Nineteen species of bryophytes are reported for the first time for Réunion. Especially remarkable are the first African record of Anastrophyllum leucocephalum (Tayl.) Steph. and the first records for the East African Islands of Cephaloziella cf. vaginans Steph., Fossombronia foveolata Lindb., Riccia huebeneriana Lindenb., Barbula unguiculata Hedw., Bryum pseudotriquetrum (Hedw.) Schwägr., Chenia leptophylla (Müll.Hal.) R.H.Zander, Dicranella subsubulata (Hampe ex Müll.Hal.) A.Jaeger, Ditrichum punctulatum Mitt., Physcomitrium spathulatum (Hornsch.) Müll.Hal. var. spathulatum and Trichostomum crispulum Bruch var. crispulum. Brachymenium gemmiferum A.J.Shaw & W.R.Buck is regarded as synonymous with Brachymenium exile (Dozy & Molk.) Bosch & Sande Lac

Müller, Frank

English

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91TROPICAL BRYOLOGY 18 (2000)Bryophyte flora of RéunionTropical Bryology 18: 91-95, 2000Additions to the bryophyte flora of Réunion (East African Islands)Frank MüllerInstitut für Botanik, Technische Universität Dresden, Mommsenstr. 13, D-01062 Dresden,GermanyAbstract: Nineteen species of bryophytes are reported for the first time for Réunion. Especiallyremarkable are the first African record of Anastrophyllum leucocephalum (Tayl.) Steph. and thefirst records for the East African Islands of Cephaloziella cf. vaginans Steph., Fossombronia foveolataLindb., Riccia huebeneriana Lindenb., Barbula unguiculata Hedw., Bryum pseudotriquetrum (Hedw.)Schwägr., Chenia leptophylla (Müll.Hal.) R.H.Zander, Dicranella subsubulata (Hampe ex Müll.Hal.)A.Jaeger, Ditrichum punctulatum Mitt., Physcomitrium spathulatum (Hornsch.) Müll.Hal. var.spathulatum and Trichostomum crispulum Bruch var. crispulum.Brachymenium gemmiferum A.J.Shaw & W.R.Buck is regarded as synonymous with Brachymeniumexile (Dozy & Molk.) Bosch & Sande Lac.IntroductionIn September and October 1998 the authorcollected bryophytes on the island of Réunion, aFrench overseas department situated in theIndian Ocean 700 km east of Madagascar. Incomparison with other regions of Africa,Réunion belongs to the bryologically betterinvestigated areas. An overview of the bryophytesknown from the island is provided by thechecklists of Grolle (1995) and O’Shea (1999).There are 484 mosses and about 220 liverwortslisted for the island. For the small size of theisland (2510 km²) this is a remarkable speciesrichness. The cause of the bryophyte richness isnot only the extensive investigation in the pastbut also the very high diversity of landscape,vegetation and geology and the tropical, wetclimate. Among the bryophytes discovered by theauthor there are a lot new for the island. Most ofthe new species were found in habitats which inthe past were obviously investigated lessintensively: walls and open soil in urban areas,shores of lakes and alpine vegetation around thehighest mountain of the island, the Piton desNeiges (3069 m). The records of some additionalmosses of the Réunion trip are already publishedin Müller & Schäfer-Verwimp (1999).Annotated list of speciesThe nomenclature and systematic arrangementof the species follows mainly O’Shea (1999) forthe mosses and Wigginton & Grolle (1996) forthe liverworts.  Voucher specimens are in mypersonal herbarium. For specimens sent tospecialists for determination or confirmation,duplicates exist in their personal herbaria. Allspecies are reported for the first time for Réunion.92TROPICAL BRYOLOGY 18 (2000)MüllerHepaticaeAnastrophyllum leucocephalum (Taylor) Steph.(= A. crebrifolium (Hook.f. & Taylor) Steph.)Hell-Bourg SW: near the Gite de la CaverneDufour below Piton des Neiges, c. 2450 m; rocky,boulder rich area with fragmentary Philippiabushes, on a rocky embankment; 22.9.1998;herb. F. Müller R 556, det. J. VánaHell-Bourg SSW: between Gite de la CaverneDefour and Cap Anglais, low Philippia forest,on forest ground, c. 2300 m; 23.9.1998; herb. F.Müller R 618, conf. J. VánaThis taxa is reported for the first time for Africa.Up to now it was only known from LatinAmerica, where it shows a montane-subantarcticdistribution. In the Americas the species isknown from Tierra del Fuego, Cape Horn,Patagonian Channels, Juan Fernandez and thehigher reaches of the Andes of Colombia,Bolivia, Peru and Ecuador (Engel 1978, Vána1984).The genus Anastrophyllum contains worldwideabout 35-40 species and represents a genus withGondwanalandic origin and speciation: themajority of the taxa are known from tropicalmountains in South America and Australasia(Vána 1999). In Africa four additional taxa areknown, A. auritum (Lehm.) Steph., A. minutum(Schreb. ex Cranz) Schuster, A. piligerum (Nees)Steph. and A. subcomplicatum (Lehm. &Lindenb.) Steph. Distribution centres of thegenus in Africa are the East African mountains,South Africa and the Mt. Cameroon. From theEast African Islands hitherto only A. piligerumwas known, which occurs occasionally onMadagascar, Mauritius, Réunion and Seychelles.A. leucocephalum represents an additionalAmerican-African geoelement.Bryophytes of southern hemispheric orsubantarctic distribution occur on Réunionmainly around Piton des Neiges, the highestmountain of the island. In addition on thismountain the mosses Amphidium tortuosum(Hornsch.) Cufod., Philonotis scabrifolia(Hook.f. & Wilson) Braithw. (see Müller &Schäfer-Verwimp 1999) and Ditrichumpunctulatum (see below) were found.Cephaloziella cf. vaginans Steph.Piton des Neiges: on the way to the top on theSE side; c. 2750 m; subalpine, fragmentedvegetation-covered scree; 23.9.1998; herb. F.Müller R 549, det. J. VánaNew for the East African Islands. The specieswas hitherto known only from several countriesof continental Africa (e.g. Cameroon, Rwanda,Tanzania, Zaire and probably Sierra Leone andif C. umtaliensis Arn. is really a synonym of thistaxa, as suggested in Wigginton & Grolle [1996],also from Mozambique and Zimbabwe).Fossombronia foveolata Lindb.Hell-Bourg SE: section of Foret de Belouveeastern and southeastern of the Gite de Belouve;c. 1500 m; mountain rainforest, wet embankmenton the lake le Grande Mare; 20.9.1998; leg. F.Müller R 5, det. S. PeroldGrand Ilet NE: shores of the lake Mare a Martin,1100 m; muddy soil along the lake shores;24.9.1998; leg. F. Müller R 338, det. S. PeroldNew for the East African Islands. In sub-SaharanAfrica additional records exist only for SouthAfrica, but the genus is being revised (S. Perold,R.E. Stotler & B. Crandall-Stotler), so that thefuture should provide more information aboutthe African distribution of this taxon.Odontoschisma africanum (Pears.) SimHell-Bourg SSW: between Gite de la CaverneDefour and Cap Anglais; low Philippia forest,on dead wood, c. 2300 m; 23.9.1998; herb. F.Müller R 590, conf. J. VánaA rare African element, which was knownhitherto in the East African Islands only fromMauritius; in continental Africa is it known froma few sites in South Africa, from where thespecies was described, and Tanzania.Riccia huebeneriana Lindenb.La Plaine-des-Palmistes N: shores of the lake leGrand Etang; c. 500 m; on lake mud; 30.9.1998;herb. F. Müller R 158, R 159, R 201, conf. S.PeroldIn sub-Saharan Africa hitherto known only fromone record in Ethiopia. The new record on93TROPICAL BRYOLOGY 18 (2000)Bryophyte flora of RéunionRéunion represent the first collection for the EastAfrican Islands and probably for the wholesouthern hemisphere. R. huebeneriana is atemperate species of the northern hemisphere,which is known from scattered localities inEurope, northwards to Scandinavia and NorthRussia, northern Africa (probably Algeria), Indiaand Japan. The material from Réunion agree wellwith Central European material from Germany.Among the holarctic phanerogams of the dwarfrush communities (phytosociological unitNanocyperion) many species have disjunct singleoccurrences in tropical and southern Africa, e.g.Dichostylis micheliana (L.) Nees, Cyperusflavescens L., Juncus bufonius L., Isolepissetacea (L.) R. Br., Juncus capitatus Weigel (seedistribution maps by Lampe 1996), whichoriginate from the transport of diaspores bywaterbirds (ornithochore diaspore dispersion).Probably also R. huebeneriana belongs with thisgroup of ornithochore aquatic plants.MusciAnomodon pseudotristis (Müll.Hal.) Kindb.La Plaine-des-Cafres NW: on the way from BoisCourt to Grand Bassin, c. 800 m, on a rockfacefully exposed to the light and as epiphyte on trees;25.9.1998; herb. F. Müller R 383, R 411The material was determined with help of thekeys in the revision of the genus by Granzow-dela Cerda (1997). Haplohymenium tenuissimum(Besch.) Broth. was the only species mentionedfrom Réunion by Brotherus (1925), but this mayhave been an error, as the original description(Bescherelle 1880) quoted this species only fromMauritius. This taxon is not treated in themonographic treatment of Anomodon andHaplohymenium (Granzow-de la Cerda 1997),but O’Shea (in press) synonymised H.tenuissimum with Anomodon pseudotristis andsaw only Mauritius collections.Barbula unguiculata Hedw.Hell-Bourg SE: section of Foret de Belouveeastern and southeastern of the Gite de Belouve,c. 1500 m, mountain rainforest, on a wall;20.9.1998; herb. F. Müller R 612A species with a more or less subcosmopolitanworld distribution. The occurrence on the islandrests most likely on introduction. In sub-SaharanAfrica the species is known from South Africa,Angola, Cape Verde and Sudan. First record forthe East African Islands.Brachymenium exile (Dozy & Molk.) Bosch &Sande Lac., Bryologia javanica 1: 139. 1860.Basionym: Bryum exile Dozy & Molk., Ann. Sci.Nat. Bot., III, 2: 300. 1844.(Syn. nov.: Brachymenium gemmiferumA.J.Shaw & W.R.Buck, Bryologist 101: 601.1998. Typus: Réunion, Lorence 7512, NY)Grand Ilet N: climb to the Roche Ecrite, alongthe hiking trail to the top, c. 1500 m,embankment in rainforest along the way,associated with Anomobryum laceratum (Besch.)Broth. and Philonotis spec.; 24.9.1998; herb. F.Müller R 433A pantropical element, widely distributed intropical Africa.Brachymenium gemmiferum A.J.Shaw &W.R.Buck was described based on material fromRéunion as new for science (Shaw, Allen & Buck1998). A special characteristic of this species isthe presence of axillary gemmae; it should be„the only species of Brachymenium known toform gemmae“. This observation is incorrect.The presence of axillary gemmae is already welldocumented in the literature for anotherBrachymenium species, B. exile (see Ochi 1972& 1994; Eddy 1996). Brachymeniumgemmiferum is well described and illustrated inShaw, Allen & Buck (1998) and there is no doubtthat it is identical with B. exile. Therefore it istreated here as synonymous with this taxa.Brachymenium longicolle Thér.Foret de Bebour: Sentier de la Riviere in thevalley Riviere des Marsouins, c. 1350 m ,epiphytic in mountain rainforest; 2.10.1998;herb. F. Müller R 387A rare afromontane element, which is knownfrom higher mountains of West Africa(Cameroon, Bioko), Tanzania, Malawi andMauritius and occurs elsewhere also in the E.Himalayas. Our material agrees well with thedescription and figures in Ochi (1972) (leaveswith long acuminate apex, operculum short-conic with a pointed apex, spores over 40 µm in94TROPICAL BRYOLOGY 18 (2000)Müllerdiameter) and are clearly distinct from theMascarene endemic B. eurychelium Besch.Bryum pseudotriquetrum (Hedw.) Schwägr.La Plaine-des-Palmistes: in the village in the areaof the hotel „Les Azalees“, c. 900 m; in lawn;2.10.1998; herb. F. Müller R 30Subcosmopolitan species which is known in sub-Saharan Africa from many countries. First recordfor the East African Islands. The occurrence onRéunion rests most likely on introduction,because the species was collected in a man-madehabitat together with Pseudoscleropodium purum(Hedw.) M. Fleisch, another introduced speciesof the island.Calymperes palisotii Schwägr. ssp. palisotiiSt-Philippe W: not much north of the village LeBaril, c. 100 m, epiphytic on a roadside tree;27.9.1998; herb. F. Müller R 674, conf. S. OrbanPantropical species which is widely distributedin sub-Saharan Africa.Chenia leptophylla (Müll.Hal.) R.H.ZanderLe Plaine-des-Cafres NW: along the way fromBois Court to Grand Bassin, c. 600 m, on soil bythe wayside in the village Grand Bassin;25.9.1998; herb. F. Müller R 443Subcosmopolitan species, easily overlooked,which is known in sub-Saharan Africa especiallyfrom the southern part. First record for the EastAfrican Islands.Dicranella subsubulata (Müll.Hal.) A.JaegerLa Plaine-des-Palmistes: in the village, c. 900m, rocky embankment of a road; 1.10.1998; herb.F. Müller R 47Hell-Bourg SE: section of the Foret de Belouveeastern and southeastern of the Gite de Belouve,c. 1500 m , mountain rainforest, wetembankment on the lake le Grande Mare;20.9.1998; herb. F. Müller R 249The species is known from many countries oftropical and southern Africa. The material fromRéunion agrees well with the description andfigures in Magill (1981). First record for the EastAfrican Islands.Ditrichum punctulatum Mitt.Piton des Neiges: climb to the top on thesoutheastern side, c. 2750 m; subalpine,fragmented vegetation-covered scree; 23.9.1998;herb. F. Müller R 576, R 577Primarily a southern hemisphere species, whichis known from Australia, New Zealand, Ecuadorand from the higher mountains aroundDrakensberg and Gaika’s Kop in South Africaand Lesotho. First record for the East AfricanIslands. Like the South African material (Magill1981) the Réunion collections are also only insterile condition.Erythrodontium spec.Hell-Bourg: on the western edge of the villageby the thermal springs, 900 m, epiphytic in apark-like area; 21.9.1998; herb. F. Müller R 356In the literature there is no information aboutthe occurrence of Erythrodontium species onRéunion. Our material cannot determined tospecies level because of its sterile condition andthe lack of a taxonomic treatment of the Africanspecies of the genus.Leucomium strumosum (Hornsch.) Mitt.St-Philippe NW: Reserve Naturelle de MareLongue, along the Sentier botanique, c. 250 m ,lowland rainforest, on dead wood; 29.9.1998;herb. F. Müller R 164Pantropical element, widely distributed intropical Africa.Physcomitrium spathulatum (Hornsch.)Müll.Hal. var. spathulatumLa Plaine-des-Palmistes N: shores of the lake leGrand Etang; c. 500 m; on lake mud; 30.9.1998;herb. F. Müller R 205The material was compared with an isotype ofP. spathulatum in BM (herbarium Schimper,Cape) and with the description and figures inMagill (1987) and agree well. In Africa thespecies was hitherto known from the southernand eastern part. First record for the East AfricanIslands.Pseudephemerum nitidum (Hedw.) LoeskeLa Plaine-des-Palmistes N: shores of the lake leGrand Etang; c. 500 m; on lake mud; 30.9.1998;herb. F. Müller R 115Grand Ilet NE: shores of the lake Mare a Martin,1100 m , on lake mud; 24.9.1998; herb. F. Müller95TROPICAL BRYOLOGY 18 (2000)Bryophyte flora of RéunionR 332A species of Europe and North America whichis also known from few, scattered localities innorthern, central and eastern Africa and probablyalso occur in eastern Asia. Like Ricciahuebeneriana (see above) this species belongsto a group of water plants with ornithochorediaspore dispersion.Trichostomum crispulum Bruch var. crispulumHell-Bourg NW: mountain Piton d’Enchaing,along the hiking trail to the top, 900 m ,  onboulders in cultivated area; 21.9.1998; herb. F.Müller R 450Hell-Bourg: on the western edge of the villagealong the track to the thermal springs, 900 m ,on a wall; 21.9.1998; leg. F. Müller R 329Subcosmopolitan species, which is known in sub-Saharan Africa from South Africa, Djibouti,Chad, Tanzania and the islands of Cape Verdeand St. Helena. First record for the East AfricanIslands.AcknowledgementsFor determination or revision of material I wish tothank the following persons: J. Vána (Praha) -Anastrophyllum leucocephalum, Cephaloziellavaginans, Odontoschisma africanum,  S. Perold(Pretoria) - Fossombronia foveolata, Ricciahuebeneriana and S. Orban (Eger) - Calymperespalisotii. My special thank goes to Mr B.J. O‘Shea(London) for correcting the English text and helpfulremarks.LiteratureBescherelle, E. 1880. Florule bryologique de laRéunion et des autres iles austro-africaines, del’ocean Indien (Deuxieme partie). Annales desSciences Naturelles, Botanique, séries 6 , 10: 233-332.Brotherus, V.F. 1925. Musci (Laubmoose), 2. Hälfte.In: Engler, A. (ed.): Die natürlichenPflanzenfamilien, Band 11. Berlin: Duncker &Humblot.Eddy, A. 1996. A Handbook of Malesian Mosses.Volume 3.  London: HMSO.Engel, J.J. 1978. A Taxonomic and PhytogeographicStudy of Brunswick Peninsula (Strait of Magellan)Hepaticae and Anthocerotae.  Fieldiana, Botany41: 1-319.Granzow-de la Cerda, I. 1997. Revision andphylogeny of Anomodon and Herpetineuron(Anomodontaceae, Musci).  Contributions of theUniversity of Michigan Herbarium 21: 205-275.Grolle, R. 1995. The Hepaticae and Anthocerotae ofthe East African Islands. An Annotated Catalogue.Bryophytorum Bibliotheca 48: 1-178.Lampe, M. v., 1996. Wuchsform, Wuchsrhythmus undVerbreitung der Arten derZwergbinsengesellschaften.  DissertationesBotanicae 266: 1-357.Magill, R.E. 1981: Bryophyta, Part 1 Mosses,Fascicle 1 Sphagnaceae-Grimmiaceae.  In:Leistner, O.A. (ed.): Flora of Southern Africa.Pretoria: Botanical Research Institute.Magill, R.E. 1987: Bryophyta, Part 1 Mosses,Fascicle 2 Gigaspermaceae-Bartramiaceae.  In:Leistner, O.A. (ed.): Flora of Southern Africa.Pretoria: Botanical Research Institute.Müller, F. & Schäfer-Verwimp, A. 1999. Newbryophyte taxon records for tropical countries III.Tropical Bryology 16: 195-201.Ochi, H.1972. A Revision of African Bryoideae,Musci (First Part).  Journal of the  Faculty ofEducation of Tottori University, Natural Science23: 1-126.Ochi, H. 1994. Brachymenium. In: Sharp, A.J., Crum,H. & Eckel, P.M. (ed.): The Moss Flora of Mexico.Part 1. - Memoirs of The New York BotanicalGarden 69.O’Shea, B.J. 1999. Checklist of the mosses of sub-Saharan Africa (version 3, 11/99). TropicalBryology Research Reports 1: 1-133.O’Shea, B.J. (in press). Taxonomic notes onAnomodon (Anomodontaceae, Bryopsida) inAfrica.   Journal of Bryology.Shaw, A.J., Allen, B. & Buck, W.R. 1998. NewSpecies of Bryaceae (Mielichhoferia,Brachymenium) from Costa Rica and Réunion.Bryologist 101: 600-604.Vána, J., 1984.  Anastrophyllum (Spruce) Steph. inLatin America - preliminary information.Proceedings of the Third Meeting of theBryologists from Central and East Europe, Praha,14th - 18th June 1982; p. 99-106.Vána, J., 1999.  Phytogeography of the Junger-manniineae (Hepaticae).  Bryobrothera 5:253-267.Wigginton, M.J. & Grolle, R.  1996.  Catalogueof the Hepaticae and Anthocerotae of Sub-Saharan Africa.  Bryophytorum Bibliotheca50: 1-267.96TROPICAL BRYOLOGY 18 (2000)Müller

Additions to the bryophyte flora of Réunion (East African Islands)

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