizontal riblets . . . . . . . . . . . . . . . . . . F. parkii 2 Vaginant laminae strongly unequal with the smaller lamella ending at or near the costa; laminal cells pluripapillose; peristome ornamentation different . . . . . . . . . . . . . . . . . . . F. gardneri 3 Laminal cells pluripapillose . . . . . . . . . . . . . . . 4 3 Laminal cells smooth or mammillose . . . . . . . . 7 4 Smaller lamella of the vaginant lamina ending at the costa (vaginant lamina open); leaves elimbate, except for the upper leaves of perichaetial stems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. gardneri 4 Smaller lamella of the vaginant lamina ending near the leaf margin; limbidia various . . . . . . . . 5 5 At least some leaves limbate on all laminae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. dasyphus 5 Limbidia restricted to the vaginant laminae . . . . . .6 6 Limbidia reaching the apex of the vaginant lamina to extending slightly onto the apical lamina . . . . . . . . . . . . . . . . . . F. intramarginatus 6 Limbidia shorter, extending 1/10–1/3 of the vaginant laminae . . . . . . . . . . . . . . . . . . . F. marthae 7 Large axillary nodules present . . . . . . F. crispulus 7 Large axillary nodules lacking . . . . . . . . . . . . . 8 8 Limbidia present on all well-developed leaves, extending the length of the vaginant laminae; laminal cells highly and sharply mammillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. submarginatus 8 Limbidia shorter, not present on all leaves . . . . 9 9 Laminal cells smooth to inconspicuously mammillose; peristome teeth short and undivided with an ornamentation of horizontal lines . . . . . . F. parkii 9 Laminal cells mammillose; peristome teeth divided into two long spirally ornamented filaments (scariosus type of peristome) . . . . . . . . F. ramulosus Acknowledgements Many thanks are due to two referees for their constructive comments on the manuscript. Taxonomic Additions and Changes: Fissidens dasyphus Welw. & Duby in Duby (= F. glauculus var. circinicaulis (Cardot) P.de la Varde), reinstated. 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