Cryptogamie,Bryologie,2010,31(1):31-66 ©2010Adac.Tous droits réservés
Furthertaxonomicstudieson the familiesCalymperaceae (Musci) and Orthotrichaceae(Musci) in the bryofloraof Réunion Island,withnotes on taxafrom otherislandsin the westernIndianOcean
JoannaWILBRAHAM* &Len ELLIS
Departmentof Botany,The NaturalHistory Museum,Cromwell Road, London SW75BD, United Kingdom
(Received 3November2008,accepted 28January 2009)
Abstract –The systematicsand geographicaldistribution of some taxain the moss families Calymperaceaeand Orthotrichaceaethatoccur in Réunion and neighbouring islandsare revised. Inthe familyCalymperaceae,three newsubspeciesareproposed: Calymperes taitense (Sull.) Mitt.subsp. pachyloma (Hampe ex Müll. Hal.) L.T.Ellis, comb.etstat.nov. , Syrrhopodon hispidocostatus Renauld et Cardotsubsp. artsii L.T.Ellis, subsp. nov. and S.flexifolius Mitt.subsp .reunionensis L.T.Ellis, subsp. nov .;three furthernewrecordsfor Réunion Island arecited: Calymperespallidum Mitt., Syrrhopodon dimorphophyllus L.T.Ellis, S.vardei L.T.Ellis,and the presenceand geographicaldistribution in the East AfricanIslandsisdiscussed of Calymperescouguiense Besch., Syrrhopodon asper Mitt., S.africanus (Mitt.) Paris, S.gardneri (Hook.) Schwägr.,and S.pulcher W.D.Reese. Inthe familyOrthotrichaceae,newsynonymyand distribution recordsarereported forthe genus Macromitrium. Severalnewsynonymsareproposed under Macromitriummicrostomum (Hook. &Grev.) Schwägr., M.serpens (Hook. et Grev.) Brid., M.mauritianum Schwägr.and M.pallidum (P.Beauv.) Wijk et Margad .Macromitriumfimbriatum (P.Beauv.) Schwägr. var. chloromitrium Besch. isrecognised asaspeciesin its ownright, M.chloromitrium (Besch.) Wilbraham, comb.etstat.nov .,and isnewlyrecorded from Réunion. Macromitriumbelangeri Müll. Hal. isexcluded from the Réunion Island’smoss flora.
Calymperaceae/Orthotrichaceae/ Syrrhopodon / Calymperes/Macromitrium/ Réunion Island /Mascarene Islands/taxonomy/phytogeography
INTRODUCTION
Ah-Peng &Bardat(2005) published the first modern,comprehensive checklist of bryophytesoccurring in Réunion Island. Subsequently,Ellis& Wilbraham(2008) proposed some alterationstothe taxalisted in the moss familiesOrthotrichaceaeand Calymperaceae. Continuing researchonthese familiesindicatesthe need foryetfurthermodification tothe reported bryoflora of Réunion and alsotothe bryoflorasof some of the otherislandsin the western IndianOcean. Inthe familyCalymperaceae,thesechangesinclude newrecords, proposalsfornewsynonymy,the reclassification of various taxa, and the
*Correspondenceand reprints:[email protected] 32 J.Wilbraham&L.Ellis recognition of three newsubspecies.Theyparticularlyconcerntaxain the genus Calymperes , Syrrhopodon subg. Pseudocalymperes and the Syrrhopodon prolifer Schwägr.complex.Most significantly,among the collectionsof the lattergroup from Réunion,adistinctmoss closelyrelated to S.hispidocostatus Renauld & Cardotand S.apertifolius Besch.,hasbecome evident,and presentlyisprobably best regarded asasubspeciesof S.hispidocostatus.Below,thisnewsubspeciesis named afterthe lateTheo Arts who made importantcontributionstoour knowledge of the bryofloraof Réunion Island and collected the type materialof the taxon. Additionally,the presenceand distribution in the East Africanislandsof Syrrhopodon gardneri (Hook.) Schwägr.and S.asper Mitt.isdiscussed,and the proposed recordsfortheseislandsof Calymperescouguiense Besch. and Syrrhopodon pulcher W.D.Reeseareredetermined. Ofthe non-leucobryoid members of the familyCalymperaceaeoccurring in Réunion island there presentlyappeartobeeightdistincttaxaof Calymperes and fifteen of Syrrhopodon (see Appendix). Thisisperhapsmorethan60%ofthe accepted taxa in the non-leucobryoid Calymperaceaerepresented in the East Africanislandsas awhole. Macromitrium (Orthotrichaceae) represents one of the largest moss generain the East Africanislands,with26 taxapresentlyrecognised from the region. Although the numberof accepted speciescontinuestobereduced with criticalrevision,thereisclearlyanaturallyhigh diversity of Macromitrium in the East Africanislands.Nine speciesof Macromitrium arecurrentlyknownfrom Réunion island (see Appendix). Inthispaper,severalnamesused fortaxaknown from the Mascarenesareplaced in synonymyunder Macromitriummauritianum Schwägr., M.microstomum (Hook. &Grev.) Schwägr., M.pallidum (P.Beauv.) Wijk &Margad. and M.serpens (Hook. &Grev.) Brid. The status of M.fimbriation var. chloromitrium Besch. isre-evaluated and recombined asthe species M.chloromitrium (Besch.) Wilbraham, comb.&stat.nov.Macromitrium lanceolatum Broth. wasoriginallyrecorded from Réunion byEen (1978) though thisrecordhasbeen overlooked in subsequentchecklists of Réunion Island mosses.
REVISED SYSTEMATICS AND GEOGRAPHICAL DISTRIBUTION
CALYMPERES (Calymperaceae) Calymperestaitense (Sull.) Mitt. subsp. pachyloma (Hampe) L.T.Ellis, comb. &stat.nov. Basionym: Calymperespachyloma Hampe ex Müll. Hal., Linnaea 40:247.1876. Type citation. Comoro-insulaJohanna, inter RhynchostegiumComorae ,600 met. supramare,adtruncosarborumsylvae:J.M.Hildebrandt1875cespitulumunicum misit. Type specimen. ComoroIslands,Anjuoan[Johanna], Hildebrandts.n. (BM000555463 –Hb.Hampe! – lectotype, vide Ellis,1995). Syrrhopodon nossibeanus Besch., Ann. Sci. Nat.Bot., Sér . 6,9:3471880. ≡ Calymperesnossibeanum (Besch.) Broth. in Engl. &Prantl, Nat.Pflanzenfam . 1(3):389. 1901. Type citation. Nossi-bé,forêtduLoucoubé,mars 1851,Boivin (hb. Mus.Par.). Type specimen.Nossi-bé,forêtduLoucoubé,mars 1851, Boivin s.n. (BM000555467–Hb.Bescherelle! – lectotype,selected here ). Calymperaceaeand Orthotrichaceaeinthe Mascarene Islands33
Syrrhopodon nossibeanus var .borbonicus Renauld &Cardot, Bull. Soc. Roy.Bot.Belgique 33:117.1895. Type citation. Bourbon,secus rivulos(rev. Rodriguez). Type specimen. Bourbon [Réunion], Rodriguezs.n. (PC0105626 –Hb.Renauld! – lectotype,selected here ;isolectotype –“Exherb.F.Renauld”! – PC0105623)
Discussion – Calymperestaitense isdistributed throughout the Palaeotropics,with its most typicalformoccurring in Asiaand Oceania.However,atypicalforms predominateatthe westernextreme of its geographicalrange in Africa and the East AfricanIslands.Forexample,aformrepresented bytwocollectionsfrom the islandsof Mahé and Silhouetteinthe Seychelles( Norkett 17947&17790 ,BM) (“Seychellesform”) isunusualinpossessing leaveswithaparticularlynarrowband of hyaline cellsforming the margin in the proximalleafbase(1rowof cellsbroad). Intypicalcollectionsthisfeatureisconsistently2-5 rows of hyaline cellsbroad. Ofall the collectionsof Calymperestaitense examined from the East Africanislandstwospecimens( Pócs&Kiss 9450/FK &9450/EM,EGR)from MangabeinNE Madagascar(“Mangabeform”) havefeaturesclosest tothoseof the typicalform,differing onlyin the possession of particularlyshort,broad leaves.Thereisinsufficientreason tojustifyattaching infraspecificnamestomost of the novel formsof C.taitense occurring in the westernIndianOceanregion. However,the most significantof theseformswasdescribed byMüller(1876)asa distinctspecies, Syrrhopodon pachyloma Hampe ex Müll. Hal. Later,Bescherelle (1880)assigned the name S.nossibeanus Besch. tothe same formbut based on a differenttype specimen. Reese(1987)placed the latername, C.nossibeanus,in synonymywith Calymperestaitense proposing thatthe specieswas “indistinguishable from generalrunof Calymperestaitense ”. Ellis(1995) recognised Syrrhopodon pachyloma Hampe ex Müll. Hal. and S.nossibeanus Besch. asrepresenting adistinctformwithin Calymperestaitense but did not assign itaformalinfraspecificrank. Subsequently,ithasbecome evidentthatthis unusual“pachyloma ”formoccurs in amoreorless discretegeographicalareaand hassufficientconsistentlydistinctivefeaturestojustifyits recognition asaformal subspecieswithin C.taitense . Anobscuretaxon, Syrrhopodon nossibeanus var. borbonicus was properlydescribed from Réunion byRenauld &Cardot(1895),but first published asa nomen nudum (Renauld,1891). Thisvariety hasbeen largelyoverlooked and omitted from subsequentliterature,but its type and authenticmaterialisidentical tocollectionsof Calymperestaitense subsp. pachyloma . Inleavesof Calymperestaitense subsp. taitense the distalhyaline base incorporatesadifferentiated intramarginalrib(teniole) of thick-walled linearcells; anarrowunistratoseband of small chlorophyllosecells(extrateniolarlamina)forms the leafmargin (Fig. 2). The latterisabout 12.5-35µmbroadand usuallyformed by 2-7rows of cells.The chlorophylloselaminain the leaflimbisunistratose(Fig. 4) withthe cellsin surfaceviewmostly(3-)5-7.5(-12) × (3-)5-7.5 µm. In Calymperestaitense subsp. pachyloma ,thesefeaturesconsistently differfrom thoseofthe type subspecies.The extrateniolarlaminain the hyaline basemostlyreaches50-75µmbroad(Fig. 5),and isformed byaround 10-15 rows of cells.The chlorophylloselaminaismostlybistratose(Figs7,8) withcellsin surfaceview(2.5-)3-5 × (2.5-)3-5 µm. Thissubspeciesoccurs in Madagascar,the Mascarenesand Tanzania.Apublished recordof C.taitense from Zaire(Orbán, 1995) represents C.afzelii Sw.All specimensfrom Réunion examined forthis studybelong tosubsp. pachyloma .Collectionsof C.taitense from eastwardofthe Mascarenes(e.g. Thailand,AndamanIslands,Philippines,Sarawak,New 34J.Wilbraham&L.Ellis
Guinea, etc.) intoOceania, havethe featuresof the type subspecies.Most collectionsof C.taitense subsp. pachyloma havebeen sampled from shaded rock in forest between 200-1390malt.The type subspeciesisknownfrom abroader range of habitats.Occurring in damp,shaded situationsin forests,itissaid tobe morecommon atloweraltitudes(Eddy,1990; Ellis&Tan,1999),forming tufts and mats on tree boles,exposed roots,rock(frequentlycollected from limestone), rotting wood and compacted earth.
Specimensexamined Calymperestaitense (Sull.) Mitt.subsp .taitense . ANDAMAN ISLANDS: Kurz s.n. (BM000518458,holotype of C.andamense ). THAILAND: Süd-Thailand,prov.Krabi, NaturparkThanBok Koroni bei AoLuk,21July1994, Schäfer-Verwimp &Verwimp 16178 (BM000518457). PHILIPPINES: PalawanIsland,ApuranBarangay,DaanSitio,vicinity of MtTinikbasan,27 April 1992, Tan92-243 (BM000726628,FH). SARAWAK: 1888, Everett E510 (BM000851681). Gunong MuluNationalPark,around Sungei BerarCamp,SWof BukitBerar,14 March1978, Jermy13788 (BM000517791). Gunong MuluNationalPark, MelinauRiver,8August 1992, Stevenson 69 (BM000728683). INDONESIAN BORNEO: SKalimantan,Tabalong District,1°36’0’’S, 115°31’27’’E, 3April 2000, EllisK5.14.00 (BM000725510). PAPUA NEW GUINEA: Sepik District,between Sumo and Mafoka, 4July1961, Darbyshire&Hoogland 8067 (BM000919251). NEW IRELAND: DanfuValley, February 1970, Eddy6328proparte (BM000517794). SOLOMON ISLANDS: Russell Island, Lingatu,December1935, Levers.n.(BM000518486). FIJI: Labasa, 8July1923, Greenwood 503 (BM000518484). SOCIETY ISLANDS: Tahiti,1838-1842, WilkesExpedition s.n. (BM000518474,isotype of Syrrhopodon taitense [ ≡ Calymperestaitense subsp. taitense ]). Calymperestaitense “Seychellesform”:SEYCHELLES: Mahé,Ridge of Brulee, 29November1973, Norkett 17947 (BM000555466);Silhouette,forest belowCorgate, 11 November1973, Norkett 17790 (BM000555465). Calymperestaitense “Mangabeform”: MADAGASCAR: Antongil Bay,Nosy Mangabe Island, Pócs&Kis9450/FK &9450/EM (EGR). Calymperestaitense subsp. pachyloma (Hampe ex Mull. Hal.) L.T.Ellis TANZANIA: Uluguru Mts.MtTumbako,,5February 1973, Pócs&Lungwecha6875/E (EGR). COMORES: Mayotte, Marie s.n. (BM000555470)and Marie 123 (BM000555471). Mayotte,Magi M’Bien,August 1880, Marie 166 (BM000555468). MADAGASCAR. Tamatave,25October1972, Crosby&Crosby6692 (EGR). Tamatave,Ivaloina, 17February 1977, Tixier10454 (EGR). MasoalaPeninsula,9-11 September1994,[Pócs& Kis ] 9447/FB, 9447/FC & 9447/FA (EGR). Antongil Bay,8,10September1994, Pócs&Kis 9446/EX (EGR). ToamosinaProv.Lohatanjon Peninsula, 12-18 August 1998, Pócs&Szabo 9871/BE (EGR). ToamasinaProv.MananaraNordBiosphereReserveand NationalPark , 14-16August 1998, Pócs&Szabo9877/BS (EGR). RÉUNION: StePhilippe,1891, Rodriguezs.n. (Hb.Bescherelle,BM);same locality,1889, Rodriguezs.n. (PC01056254); same locality sine dato , Rodriguezs.n. (PC01056255). 14 km Wof SteAnne:Ravine d’Etang. AnnetteFalls,21°5’20’’S, 55°38’10’’E, 29August 1994, Orbán9434/C (EGR). Takamakagorge 13km Wof SteAnne,around Cascade de l’Arcen Ciel. 21°4’35-45’’S, 55°37’10-40’’E, 30 August 1994, Orbán9436/CF (EGR). Grand Etang,27 September1997, Arts RÉU53/36 (BR-BRYO 316692-84;BM000555464). ForêtduBrulé,atthe Cascade Maniquet,Boisde Couleur,3November1998, Arts RÉU70/62 (BR-BRYO 316693-85). AboveLa Montagne,routeD41,atthe junction toStBernard,forest pathtoIletà Guillaume,Grid:A3.e5,5November1998, Arts RÉU80/32 (BR-BRYO 316694-86). Valley of the rivièreLangevin from Grand GalettoChap. duCapBlanc, Grid:F6.b5,7November 1998, Arts RÉU90/15 (BR-BRYO 316695-87). CircuitdesRavenales,ca1kmwest- northwest of Chemin de Ceinture,grid:C7.d2,10November1998, Arts RÉU97/20 proparte (BR-BRYO 316662-54). Gorge de laRivièreduMat,Brasde Caverne forest pathtothe Cascade Blanche,Grid:B6.e1,11 November1998, Arts RÉU101/08 (BR-BRYO 316696-88). Plaine desGrégues,Grid:F6.d2,21February 2000, Arts RÉU131/41 (BR-BRYO 316697-89) and RÉU131/54 (BR-BRYO 316698-90). Calymperaceaeand Orthotrichaceaeinthe Mascarene Islands35
Figs1-7. Calymperestaitense (Sull.) Mitt.subsp .taitense (1-4) and C.taitense subsp. pachyloma (Hampe) L.T.Ellis(5-7). 1. Semi-diagramofleaf. 2,5. Detail of distalhyaline leafbase. 3,6. Costain chlorophylloselimbin cross-section. 4,7. Chlorophylloselaminaand marginalribin cross-section. (2from Darbyshire&Hoogland 8067,BM;3,4from Everett E510 ,BM;5from Orbán9434/C ,EGR;6,7from Orbán9436C/J ,EGR).
Calymperesafzelii Sw. ZAIRE: 20 km Nof Kisangani nearthe village of Bawombi, 11 September1977, Lisowski 50339 (EGR). Calymperespallidum Mitt. (newtoRéunion) Calymperespallidum (Mitten,1879) iswidespreadinboththe New World and Old World tropicsand hasbeen recorded from severalofthe islands in the westernIndianOcean,including Madagascar,Seychelles,and Rodriguesin the Mascarenes.Its presenceonRéunion,newlyrecorded here,isnotunexpected. The leavesin thisspeciespossess featuresvery similartothoseinthe pantropical species C.erosum Müll. Hal.,but differin thatthe costalacksstereids(Fig. 10) and hasaninflated appearance. Thisspeciesisdescribed and illustrated byEllis (2007). 36 J.Wilbraham&L.Ellis
Figs8-10. Calympereshispidum Renauld et Cardot(8), C.couguiense Besch. (9) and C.pallidum Mitt.(10). 8,9. Apicesof gemmiferous leaves(gemmaebroken off). 10. Cross-section of leaf, chlorophylloselimb.(8from Orbán9357/F ,EGR;9from Greenwood 474 ,BM; Arts RÉU152/13 , BR).
Specimen examined RÉUNION: BoisBlanc, sitedelaSource,Grid:E8a5,26 Feb.2000, Arts RÉU 152/13 (BR-BRYO 316679-71). Calymperescouguiense Besch. Calymperescouguiense ,described from NewCaledonia(Bescherelle, 1873)and largelyoccurring in the islandsof Oceaniaand in northeastern Australia(Ellis,2002),hasbeen erroneouslyrecorded forMauritius (Reese& Stone,1987)and the Seychelles(Orbán,1995a). The collectionsupon whichthe recordsareapparentlybased belong to C.hispidum Renauld et Cardot,aspecies withawell established distribution in the East AfricanIslands(Ellis,1988; O’Shea, 2006). Calymperescouguiense and C.hispidum havedimorphicleaves, gemmiferous and non-gemmiferous.The dimorphismisparticularlyextreme in C.couguiense in whichthe gemmiferous leavesarelinearand exserted from the shoot,well abovethe lingulatenongemmiferous leaves.Shoots and leavesin C.hispidum cansometimeshaveasimilarappearancetothosein C.couguiense (e.g. asin Seychelles, Orbán9357/F ),but aremost plainlydistinguishable bythe formofthe apicesin theirgemmiferous leaves.Ingemmiferous leavesof C.couguiense ,the laminarunsintothe costabelowits apexand the apicalcells of the leafaredrawnout asblunt,finger-like projectionsthatformadense fringe atthe baseofacrown-like clusterof gemmae(Fig. 9). Ingemmiferous leavesof C.hispidum ,anarrowband of laminacompletelyborders the costal apex(Fig. 8),and gemmaeareproduced in aradiating clusteronlyfrom its ventralsurface.
Specimensexamined Calymperescouguiense Besch. FIJI: Interiorof VanuaLevu,25December1922, Greenwood 474 (BM000675201,holotype of C.marginatum Dixon [= C.couguiense ]). Calymperaceaeand Orthotrichaceaeinthe Mascarene Islands37
Calympereshispidum Renauld et Cardot MAURITIUS. Balfour s.n. (NY). SEYCHELLES: Praslin,Grand Fond,summitatthe NW part,4°18’15’’S55°42’21’’E, 31August 1993, Orban9357/F (EGR).
SYRRHOPODON PROLIFER-LIKE TAXA (Calymperaceae) Ofthe several Syrrhopodon prolifer -like taxa, withlinear-ligulate, limbateleaves,thatoccur on Réunion,four havebeen particularlymisunderstood orover-looked – S.apertifolius Besch., S.hispidocostatus Renauld et Cardotwith anewlyrecognised subspecies(see below),and the newlyrecorded S.dimorphophyllus L.T.Ellis.Manycollectionsof thesetaxafrom Réunion have been misidentified variouslyas Syrrhopodon spiralis Renauld et Cardot, S.prolifer Schwägr.var. seychellarum Orbán[= S.albidus Thwaites et Mitt.subsp. integrifolius (E.B.Bartram) L.T.Ellis], S.prolifer var. prolifer and S.prolifer var. acanthoneuros (Müll. Hal.) W.D.Reese(see “specimensexamined”). Thereis presentlyno evidenceforthe occurrenceoftheselattertaxaon Réunion (Ellis, 2005;Ellis&Wilbraham,2008).
Syrrhopodon hispidocostatus Renauld et Cardot and S.apertifolius Besch. Syrrhopodon hispidocostatus ismost likelytobeconfused withits sibling species– S.apertifolius.Bothtaxaaredescribed,illustrated and discussed byEllis (2005). Leavesin S.hispidocostatus haveanextensivelyspinycosta, characteristicallywitharowof spinesalong eitherside of the dorsalsurface (dorsal-lateralcostalspines,Fig. 19) and often withtall (sometimesmultifid) spineson the ventralsurface(Fig. 20). The cellsof the chlorophylloselaminaare relativelylarge (6-17.5(-20) × 7-12.5 µm) and mostlypossess tall multifid projections(Figs20-22)through whichthe underlying cellsremain visible. Inmost leavesof S.apertifolius the costahasmostlysimple spinesonlytowardsthe leaf apex; proximally,ventralordorsal-lateralprojectionsfrom the costatend tobe short and multifid. The cellsof the chlorophylloselaminaappearmuchsmaller, (5-)7-12.5(-15) × (5-)7-10(-12.5) µm) thanthosein S.hispidocostatus;dorsallyand ventrallytheyproducerelativelysmall,low,multifid papillae(Fig. 23)thatareso denselydistributed astoalmost entirelyobscurethe underlying cellsand make the laminaratheropaque. Syrrhopodon hispidocostatus and S.apertifolius havenotbeen found outside of Madagascar,Mauritius and Réunion. Ellis(2005) suggested that S.hispidocostatus appeared generallytopreferhabitats athigheraltitudes.This proposition isundermined bythe severalcollectionsmade from lowland localities on Réunion byArts (mostlyheld in BR). Syrrhopodon hispidocostatus iswell distributed in Réunion whereithasbeen recorded from tree trunksand stumps in forest at110-1200 malt.Itappears tobeatleast ascommon as S.apertifolius withwhichitsometimesoccurs. Syrrhopodon apertifolius prefers lowland habitats, having been most commonlycollected in Réunion on treesand rotten stumpsin forest between 40and 340malt.
Specimensexamined Syrrhopodon hispidocostatus Renauld et Cardot MAURITIUS: Petrin HeathNature Reserve,(loc.5),3October1962, G.Een M198 (S, originallyidentified as Syrrhopodon prolifer ). RÉUNION: CircuitdesRavenales, ca 1kmwest-northwest of Chemin de Ceinture,Grid:C7.d2,10November1998, Arts RÉU97/12proparte (BR-BRYO 319366-42), RÉU97/57 (BR-BRYO 319369-45) and RÉU97/60 proparte (BR-BRYO 319370-46). Gorge de laRivièreduMat,Brasde Caverne forest pathtothe Cascade Blanche,Grid:B6.e1, 38J.Wilbraham&L.Ellis
Figs11-23. Syrrhopodon hispidocostatus Renauld et Cardotsubsp. artsii L.T.Ellis(11-17), S. hispidocostatus subsp. hispidocostatus (18-22)and S.apertifolius Besch. (23). 11,12,18. Leaves. 13,19. Dorsal-lateralcostalprojectionsin proximalleaflimb. 14,20. Chlorophylloselimbin cross-section. 15,21. Cellsof chlorophylloselaminaand papillaeinsurfaceview. 16,17,22, 23. Chlorophylloselaminain cross-section. (11 from Arts RÉU97/42 ,BR;12-16from Arts RÉU146/91;17from Arts RÉU97/12 ,BR;18,19,21from Arts RÉU102/168 ,BR;20,22 from Arts RÉU102/152 ,BR;23from Arts RÉU54/13 ,BR). Calymperaceaeand Orthotrichaceaeinthe Mascarene Islands39
11 November1998, Arts RÉU101/16proparte (BR-BRYO 319372-48). From the “Airede Pique-niquedelaPlaine desLianes”tothe “Sitetouristiqued’Eden”,Grid:C6.a3, 12November1998, Arts RÉU102/81 (BR-BRYO 319373-49). ForêtDomaniale de Bras Panon,atthe “Sitetouristiqued’Eden”,aformerthee plantation,Grid:C6.a3,12November 1998, Arts RÉU102/154 (BR-BRYO 319374-50), RÉU102/168 (BR-BRYO 319375-51) and RÉU 102/170 proparte (BR-BRYO 319376-52). L’Oasis,west of StBenoit,Grid: C7.c1,15 November1998, Arts RÉU110/05proparte (BR-BRYO 319377-53). Tremblet, Grid:F8.c3,8December1998, Arts RÉU92/07 (BR-BRYO 319363-39); Arts RÉU92/15 (BR-BRYO 319364-40). BasseVallée forest roadin Cryptomeria forest,Grid:F7.e3, 24February 2000, Arts RÉU144/12 (BR-BRYO 319379-55). BasseVallée,northofdeGite de BasseVallée forest roadin Cryptomeria forest,Grid:F7.e3,24February 2000, Arts RÉU144/67 proparte (BR-BRYO 319380-56)and RÉU144/69 (BR-BRYO 319381-57). ForêtMourouvin,abovethe “troisciterne” (waterreservoirs),bushvegetation,Grid:D8d1, 26 February 2000, Arts RÉU150/61 (BR-BRYO 319382-58). LeDimitile,Sentierde Dimitile parLa Chapelle,Grid:E4.b5,along pathin Erica forest,1March2000, Arts RÉU162/43 (BR-BRYO 319384-60). Syrrhopodon apertifolius Besch. RÉUNION: Rodriguezs.n.(exHb.Möllercom. Brotherus)(S, originallyidentified as S.prolifer )and Rodriguezs.n.(exHb.Rothcom. Brotherus)(S, originallyidentified as S.prolifer ). SentierduTrembletaunorddeSte Philippe,forêtprimairehumide d’egradée,10September1971, Onraedt71.R.9453 (BR- BRYO 234495-46). Ravine duTremblet,forêtdomaniale de StePhillipe,10September 1971, Onraedt71.R.9353 (BR-BRYO 234496-47)and 71.R.9353 (Hb.Bizot19297) (PC0092698,BR-BRYO 052400-20,originallyidentified as S.spiralis ). Lavesrécentedu Grand Brule,12April 1972, Cadet435 (Hb.Bizot19420)(PC0092700,originallyidentified as S.spiralis ). Grande Brule,27 April 1972, Cadet458 (Hb.Bizot19421) (PC0092699, originallyidentified as S.spiralis ). RéserveNaturelle MareLongueonthe slope of Piton de laFournaisevolcano,5September1994, Vojtko &Vojtko 9432/CB (RÉU, originally identified as S.prolifer var. seychellarum ). StePhilippe,RouteforestièredeMareLongue’ RéserveNaturelle,7March1997, Arts RÉU11/50 (BR-BRYO 319360-36).). StPhilippe, RéserveNaturelle de MareLongue,forest trail,7March1997, Arts RÉU11/87 (BR-BRYO 319361-37). La Côteaux vents,LeGrand Brulé,Ravine duTremblet,28September1997, Arts RÉU54/13 (BR-BRYO 319362-38). Forêtde BoisBlanc, Grid:E8.b4,8November 1998,Arts RÉU93/18 (BR-BRYO 319365-41). CircuitdesRavenales, ca 1kmwest- northwest of Chemin de Ceinture,Grid:C7.d2,10November1998, Arts RÉU97/60 proparte (BR-BRYO 319370-46)and RÉU97/69 (BR-BRYO 319371-47). Gorge de laRivièredu Mat,Brasde Caverne forest pathtothe Cascade Blanche,Grid:B6.e1,11 November1998, Arts RÉU101/16proparte (BR-BRYO 319372-48). L’Oasis,west of StBenoit,Grid:C7.c1, 15 November1998, Arts RÉU110/05proparte (BR-BRYO 319377-53). Forest pathfrom routeN2toPointedeLa Table,Grid:F8.e4,28February 2000, Arts RÉU156/04 (BR-BRYO 319383-59). Syrrhopodon hispidocostatus subsp. artsii L.T.Ellis, subsp. nov. Subspecieshaecsubspeciei typicosimilissed papillislaminaehumilibus recedit. Type specimen.Réunion. Leserré,northofSaint-Joseph,Grid:F6.d4,25February 2000, Arts RÉU146/91 ( holotype -BR-BRYO 319385-61). Shoots mostly0.5-2.0cmtall. Leaves<3-5 mm long,withanarrowlysubelliptical semisheathing hyaline baseand alineartoligulatechlorophylloselimb.Costa ending just short of leafapex; dorsalsurfacemostlysmoothand formed by stereids,but generallyspinosenearleafapex,alateralrowof chlorophyllose cellsoften projectasspinesormultifid protuberancesalong eitherside of the costa;ventralsurfaceformed bystereidsorapartiallayerof projecting, multipapillosechlorophyllosecells(projectionsoften morespine-like nearleaf apex). Hyaline laminasharplydefined,occupying less thanathirdtohalf of the leaflength. Cellsof chlorophylloselaminapolygonalwith4-6sidesorrounded, 40J.Wilbraham&L.Ellis mostly7.5-17.5 × 7-10(-12.5) µm,slightlylongerthanbroadtoslightlybroader thanlong,dorsallyand ventrallymultipapillosewithlow,knobblymultifid papillae. Leafmargin differentiated from leafbasetoapex; distallyfrom around distalhyaline baseformed byastrand of stereids,mostlyentirebut usually toothed nearleafapex.Gemmaefusiform,uniseriateproduced from dorsaland ventralsurfacesof costalapex.Sporophytesnotseen.
Discussion –The leavesof Syrrhopodon hispidocostatus subsp. artsii most consistentlydifferfrom thoseofthe type subspeciesin theirornamentation. This subspeciespossessescellsin the chlorophylloselaminacomparable in size (mostly7.5-17.5 × 7-10(-12.5) µm) tothoseintypical S.hispidocostatus but which lackthe tall,multifid projectionsfound in the latter.Instead,the laminalcells bearsmall,lowmultifid papillae(Figs15-17). The ornamentation of the costaisa less constantfeaturedistinguishing subsp. artsii from the type subspecies.In some leavesin subsp. artsii,dorsal-lateralspinesoccur extensivelyalong the costa(asin the type subspecies),but in manyleavesthe equivalentprojections, atleast in the proximalchlorophylloselimb, often tend tobeshort and multifid ratherthanspine-like (Figs13,14). Inaddition,chlorophyllosecellsatthe ventralsurfaceofthe costaoften projectaslowmultifid papillae,whereasin the type subspecies,thesecellsmorefrequentlyprojectastall simple ormultifid spines.Finally,leafshape isavariable featureinsubsp. artsii;in some specimens (including the type collection) the leavestend towardsbeing moreligulatethan linear(Figs11,12),the latterisamorecommon shape in the type subspecies (Fig. 18). The leavesin Syrrhopodon hispidocostatus subsp. artsii arealso comparable to,but areeasilydistinguished from,thosein S.apertifolius.Inthe latter,the chlorophylloselaminaisrendered opaquebydenselycrowded multifid papillae;the tinyunderlying cellsseldom reachmorethan12.5 µm long. In contrast,the chlorophylloselaminain subsp. artsii ispellucid,withsmall multifid papillaethatarewell spaced out; the underlying cellsareeasilyvisible and can reach17.5 µm long.Syrrhopodon albidus subsp. integrifolius (E.B.Bartram) L.T.Ellisisarelated moss occurring in the Seychellesand Malesia(Ellis,2005), but canalsobedistinguished from S.hispidocostatus subsp. artsii bythe sizeofits laminalcells.The cellsforming the chlorophylloselaminain S.albidus subsp. integrifolius areconsistentlymuchlargerthanthosein S.hispidocostatus subsp. artsii,withmanyreaching 20->25µmlong. Syrrhopodon hispidocostatus subsp. artsii appears tobeendemicto Réunion. Itoccurs between 650and 920 malt.,apparentlypreferring earthy substratesand hasoften been collected in association withthe type subspeciesand S.apertifolius,whichareotherwiselargelycorticolous mosses.
Specimensexamined Syrrhopodon hispidocostatus subsp. artsii L.T.Ellis RÉUNION: 14 km Wof SteAnne, Ravine d’Etang,29August 1994, Orbán9434/CH &9434/ CG (REU, originallyidentified as S.prolifer var. seychellarum ). CircuitdesRavenales,ca1kmwest-northwest of Chemin de Ceinture,Grid:C7.d2,10November1998, Arts RÉU97/12proparte (BR-BRYO 319366- 42), RÉU97/42 (BR-BRYO 319367-43)and RÉU97/47 (BR-BRYO 319368-44). Forêt Domaniale de BrasPanon,atthe “Sitetouristiqued’Eden”,aformerthee plantation,Grid: C6.a3,12November1998, Arts RÉU 102/170 proparte (BR-BRYO 319376-52). L’Oasis, west of StBenoit,Grid:C7.c1,15 November1998, Arts RÉU110/05proparte (BR-BRYO 319377-53). BasseVallée,northofdeGitedeBasseVallée forest roadin Cryptomeria forest,Grid:F7.e3,24February 2000,Arts RÉU144/67 proparte (BR-BRYO 319380-56). Calymperaceaeand Orthotrichaceaeinthe Mascarene Islands41
Figs24-28. Syrrhopodon dimorphophyllus L.T.Ellis. 24. Non-gemmiferous leaf. 25. Gemmiferous leaf. 26. Apexof gemmiferous leaf(gemmaebroken off). 27. Chlorophyllose laminain gemmiferous leafincross-section. 28. Chlorophylloselaminain nongemmiferous leafin cross-section. (24-28from Arts RÉU142/74 ,BR).
Syrrhopodon dimorphophyllus L.T.Ellis (newtoRéunion) Syrrhopodon dimorphophyllus, described from Malawiand Madagascar (Ellis,2005),isnewlyrecorded forRéunion Island. The speciesappears tohave been often overlooked ormisidentified,but nine collectionshavenowbeen determined from diverselocalitieson the island. Manytaxain the Calymperaceae, including S.dimorphophyllus,characteristicallyexhibitleafdimorphism,with leavesproducing gemmaebeing significantlymodified relativeto“normal” leaves. Asin some othertaxa, shoots of S.dimorphophyllus usuallypossess some leaves withaformtransitionalbetween thatof theirgemmiferous and nongemmiferous leaves.Insome specimens,the most characteristicextreme expression of the gemmiferous leafmaynotalways beattained. Thiscancausedifficulty withthe identification of material. The gemmiferous leavesin Syrrhopodon dimorphophyllus,attheirbest developed,arelinearwithamodified gemmae-bearing apex(Fig. 25);the cellsof the chlorophylloselaminaarerelativelylarge and spinosethroughout the leaf limb(Fig. 27). Non-gemmiferous leavesareshorter,ligulate(Fig. 24) and have smallerlaminalcellswithlowpapillae(Fig. 28). Leaveswitha“transitional” form aremodified from the lattertovarious degrees.Inmost collectionsof S.dimorphophyllus examined from Malawi,the most extremelymodified formof gemmiferous leafiswell represented. However,in some collectionsexamined from Réunion (e.g. Onraedt73.R.8777),gemmiferous leavesoften appearless completelydeveloped. Although theseleavescanbelinearwithadistinct gemmiferous apex,the cellsof the chlorophylloselaminatend tobesmall with 42J.Wilbraham&L.Ellis lowpapillae,and onlyrarely,become largerand spinosenearthe leafapex.In otherspecimens(e.g. Arts REU142/74 ),atleast afewgemmiferous leavesare sufficientlydeveloped topossess spinoselaminalcellsthroughout the leaflimb. Fortunately,non-gemmiferous leavesin S.dimorphophyllus possesfeaturesin the chlorophylloselaminathatdistinguishthem from the leavesof the otherlocaltaxa in the S.prolifer -like group. In S.dimorphophyllus,the cellsof the chlorophyllose laminain nongemmiferous leavesarerelativelytiny,on average 5-7.5 µm diam. in surfaceview.Eachcell possessesseveralsmall,low,knobblypapillae(dorsally and ventrally),whichtend toleanoutwardsoverthe cell walls.Inotherlocal S.prolifer -like taxa, the cellsof the chlorophylloselaminaaregenerallylarger, seldom less than7.5 × 7.5 µm,and aremostlymorecoarselypapillose,often with eachcell appearing topossess asingle,large multifid papilla(see Ellis,2005). Syrrhopodon dimorphophyllus occurs in Malawi,Madagascarand Réunion. Throughout its range ithasbeen collected between 1200 and 2375m alt.,mostlyon the trunksof treesand tree fernsin humid,shaded situationsin forest.
Specimensexamined Syrrhopodon dimorphophyllus L.T.Ellis MALAWI: Mulanje,Mulanje Mountain, LichenyaRiver,15°58’50’’S35°33’19’’E, 28June 1991, Wigginton 1683a (BM000745235, holotype of S.dimorphophyllus). RÉUNION: Plaine desCafres,forêtde Bébour, 23 December1973, Onraedt73.R.8777 (BR-BRYO 234494-45,originallydetermined as S.spiralis ). Forêtde Bélouve,sentierde laTamaraies,SWof the GitedeBélouve,grid: C5.d3,23 February 2000, Arts RÉU 142/74 (BR-BRYO 319378-54). Plaine desFougères, trackofthe GR towardsthe Piton BéMassoune,16March2004, Ah-Peng R49-B2proparte (PC). Forêtde Belouve,Troude Ferpath,21°3’40’’S55°32’10’’E, 12September2008, Ellis &WilbrahamR08-159 (BM000890375),Forêtde Belouve,Sentierde L’Ecole Normale, 21°4’39.9’’S55°32’34’’E,13September2008, Ellis&WilbrahamR08-202 (BM000890379) & R08-209 (BM000890377). Plaine desChicots,ReserveNaturelle Roche Ecrite,20°57’7.4’’S 55°26’13’’E, 14 September2008, Ellis&WilbrahamR08-233a (BM000890380). Salazie, routedePiton Marmite,21°3’40’’S, 55°27’3’’E, 16September2008, Ellis&WilbrahamR08- 263 (BM000890376); R08-264 (BM000890378). Syrrhopodon vardei L.T.Ellis (newtoRéunion) and S.pulcher W.D.Reese Syrrhopodon vardei,until nowregarded asarareMadagascanendemic, isherenewlyrecorded forRéunion Island. Unfortunately,the collection ( Ellis& Wilbraham314b ,BM)consists of asingle shoot.Itwasfound in association with S.gardneri and S.hispidocostatus on the shaded underside of roots on avertical stream-side bank at1220 malt. Syrrhopodon pulcher W.D.Reesewasdescribed from Seramand SulawesiinSE Asia(Reese etal.,1991). Orbán(1995b)cited twocollectionsas newrecordsof thisspeciesforMadagascar[“ Pócs,Magill &LaFarge England” [ Magill &Randrianasolo] 90114/AS & 90114/AV (EGR)].The presenceofthe speciesin Madagascarwasreiterated byOrbán(2007),but subsequently,he correctlyredetermined and annotated the previouslycited specimensas Syrrhopodon vardei.Thereisno evidenceforthe occurrenceof S.pulcher in Madagascar. S.pulcher and S.vardei aresuperficiallyvery similar,but theirleavesare easilydistinguished underthe microscope. In S.pulcher ,the cellsof the chlorophylloselaminaarestronglyprotuberantfrom the ventralleafsurface,each withanacute,simple ormultifid papilla.Dorsallythe cellsof the laminaarenot protuberant,but smoothorunipapillose. Amarginalrib(stereome) in the chlorophylloselimbisthickand entirelycomposed of stereids.Incontrast,the Calymperaceaeand Orthotrichaceaeinthe Mascarene Islands43 cellsof the chlorophylloselaminain S.vardei areflattobarelyprotuberantand eachiscovered dorsallyand ventrallywithseveralsmall,knobblypapillae. The marginalstereome often includesamedianlayerof guide cells,partiallydividing dorsaland ventralgroupsof stereids(see Ellis,2005).
Specimensexamined Syrrhopodon vardei L.T.Ellis MADAGASCAR.Antsiranana.ReserveIntergrale Nationale de Matojezy.Elfin forest –onridge between Camp II and III at1000-1400(-1800)malt., 26-28March1990, Magill &Randrianasolo 90114/AS &90114/AV (EGR). RÉUNION. Plaine desFougères,20°58’30’’S55°31’13’’E, 19 September2008, Ellis & WilbrahamR08- 314b (BM000890381).
SYRRHOPODON SUBG. PSEUDOCALYMPERES BROTH.(Calymperaceae) Syrrhopodon subg. Pseudocalymperes includesadistinctivegroupof speciespreviouslyreferred tothe genus Calymperopsis (Müll. Hal.) M.Fleisch. The group’sgenericstatus wasdiscarded byReese(1978) and its correctname established byReese(1993). Reese(1994a, 1994b)revised the taxadescribed from West Africa, and laterReese(1995) usefully,but briefly,rationalised the taxain the entiresubgenus. Mossesof Syrrhopodon subg. Pseudocalymperes characteristically possess erectshoots withleavessomewhatresembling thoseof Mitthyridium .In some speciesthereisastrong degree of leafdimorphism,gemmae-bearing leaves aresomewhatmodified relativetonormalleaves.The latterareusuallylingulate and in most specieshaveadifferentiated margin formed byanarrowstrand of stereids(stereome). Gemmae-bearing leavestend tobeshorterand broader,in some casesmoredeltoid thanlingulate. Theymostlyformasmall cupatthe apicesof shoots (comae)1 .Long,filamentous,uniseriategemmaearecommonly produced in afan-shaped weftfrom the ventralleafsurfacedistallyof the hyaline lamina. Three speciesof Syrrhopodon subg. Pseudocalymperes havebeen recorded forRéunion Island: S.africanus (Mitt.) Paris, S.crenulatus (P.Tix.) W.D.Reese(Arts,2005;O’Shea, 2006)and S.parasiticus (Brid.) Paris(Müller, 2002). The recordfor S.africanus appears tobecorrectlydetermined,but as demonstrated below,the recordof S.crenulatus iserroneous,and the material upon whichitisbased hasfeaturessimilartothoseof S.flexifolius Mitt.The proposed recordof S.parasiticus from Réunion hasnotbeen available for examination. Syrrhopodon africanus (Mitt.) Paris Syrrhopodon africanus asrecognised byReese(1995) hassomewhat variable features.Most collectionsfrom continentaleasternAfrica (including Tanzania, Last s.n. (BM, isotype of S.africanus)possess lingulateleaveswith broadlyacutetoobtuse(often apiculate) apices(Fig. 29);the marginalstereome and the costausuallyfail short of the leafapex.The margin of the leafapexis often papillose-crenulate,but sometimesentireorirregularlydenticulate(Figs31, 36,40). Atthe apexof some shoots agemmae-cup(coma)isformed bymodified
1 Newgrowthwithunmodified leavesoften developsfrom within acoma, sothatthe comano longerappears tobeapical. The newgrowthmayproduceagemmiferous apicalcomaof its own,whichinturnmaygiverise tofurthernewgrowth,ultimatelyproducing anothernewcoma.Inthisway,afterseveralseasonsof growth, groupsof gemmiferous and nongemmiferous leavescanappeartoalternatealong ashoot. 44 J.Wilbraham&L.Ellis
Figs29-40. Syrrhopodon africanus (Mitt.) Parisvar. africanus. 29,34. Non-gemmiferous leaves and 30,35. detailsof apices. 31,36,40. Leafmargin atapexand 32,37. in distalchlorophyllose limb. 33,38. Surfaceofchlorophylloselaminashowing papillae. 39. Cross-section of lamina (29-33 from Last s.n.,BM, isotype of S.africanus;34-39from Arts RÉU92/28 ,BR;40from Porley 269a ,E). Calymperaceaeand Orthotrichaceaeinthe Mascarene Islands45 gemmiferous leaves.Collectionsfrom West Africa in BM, annotated as S.africanus byReese,differfrom the East Africanmaterialinpossessing narrowerleaveswithnarrowlyacuteapicesand anexcurrentcosta, the marginal stereome usuallyreachesthe leafapex.Theselattercollectionsappearto representadistinctlocalformof S.africanus. Yetanothervariantof thisspecies, Syrrhopodon africanus var. mandrakensis (P.Tix.) W.D.Reese,wasoriginallydescribed from Madagascaras Calymperopsismandrakensis P.Tix.(Tixier,1992). Presentlyaccepted asaformal variety of S.africanus,its type material(Madagascar, Tixier11355 [“ 11335 ”], PC) hasbeen unavailable forexamination in thisstudy.Tixier(1992)and Reese(1995) distinguishits leavesfrom thoseofrelated taxabythe possession of acrenulate apicalmargin. Inthe context of the apparentvariability of leafapiceswithin S.africanus,thisfeaturemayseem of ratherdubious criticalvalue. However, Tixier(1992)describesthe cellsof the chlorophylloselaminain his C.mandrakensis ashaving adiameterof about 5µm,very muchsmallerthan thoseinmost othertaxa(including S.africanus)inthe subgenus Pseudocalymperes .Rediscovery of the type materialwill berequired toproperly assess the status of S.africanus var.mandrakensis . Arts (2005) recorded Syrrhopodon africanus forRéunion Island based on three collections– Arts REU92/28 (BM, BR), REU102/150 (BR)and REU70/43 (BR). Thesespecimensappeartorepresentanotherdistinctformofthe species. Theirfeatureslargelyagree withthoseofcollectionsfrom easterncontinental Africa.However,the leavesconsistentlydifferfrom thoseofthe latterin some featuresof the chlorophylloselaminaand leafapex.Forexample,the apicesof leavesin Arts’scollections(Fig. 35) tend tobeslightlymoreobtusethanthose found in most specimensof S.africanus.Moresignificantly,in S.africanus the cellsof the chlorophylloselaminaeachpossess afewsimple,lowpapillae,in nongemmiferous leavesthesecellsare(7.5-)10-20 × (7.5-)10-15 µm in surface view,while in gemmiferous leavestheyreach10-25(-27.5) × 10-20 µm. The cells of the chlorophylloselaminain Arts’scollectionspossess morestronglydeveloped papillae(especiallyon the dorsalsurfaceofthe lamina)(Figs38,39) and are generallyslightlysmaller.Innongemmiferous leavesthe cellsreach(7.5-)10-17.5 × (7.5-)8-12.5 µm,and in gemmiferous leaves10-20 × 8-15 µm. Considering the apparentplasticity of S.africanus (ascurrentlyrecognised),Arts’collectionsfrom Réunion probablyrepresentasmall,obtuseleaved formofthe specieswithwell developed laminalpapillae,and therefore,werecorrectlyidentified byArts (2005). Syrrhopodon africanus iswidespreadacross tropicalAfrica (O’Shea, 2006),and hasbeen collected on trunks,branchesand twigsof treesin forest between 230 and 1650malt.
Specimensexamined Syrrhopodon africanus (Mitt.) Paris SIERRA LEONE: KaseweForest Reserve, 27 February 1971, RichardsR7207 (BM000663242). NIGERIA: Benin,OkomuForest Reserve,9December1947, Richards3628 (BM000663243),17December1947, Richards 3681 (BM000663244) and 24December1947 ,Jones3894 (BM000663240). Agbadi village nearSapoba, 14 November1949, Keay&Meikle UD23 (BM000736191). CAMEROON: Dusén exsicc.no. 500 (BM000663246,PC0094867,isolectotypesof Syrrhopodon spurio- disciformis Dusén [= S.africanus, vide Reese,1994]). CONGO: Pengbe,3February 1914, Bequeart s.n. (BM000677677). UGANDA: Bushenyi,Kalinzu FR, nearNkombesawmill, 0°23’S30°5’E, Hodgetts U4557a (E). Bwindi NationalPark,2February 1996, Porely269a (E). TANZANIA: UsagaraMts., Last s.n.(BM000663239,isotype of S.africanus). RÉUNION: ForêtduBrulé,footpathtoCascade Maniquet,Boisde Couleur,3November 46J.Wilbraham&L.Ellis
1998, Arts RÉU70/43 (BR-BRYO 319331-07). Tremblet,Grid:F8.c3,8November1998, Arts RÉU92/28 (BM000678107; BR-BRYO 319332-08);ForêtDomaniale de BrasPanon: atthe “Sitetouristiqued’Eden”,aformerthee plantation. Grid. C6.a3,12November1998, Arts RÉU102/150 (BR-BRYO 319333-09).
Syrrhopodon flexifolius subsp. reunionensis L.T.Ellis , subsp. nov. Subspecieshaecsubspeciei flexifolio similissed costain basehyalinafolii incrassata, cellulissuperficialibus parietibus tenuibus differt. Type specimen. ForêtDomaniale de BrasPanon:atthe “Sitetouristiqued’Eden”, aformerthee plantation,Grid:C6.a3,12November1998, Arts RÉU102/192 (BR- BRYO 319335-11, holotype;BM000678105, isotype). Shoots curved toerect,0.5-2.0cmhigh,denselyleaved,oldergrowth brown,often withdensetufts of rhizoidsbetween leafbases,youngergrowth yellowishgreen. Leavesdimorphicasgemmiferous (modified) and non- gemmiferous (normal) leaves(modified gemmiferous leavesforming terminal gemmaecups(comae)). Normalleaveswhen dry incurved (somewhatappressed in youngergrowth) and spirallytwisting forming apointed shoottip,when moist erecttopatent,mostly2.5-4.0mm long,lingulatetoelliptical-lingulatewithan obtusetoobtuse-apiculateapex; consisting of semi-sheathing hyaline baseand a broadchlorophylloselimb(laminain distalleafbaseoften slightlybut abruptly broadening intolimb). Costarobust (especiallyin hyaline base),ending immediatelybelowleafapex,smooth,dorsalsurfacelargelyformed bystereids, ventralsurfaceindistalchlorophylloselimbalsoformed bystereids,but in proximalchlorophylloselimbsuperficialcellssometimesbecoming shortly rectangularwiththick,porosewalls,in hyaline base(mostly60-100 µm wide) cells becoming longerand broader,hyaline withthin,porosewalls(like thoseofcells in adjacenthyaline lamina). Chlorophylloselaminagentlyundulating;cellsin surfaceviewrounded toshortlyrectangular,chollenchymatous (onlydistinctly porosenearapexof hyaline lamina),(10-)12.5-25(-30) × (10-)12.5-15(-20)µm, smooth,in transversesection rectangular.Hyaline laminausuallysharplydefined, occupying less thanhalf of the leaflengthand withanacuteapex,most cellswith asuperficialporeatthe proximalend. Leafmargin slightlyundulate, differentiated from leafbasetonearapex; in leafbaseaflat,narrowunistratose ribof often orange-brown,thick-walled,porose,linearcells,in distalhyaline base ribbordered bymarginalstrand of stereidsthatcontinuesdistallyinto chlorophylloselimbending nearthe leafapex,strand largelyentiretodistantly toothed;apicalmargin formed bycellsof chlorophylloselamina, usuallyentire, sometimessparselytoothed. Modified “gemmiferous”leavesforming apical comae, ca 3mm long,similartonormalleavesexceptmoreconcaveand broadly lingulateand withamoreincrassatecosta, cellsof chlorophylloselaminaslightly larger,reaching >30 µm long. Gemmaeproduced ventrallyfrom aboveapexof hyaline laminatobeyond midleaf,developing from lateralcostaland juxtacostal laminalcells,filamentous,uniseriate,mostly400-650µm long,darkgreen to reddishbrown. Sporophytesnotseen.
Discussion –Apublished recordof Syrrhopodon crenulatus on Réunion Island (Arts,2005) wasbased on Arts RÉU102/192 (BR, BM). Using Reese(1995),the best available keyto Syrrhopodon subg. Pseudocalymperes ,thiscollection can indeed beidentified as S.crenulatus.However,neitherthe authorof the keynor the collectorof the specimen wasable toexamine the type materialof S.crenulatus;its whereabouts arestill amystery.Consequently,the truenature of the speciesisuncertain. Calymperaceaeand Orthotrichaceaeinthe Mascarene Islands47
Figs41-52. Syrrhopodon flexifolius Mitt.subsp .reunionensis L.T.Ellis(41-46), S.flexifolius subsp. flexifolius (47-50)and S.parasiticus (Brid.) Paris(51,52). 41. Non-gemmiferous leaves. 42,48. Leafapices. 46,47. Ventralsuperficialcellsof costain mid-hyaline leafbase 43,51.Cellsof chlorophylloselaminain surfaceviewand 45,50,52. in cross-section. 44,49.Leafmargin in surfaceview.(41-45 from Arts RÉU102/192 ,BR;46from Porley9728 ,BM;47from Allen 23571 , MO;48,50from Casado 585,MO;49 from Allen 23164 ;51,52from Swartz s.n. [ H2269 ], BM, isotype of S.parasiticus). 48 J.Wilbraham&L.Ellis
Comparing the featuresdescribed in the protologueof Calymperopsis crenulatus P.Tix.[= Syrrhopodon crenulatus](Tixier,1967)withthoseof Arts RÉU102/192 indicatesthatthe latterdoesnotbelong tothisspecies.Forexample, the cellsof the chlorophylloselaminain leavesof S.crenulatus aredescribed as “cellulesisodiamétriquesd’environ 15 µmuniesd’une papille àlafacedorsale” (notunlike thosein S.parasiticus,Figs51,52),whereasthe cellsof the chlorophylloselaminain Arts RÉU102/192 aremostlysignificantlylarger,and flat and smoothonbothdorsaland ventralsurfaces.Thisisanunusualfeatureinthe subgenus Pseudocalymperes .Inotherspeciescellsof the chlorophylloselamina areusuallypapilloseand/orprotuberant.Large,smooth,flatcellsareapparently characteristicof onlyone otherspecies, Syrrhopodon flexifolius from Central America.Thisspecieswasformerlyerroneouslyconsidered conspecificwith S.parasiticus but latterlyrecognised byReese(1993)asagood species. Unfortunately,shoots in the twoavailable duplicatesof Arts RÉU102/192 (BR, BM)arerelativelysparse,but thiscollection hasvery recentlybeen supplemented by Porley9728 (BM),aspecimen withidenticalfeaturesfrom adifferentlocality on Réunion. Some consistentfeaturesof thesecollectionsappeartodistinguish them from thoseof S.flexifolius in the NewWorld,and justifytheirrecognition asrepresenting anewsubspecies– S.flexifolius subsp. reunionensis L.T.Ellis. Generally,plants of the Americantype subspeciesarelargerthanthose in the collectionsfrom Réunion,withshoots reaching around 1-3cmhigh. Unmodified,nongemmiferous leavesaremostly3.0-4.5 mm long and lingulate, withstronglyundulating margins.When dry,theyappearvariouslycurled and twisted,when moist erecttopatent(toreflexed in well-developed material,e.g. Allen 20668 ,MO). The costain mid-hyaline baseisnarrow(usually ca 50µm wide),withaventralsurfaceformed bylong rectangular,thick-walled,porose cells(Fig. 47). Cellsin the chlorophylloselaminaarechollenchymatous,flat and smooth,(10-)12.5 -27.5(-37.5) × (10-)12.5-22.5(-25) µm,mostlywiththick, distinctlyporosewalls(Figs49,50). The marginalstereome often continues distallytothe leafapex.Atthe apicesof some shoots shorter,broader gemmiferous leaveswiththickcostaeformdistinctcomae;gemmaereacharound 650µm long. Inafewspecimens(e.g. Allen 23144,MO)atypical,tall shoots are present,bearing short,incurved leaves.Theseareneatlyarranged on the stem in aslightspiral,but areotherwiseidenticaltoleaveson normalshoots in typical Syrrhopodon flexifolius. Thereareseveralfeaturesof Syrrhopodon flexifolius subsp. reunionensis thatconsistentlycontrast withthoseofthe type subspeciesasoutlined above. Shoots in thissubspeciesarealittle reminiscentof the atypicalshoots in Allen 23144.Theyreach0.5-2.0cmhigh and possess broadlylingulateleaves(Fig. 41), whicharemostly2.5-4 mm long. When dry theyareincurved and appressed, arranged on the stem in aslightspiraltwist; when moist theyareerecttopatent, theirmarginsslightlyundulating. The curled and twisted leavescharacterising dry shoots of typical S.flexifolius areabsent.Thereisatendencyin manyleavesin the Réunion collectionstoslightlybut abruptlybroaden from the distalhyaline baseintothe chlorophylloselimb, aleafshape notgenerallyapparentin New World collectionsof S.flexifolius.Infurther,moresignificantcontrast tothe latter,the costaaround mid-hyaline baseisoften relativelythick(<60->100 µm wide),withthe cellsforming the ventralsurfacelong rectangularand hyaline with very thin,porosewalls(Fig. 46). Cellsin the chlorophylloselaminain leavesof S.flexifolius subsp. reunionensis arechollenchymatous,flatand smooth,mostly (10-)12.5-25(-30) × (10-)12.5-17.5(-20)µm(Figs43-45),relativelythin-walled compared withthoseoftypical S.flexifolius and,unlike in the latter,areonly Calymperaceaeand Orthotrichaceaeinthe Mascarene Islands49 obviouslyporosenearthe hyaline leafbase. Although the range of sizesof the chlorophyllosecellsiscomparable tothatin the type subspecies , most cellsfall within the lowerend of the range and none reachthe extreme largerend of the range. Thisisparticularlyevidentcomparing cellsnearthe leafmargin;thosein NewWorld specimensof S.flexifolius appeargenerallylargerthanthoseinthe Réunion collections(Figs44,49). The marginalstereome in leavesof the latter collectionsfailsshort of the leafapex.Here,the margin isformed bylaminalcells and isusuallyentireormayhaveone ortwoteeth(Fig. 42). Inthe extreme leaf base,the linearmarginalcellsarestronglydifferentiated,mostlytinted orange and withthick,porosewalls; thoseinNewWorld collectionsof S.flexifolius are seldom distinctlycoloured. Discretegemmiferous comaearepresenton shoots in bothofthe available collectionsof subsp. reunionensis .The gemmiferous leaves in Arts RÉU102/192 areaccuratelyillustrated byArts (2005). Gemmaearesimilar tothoseintypical S.flexifolius,mostly400-650µm long. The type collection of Syrrhopodon flexifolius subsp. reunionensis was collected on anearthslope and on fernstolons( Dryopteris )at650malt.,while the otherknowncollection, Porley9728 ,occurred on the stiltroots atthe baseof a Pandanus trunk in miresomewhatbelow900 malt.Thesehabitats arewell within the range of thoseoccupied by S.flexifolius in the NewWorld,whichhas been recorded on treesand herbaceous stemsbetween 450and 900 malt.The evidentlysparsematerialinthe available collectionsof S.flexifolius subsp. reunionensis suggest thatitprobablymostlyoccurs asscattered shoots ratherthan in densetufts ormats.
Selection of specimensexamined Syrrhopodon flexifolius Mitt.subsp. reunionensis L.T.Ellis RÉUNION: La Plaine des Palmistes,PlacedelaUniversité,21°6’56’’S, 55°38’41.3’’E, 12September2008, Porley9728 (BM000890374). Representativeselection from 45 specimensin MO: Syrrhopodon flexifolius Mitt.subsp. flexifolius COSTA RICA: Puntarenas,9.5 km northwest of Rincon de Osa, 08°34’N, 083°31’W, 17July1967, Bowers 327a (MO). BRAZIL.SãoGabriel, Spruces.n. (MO, isolectotype of S.flexifolius). Rio Negro, Spruce‘12’ [= 11?](BM000663854;BM000663855). Roraima, vicinity of armybase, Acampamentodo6*BEC-Jundia, atKm328,16-17November1977, Bucketal. 1848 (MO). Amazonas,along Rio Marié,atManauná, 00°40’S, 66°45’W, 5July1979, Buck2386 (MO). SerraCuricuriari,from Igarapé Arabúof the Rio Curicuriaritothe summit,00°20’S, 60°50’W, 9-12July1979, Buck2442 (MO). Amazonas,along road(U.H.E.deBalbina)to Balbinahydroelectricdamprojectfrom Manaus-CaracaraiRoad(BR 174),02°03’S 59°50’W, 8,11 August 1979, Buck2737J (MO). VENEZUELA: Bolivar,0-6km SE of El Puaji,04°30’N, 61°35’W, 9November1985, Liesner19795 (MO). SURINAME: Sipaliwini, TafelbergNationalPark,northrim of Tafelberg,LisaCreek –savannaarea, 03°54’26’’N, 56°12’45’’W, 26 June 1998, Allen 20525 (MO);LisaCreek –GraceFallstrail,between Lisa and Augustus Creeks,03°54’22’’N, 56°11’26’’W, 30 June 1998, Allen 20668 (MO);upper reachesof GraceCreek,03°53’34’’N, 56°09’35’’W, 5July1998, Allen 20780 (MO);South East LisaCreek,03°54’26’’N, 56°12’45’’W, July1998, Casado 585 (MO). Sipaliwini, Tafelberg,trail from Augustus Creek toNorthRidge,03°55’30’’N, 56°11’55’’W, 20 June 2001, Allen 23144 (MO);trail from Augustus FallstoOld Augustus Creek Camp, 03°55’10’’N, 56°11’15’’W, 21June 2001, Allen 23164 (MO);trail from CaimanCamp toBothropsHill,03°53’30’’N, 56°10’39’’W, 1July2001, Allen 23471 (MO);trail from SouthRim Lookout west along southrim,03°52’53’’N, 56°10’31’’W, 6&7July2001, Allen 23571 (MO). Syrrhopodon parasiticus (Brid.) Paris JAMAICA: Swartz s.n. [ H.2269 ](BM000664063, isotype of S.parasiticus) 50J.Wilbraham&L.Ellis
SYRRHOPODON GARDNERI-LIKE TAXA (Calymperaceae) Syrrhopodon asper Mitt. and S.gardneri (Hook.) Schwägr. Syrrhopodon asper ,asits name suggests,hasleavesthatbristle with acuteprojections.Itoccurs largelyin continentalAfrica and evidenceforits presenceinEast AfricanIslandsissparse. Acollection from Grand Comoro Island,NWof Madagascar( Pócs9268/M ,MO),wascorrectlyannotated as S.asper byW.D.Reese. Reese(1991) alsoreported S.asper from the island of Madagascar(based on Crosby&Crosby5315 and 6676,MO),and from Réunion Island (based on Crosby&Crosby8905 ,MO). The lattercollection represents the distinctivelocalspecies S.mahensis Besch. (discussed and illustrated byEllis &Wilbraham,2008). Although leavesof S.asper haveastructuresimilartothose of S.mahensis theyareeasilydistinguished. Inthe latterspecies,most superficial cellsof the marginalribin the chlorophylloselimbaremultipapillose(Figs65, 66),whilst thosein S.asper aremostlyexserted assmall,smoothacuteteeth (Figs62,63). Ofthe Madagascancollectionsoriginallyidentified as Syrrhopodon asper , Crosby6676 hasnotbeen available forexamination,but Crosby&Crosby 5315 represents ahispid leaved formof Syrrhopodon gardneri (see below). Presently, S.asper isunknowninRéunion Island,and in Madagascar,its presence requiresconfirmation. Syrrhopodon gardneri ,anearlypantropicalspecies,iswidelydistributed across sub-SaharanAfrica (O’Shea, 2006). Leavesin S.gardneri arepapillose,but often withrelativelysmall,lowpapillae(e.g. asin Uganda ,Mugizi1317/25 ,BM) (Fig. 56). However,manycollectionsof S.gardneri from the westernIndian Oceanregion possess unusual,almost hispid leaveswithbristle-like laminal papillae(Figs60,61) thatcould easilybeconfused withthoseof,the similarly bristled, S.asper (Fig. 64). Notable examplesof thishispid formof S.gardneri from Madagascarinclude DeansCowans.n. (BM)and Crosby&Crosby5315 (MO);and from Réunion, Crosby&Crosby8860 (MO)and Lepervanche s.n. (BM, type materialof S.aculeato-serratus Besch. [= S.gardneri ]). The hispid form of S.gardneri maybereadilydistinguished from S.asper bythe structureofthe thickened ribatthe marginsof its leaves.In S.asper ,theseribsaretereteincross- section withmost superficialcellsexserted asacute,distallyleaning projections (Figs62,63). Thosein S.gardneri areangular(mostlysubtriangular)incross- section and mostlyhaveflatsuperficialcells; although intermittent,often geminate,teethareformed atthe anglesof the rib(Figs54-59). Occasionallyin specimensof S.gardneri ,teethemerge from the flat surfaceofthe marginalribadjacenttothoseatthe angles,and consequentlyform adistincttransversegroup(Fig. 55). Thesegeminateormulti-toothed groupsare usuallywell spaced out along the leafmargin,but become morefrequenttowards the leafapex. The superficialappearanceofthe leafmargin maycausesome confusion of S.gardneri withanother,localrough-leaved species, S.pottioides Orbán. This specieswasdescribed from Madagascar(Orbán,1995) and isrecorded from Réunion (Ah-Peng etal.,2005). Itresemblesatiny,depauperateformof S.gardneri, but appears tobedistinguished byits tiny Pottia -like shoots with lingulateleaves(Fig. 67). Insome specimensof otherwisetypical S.gardneri (hispid form) from Réunion (e.g. Coode 4686a ,BR)pairs orgroupsof teethoccur relativelyfrequentlyand closetogetheralong the marginalrib, and unusually, some teethmaybareone ortwosmall papillae(Fig 55). Thisstructureand ornamentation of the marginalribisvery similartothatin the leavesof Calymperaceaeand Orthotrichaceaeinthe Mascarene Islands51
Figs53-67. Syrrhopodon gardneri (Hook.) Schwägr.(53-61), S.asper Mitt.(62-64), S.mahensis Besch. (65,66)and S.pottioides Orbán(67-70). 53,67. Leaves. 54,55,62,65,68. Erectleafmargin in ventralview. 56-59,63,66,69. Marginalribin cross-section. 60,61,64,70. Chlorophyllose laminain cross-section. (53,55,59 Coode 4686a ,BR;56 Mugizi1317/25 ,BM;57,61from Crosby &Crosby5315,MO;58,60 from Lepervanche s.n.,BM, holotype of S.aculeato-serratus;63,64 from Loveridge JPL397 ,BM;62from Pócs9268/M ,MO;65,66 from Balslev637a ,MO;67-70 from Kiss 9443/ED,EGR, holotype of S.pottioides ). 52J.Wilbraham&L.Ellis
S.pottioides (Fig. 68). Featuresdistinguishing S.gardneri and S.pottioides are presented byOrbán(2007),but studyof furthercollectionsmayshowthese speciestobemorecloselyrelated thanispresentlyrecognised. Although presentin the Mascarenesand Madagascar,thereisasyetno evidenceforthe occurrenceof Syrrhopodon gardneri (hispid orotherwise) in the Seychelles.Aspecimen in MO, collected from Praslin Island ( Balslev637)and annotated as S.gardneri byW.D.Reese,is S.mahensis ;its leavesimmediately distinguishable from thoseof S.gardneri bytheirpapillose,ratherthansmooth and flat,marginalcells.
Specimensexamined Syrrhopodon asper Mitt. UGANDA: Kasese,RuwenzoriMts,aboveMinimba Camp, 22 January 1962, Loveridge JPL397 (BM000661412). COMORES: Ngasidja(Grande Comore) Island,11°45’43’’S, 043°18’40’’E, 1-3August 1992, Pócs9268/M (MO). Syrrhopodon gardneri (Hook.) Schwägr.(hispid fo.) MADAGASCAR. DeansCowan5 (BM000677020). Tananarive,along roadN-2,between Atolaonaand Mandraka, 40km Eof Tananarive,18°55’S, 47°55’E, 22 October1972, Crosby&Crosby5315 (MO),originally identified as S.asper ). RÉUNION: Lepervanche s.n. (Hb.Besch.) (BM000677018,type of S.aculeato-serratus). PetitePlaine desPalmistes,10December1969, Onraedt69.R.211 (BR-BRYO 234467-17,BR-BRYO 234467-18) and 69.R.1969 (BR-BRYO 236493-07); Plaine desCafres,auCol Bellevue,7September1971, Onraedt71.R.9156 (BR-BRYO 234489-39;BR-BRYO 234489-40; BR-BRYO 052435-55). Arrt.duVent,along trail toLa Roche Ecrite. From vicinity of forest station (gite) atNW cornerof Plaine desChicots down toparking areaatend of roadCF-1bis,8-12km Sof StDenis,20°56-58’S, 55°27’E, 26 November1972, Crosby&Crosby8860 (MO);Sentierde laRoche Ecriteaudessus de SainteDenis,25November1973, Coode 4686a (BR-BRYO 234469-20);Forêtde Bélouve, forest path:“Letrouduferparle chemin de l’école normale”,Grid:C5.d4,9November1998, Arts RÉU94/40 (BR-BRYO 319337-13);Forêtde Bélouve,sentierde laTamaraies,southwest of the GitedeBélouve,Grid:C5.d3,23 February 2000, Arts RÉU142/39 (BR-BRYO 319339-15), RÉU142/12 (BR-BRYO 319338-14) and RÉU142/77 (BR-BRYO 319340-16). Syrrhopodon gardneri (Hook.) Schwägr.(typicalfo.) UGANDA: Bwindi Impenetrable Forest NationalPark,Buhoma, HabinyanjaHill,14 March2005, Mugizi1317/25 (BM000919752). RÉUNION: Sentiervers laPlaine desChicots,2September1971, Onraedt 71.R.9177 (BR-BRYO 234468-19;BR-BRYO 052346-63). RouteForestièredelaRoche Ecrite,ForetduBrulé,aboveCamp Mamode,Zone Boisde Couleur,4November1998, Arts RÉU73/21 (BR-BRYO 319336-12). Syrrhopodon mahensis Besch .RÉUNION.Arrt.Sous le Vent:Forest Nof StPhilippe, along tractthrough RéserveTouristique. 53km bySfrom SteDenis,21°20’S, 55° 46’E, 28November1972, Crosby&Crosby8905 (MO, originallyidentified as S.asper ). Sentierdu TrembletauNorddeSaintePhilippe,18 December1973, Onraedt73.R.6607 (BR-BRYO 234470-21). SEYCHELLES: Praslin Island. Valle de mai,26 November1979, Balslev637 (MO, originallyidentified as S.gardneri ). Syrrhopodon pottioides Orbán MADAGASCAR: 69kmNW of Antananarivoalong Mahajangaroad,Nof Soaniadananavillage,18°29’54’’S, 47°17’02’’E, 4September1994, Kis 9443/ED (EGR, holotype of Syrrhopodon pottioides ).
MACROMITRIUM (Orthotrichaceae) Macromitriumchloromitrium (Besch.) Wilbraham, comb.etstat.nov. Basionym. Macromitriumfimbriatum (P.Beauv.) Schwägr.var. chloromitrium Besch., Ann. Sci. Nat.Bot. ,Sér.6,9:359. 1880. Type citation. Maurice:montagne duPavillon,BOIVIN (herb.Mus.Par.). Type specimen. Mauritius, Boivin s.n. (BM000873890!–lectotype,selected here ). Calymperaceaeand Orthotrichaceaeinthe Mascarene Islands53
Figs71-82. Macromitriumchloromitrium (Besch.) Wilbraham(71-77)and M.fimbriatum (P.Beauv.) Schwägr.(78-82). 71,78. Habit. 72,79. Branchleaves. 73,80. Branchleafapex. 74,81. Basallaminalcells. 75. Cross-section of upperlamina. 76,82. Upperlaminalcells. 77. Perichaetialleaf. (71-77 from Arts REU32/11,BR;78-82from Arts REU179/18,BR). 54 J.Wilbraham&L.Ellis
Discussion – Macromitriumfimbriatum var. chloromitrium possessesfeatures sufficientlydistinctfrom thoseofthe type variety of M.fimbriatum tobe recognised asadistinctspecies, M.chloromitrium. Boththe latterand M.fimbriatum possess branchleaveswithbluntlyacutetoobtuseapices; bulging, irregularlyrounded and multipapillosecellsin the upperlamina, and differentiated basalcellsrestricted toarelativelysmall proportion of the leafbase. However, M.chloromitrium isdistinguished bythe possession of relativelysmall upper laminalcells(< 9µmdiameter),withcell wallspartiallyobscured bypapillae (Fig. 76);basalcells<30 µm long (Fig. 74),and anisosporous sporophytes.In contrast, M.fimbriatum possesseslargercellsin the upperlamina(> 8µm diameter)withclearlyvisible walls,notobscured bypapillae(Fig. 82);ithas generallymoreelongatecellsin the basallamina(< 41 µm long) (Fig. 81),and isosporous sporophytes. Macromitriumchloromitrium isnewlyrecorded hereforRéunion Island, having been previouslyconsidered asendemictoMauritius.Ithasbeen collected from altitudesbetween 500 and 1700 ma.s.l. and grows asanepiphyteoron rocks. Macromitriumfimbriatum isknownfrom Madagascar,Mauritius,Réunion Island (O’Shea2006),Brazil (Yano,1989) and TristandeCunha ( Palisotde Beauvois, 1805). Ittypicallyoccurs atloweraltitudesthan M.chloromitrium,( 20-890m a.s.l.) and grows asanepiphyte.
Selection of specimensexamined Macromitriumchloromitrium (Besch.) Wilbraham RÉUNION: Cilaos,100 mabovethe village,29September1962, Een R453 (S, B13353). Forest aboveSteDenis,11 October1962, Een R463 (S, B13363). RouteStPierre–StBenoit prèsduMourne de l’Etang,28December1973, Onraedt73.R.6555 (BR-BRYO 236584-01). RoutedelaMontagne,W.deSteDenis,31December1973, Onraedt73.R.6851 (BR-BRYO 236550-64). Footpathfrom BoisCourt toGrand Bassin (Plaine desCafres),20 September 1997, Arts RÉU32/11 (BR-BRYO 318207-47). CirquedeCilaos,LeGrand Matarum surroundingsof La Roche Merveilleuse,29September1997, Arts RÉU57/15 (BR-BRYO 318230-70). CirquedeCilaos,sentierGR-R1from le PlateaudesChênes(LeGrand Matarum) and îlets duBoisRouge,30 September1997, Arts RÉU58/02 (BR-BRYO 318231-71). Dos-d’Âne,footpathtoCapNoir,14 Nov1998, Arts RÉU106/04 (BR-BRYO 318287-30). Le serré,northofSaint-Joseph,25February 2000, Arts RÉU146/48 (BR-BRYO 318312-55). JacquesPayet,SentierduVolcan,27 February 2000, Arts RÉU153/59 (BR-BRYO 318318-61). Macromitriumfimbriatum (P.Beauv.) Schwägr. RÉUNION: Rodriguezs.n. (BM000873893). LePontsuspendude laRivièredel’Est, 22 September1997, Arts RÉU35/06 (BR-BRYO 318214-54). AboveLa Montagne,footpath toIletàGuillaume,from routeD41,atthe junction toSt.Bernard,5November1998, Arts RÉU82/21 (BR-BRYO 318249-89). Valleyof the rivièreLangevin,Grand Galet, 7November1998, Arts RÉU 88/04 (BR-BRYO 318252-92). La Paix,west of BrasPanon, 13November1998, Arts RÉU104/16 (BR-BRYO 318277-20). Commune d’Entre-Deux,La mare,20 February 2000, Arts RÉU127/39 (BRYO-BR 318303-46). Leserré,northofSaint- Joseph,25February 2000, Arts RÉU146/27 (BR-BRYO 318310-53). Forest pathfrom route N2toPointedeLa Table,28February 2000, Arts RÉU156/25 (BR-BRYO 318326-69). Sainte-Anne,Ravine duRivièreSte-Anne nearthe Church,7March2000, Arts RÉU179/18 (BR-BRYO 318345-88). La RivièredesPluies,“La Zone de Plaisirde l’IletQuinquina”, 9March2000, Arts RÉU183/07 (BR-BRYO 318346-89). TRISTAN DE CUNHA: comm. M.Dupetit-Thouars s.n. (G00120670 –type specimen of M.fimbriatum ). Macromitriummauritianum Schwägr. , Sp. Musc.Frond.,Suppl. 2(2):127, pl. 189. 1827. Type citation. IninsulaMauritii lectummisitcl. Sieber. Type specimen. Mauritius. Siebers.n. (G00046151! – lectotype,selected here ;BM000873885!, BM000873884!,BM000873807!-isolectotypes). Calymperaceaeand Orthotrichaceaeinthe Mascarene Islands55
Figs83-93. Macromitriummauritianum Schwägr. 83. Habit. 84-85. Branchleaves. 86-87. Branch leafapices. 88. Cross-section of upperlamina. 89. Basallaminalcells. 90. Upperlaminalcells. 91-93. Perichaetialleaves.(83,91 from Arts REU102/137;84 from BM000873884,type of M.mauritianum ;86,88,89,92from BM000873885,type of M.mauritianum ;85,93from BM000878258,type of M.rhizomatosum ;87from BM000868316,type of M.subpungens ;90from H-BR 2608007,type of M.mauritianum var. viride).
Macromitriummauritianum var. viride Broth., ReiseOstafr.,Syst.Arbeit. 3:55. 1908, s yn. nov. Type citation. Mauritius [Voeltzkow]. Type specimen. Mauritius,1904, Voeltzkows.n.(H-BR 2608007!–lectotype,selected here ;H-BR 2608005!,H-BR 2608006!,H-BR 2608008! -isolectotypes). Macromitriumsubpungens Hampe ex Müll. Hal., Linnaea 40:249. 1876, syn. nov. Type citation. Comoro-insulaJohanna, 1000 met.supramare,in lingo putrido sylvestri:J.M.HildebrandtJunio–Aug. 1875. Coll. No. 1814,cum 56J.Wilbraham&L.Ellis
Macromitrio Hildebrandti associatum. Type specimen. Comores, Hildebrandts.n. (BM000868296!–lectotype,selected here ;BM000868316!,BM000878256!– isolectotypes). Macromitriumsubpungens var. madagassum Cardot, Hist.Phys. Madagascar,Mousses 239. 1915. Type citation. Zone inférieuredesforêts: Maroantsetrˇa ,danslabaie d’Antongil (Ch. Mathieu). Type specimen. Madagascar, Mathieus.n. (PC0101522!–lectotype,selected here ). Macromitriumrhizomatosum Müll. Hal. ex Besch., Ann. Sci. Nat.Bot., Sér.6 ,9:360.1880, syn. nov .Type citation.Nossi-bé:borddesruisseaux,avril 1841,PERVILLÉ, n. 789. Type specimen. Nossi-bé, Perville s.n. (BM000878259! – lectotype,selected here ;BM000878258! –isolectotype).
Discussion – Macromitriummauritianum isrecognised bythe possession of lanceolatebranchleaveswithexcurrentcostae(Fig. 84-87);cellsin the upperlamina obscured bypapillae(Fig. 88),and smoothcellsin the basallaminawithcurved lumina, whichoccupyupto1/3 of the leaf(Fig. 89). The perichaetialleavesare relativelyshort orof asimilarlengthtothe vegetativeleaves,and areoblong with rounded obtusetobluntlyacuteapices(Fig. 91-93). Sporophytesareanisosporous withovoid capsulesand sparselyhairy calyptrae. Anexamination of the type specimensof M.mauritianum , M.mauritianum var. viride Broth., M.subpungens Hampe ex Müll. Hal., M.subpungens var. madagassum Cardotand M.rhizomatosum Müll. Hal. ex Besch. hasshownthem torepresentthe same species,withcharacters closelymatching the abovedescription. Consequently,all of thesetaxaare considered hereassynonymsunder M.mauritianum ,the earliest available name. Brotherus (1908) distinguisheshisvariety Macromitriummauritianum var. viride bythe darkgreen colour of the plant,though he statesthathe found no differenceinthe leavesorperistome tothoseofthe type variety.Specimens of M.mauritianum arevariable in colour,ranging from reddishbrowntoolive green. The name var. viride hasbeen restricted in usetocollectionsfrom Madagascar,and examination of the type materialconfirmsittobeinseparable from var. mauritianum . Inhisprotologuefor Macromitriumsubpungens ,Müller(1876) distinguisheshisnewspeciesfrom related taxabythe presenceofexcurrentcostae in its branchleaves,hencethe epithet“ subpungens ”. However,the lengthto whichthe costaextendsbeyond the cuspidate-acuminateapexof the branch leavesisavariable featurein M.mauritianum ,ranging from the costaending shortlyabovethe acuminateapextobeing stoutlyexcurrent.The type materialof M.subpungens ( Hildebrandts.n.,BM)and additionalspecimensexamined from the Seychelles( Norkett 16409&18239 ,BM)possess excurrentcostaefalling atthe upperend of the range of lengthsobserved in M.mauritianum .The type material of M.subpungens var. madagassum Cardot( Mathieus.n.,PC0101522),alsohas featuresmatching thoseof M.mauritianum . Inhisprotologueof Macromitriumrhizomatosum ,Bescherelle (1880) distinguishesthe newspeciesfrom M.mauritianum byits possession of perichaetialleavesthatarelongerthanthe vegetativeleaves.However,type materialof M.rhizomatosum ( Perville s.n.,BM)hasperichaetialleavesof approximatelyequalsizetothe vegetativeleaves,and anisotype specimen of M.mauritianum ( Siebers.n.,BM000873885) hasperichaetialleavesin different perichaetiaof the same specimen ranging from shortertoapproximatelyequalin sizetothe vegetativeleaves.Therefore,the sizeofperichaetialleavesdoesnot appeartobeasatisfactory charactertodelimitthesetaxa, whicharehere considered asconspecific. Calymperaceaeand Orthotrichaceaeinthe Mascarene Islands57
Macromitriummauritianum isrecorded from the AgalegaIslands, Madagascar,Tanzania, Mauritius and Réunion Island (O’Shea, 2006). The synonymywiththe name Macromitriumsubpungens addsthe Comorosand Seychellestoit’sknowndistribution. Itiscommonlyfound atlowtomid-level altitudesand hasbeen collected from sealevel upto1200 ma.s.l.,growing asan epiphyteoron rocks.
Selection of specimensexamined Macromitriummauritanum Schwägr. COMORES: Grande Comore,SWslopesof Karthala, 26 July1992, Magill &Pócs10918, (BM000862001). MAURITIUS: 1881, Robillard1772 (BM000960608). SouthofTamarin Reservoiron banksof RivieredesAigrettes,31November1968, Barclay1379a (Priv.Herb. Townsend). RÉUNION ISLAND: StPhilippe,RéserveNaturelle de MareLongueforest trail,7March1997, Arts RÉU11/01 (BR-BRYO 318195-35) and RÉU11/02 (BR-BRYO 318196-36). La Coteaux vents,Ste-Rose,22 September1997, Arts RÉU36/04 (BR-BRYO 318215-55). Tremblet,8November1998, Arts RÉU92/29 (BR-BRYO 318258-01). Circuit desRavenales, ca 1kmNW of Chemin de Ceinture,10November1998, Arts RÉU97/82 (BR-BRYO 318269-12). ForêtDomaniale de BrasPanon,atthe “Sitetouritistiqued’Eden”, aformerthee plantation,12November1998, Arts RÉU102/137 (BR-BRYO 318272-15). ForêtMourouvin,N-NE slope of Piton de lafournaise,along the roadtoles3citernes, 13November1998, Arts RÉU105/15 (BR-BRYO 318280-23). La Grande Montagne,Route ForestièredeLa Plaine d’Affouche,20 November1998, Arts RÉU122/07 (BR-BRYO 318299-42). BoisBlanc, sitedelaSource,26 February 2000, Arts RÉU152/07 (BR-BRYO 318315-58). SEYCHELLES: Mahé,Salazie,ForêtNoirRoad,4September1973, Norkett 16409 (BM000873865). Mahé,Vingt-Cinq-Sous,22 December1973, Norkett 18239 (BM000873862). Mahé,Morne Blanc, 23 January 1974, Norkett 18740 (BM000960613). Macromitriumpallidum (P.Beauv.) Wijk et Margad., Taxon 9:190.1960. Basionym . Orthotrichumpallidum P.Beauv., Prodr.Aethéogam. 81. 1805. Type citation. Ile de Bourbon;communiqueparM.Bory-St-Vincent.Communiquepar M.Dupetit-Thouars quilarecueillie aux Ilesde Franceetde Bourbon. Type specimen. Réunion, Bory-St-Vincents.n.(G00120669! – lectotype,selected here ). Macromitriumvoeltzkowii Broth .,ReiseOstafr.,Syst.Arbeit. 3:55. 1908, syn. nov. Type citation. Mauritius [Voeltzkow]. Type specimen. Mauritius, Voeltzkows.n.(H-BR 2616005! – holotype). Discussion – Macromitriumpallidum closelyresembles M.mauritianum but is distinguished from the latterbythe possession of lanceolatebranchleaveswith sparselytuberculatebasalcells(Fig. 99);anupperlaminathatisdenselypapillose withoccasionalbistratosepatches(Fig. 100);acostaending in orbelowthe acute leafapex(Fig. 97),and short,ovate-lanceolateperichaetialleaveswithbluntly acuteapices(Fig. 101). The type materialof M.voeltzkowii (V oeltzkowis.n., H-BR),described from Mauritius,hasfeaturesmatching thoseof M.pallidum (P.Beauv.) Wijk et Margad. (Réunion, Bory-St-Vincents.n.,G–lectotype). Previous recordsof M.voeltzkowii Broth. occurring on Réunion Island (Een 1978) arereferable to M.chloromitrium (Besch.) Wilbraham(see discussion hereunder thisspecies). Macromitriumpallidum hasbeen recorded from Brazil (Yano,1981), Réunion,Mauritius and the Seychelles(O’Shea, 2006). Ithasbeen collected between altitudesof 220 and 980ma.s.l.,growing asanepiphyte.
Selection of specimensexamined Macromitriumpallidum (P.Beauv.) Wijk et Margad. MAURITIUS: Mauritius, DupetitThouars s.n. (BM000873859! –syntype of Macromitrium pallidum ). RÉUNION: Rodriguezs.n.(BM000873894). Plaine desPalmistes,de laroute 58 J.Wilbraham&L.Ellis
Figs94-101. Macromitriumpallidum (P.Beauv.) Wijk. et Margad. 94. Habit. 95-96. Branch leaves. 97-98. Branchleafapices. 99. Basallaminalcells. 100. Cross-section of upperlamina. 101. Perichaetialleaves.(94 from Arts REU102/143 ,BR;96-98,100-101from BM000873859,type of M.pallidum ;95,99 from H-BR 2616005,type of M.voeltkowii).
StPierre–StBenoit,27 December1969, Onraedt69.R.742 (BR-BRYO 235773-63). Sud- ouest de Saint-Benoit,entreleGrand Etang etlaroute,28December1973, DeSloover 17.858 ( BR-BRYO 273489-46 ). Tremblet,8November1998, Arts RÉU92/27 (BR-BRYO 318257-00). ForêtDomaniale de BrasPanon:atthe “Sitetouritistiqued’Eden”,aformer thee plantation,12November1998, Arts RÉU102/143 (BR-BRYO 318273-16). Forêt Mourouvin,N-NE slope of Piton de lafournaise,aboveles3citernes,13November1998, Arts RÉU105/47 (BR-BRYO 318282-25) and Arts RÉU105/50 (BR-BRYO 318283-26). JacquesPayet,SentierduVolcan,27 February 2000, Arts RÉU153/21 (BR-BRYO 318316- 59), RÉU153/28 (BR-BRYO 318317-60)AND RÉU153/85 (BR-BRYO 318321-64). Macromitriummicrostomum (Hook. et Grev.) Schwägr., Sp. Musc.Frond.,Suppl. 2(2):130.1827. Basionym : Orthotrichummicrostomum Hook. et Grev., Edinburgh J.Sci. 1:114, pl. 4 .1824. Type citation.VanDieman’sLand,Dr.Spence. Type specimen.Tasmania, Spence&Neill s.n. (E00011665! – lectotype, Vide Vitt & Ramsay,1985;isotype –BM000873897!). Calymperaceaeand Orthotrichaceaeinthe Mascarene Islands59
Figs102-110. Macromitriummicrostomum (Hook. et Grev.) Schwägr. 102-104. Branchleaves. 105-107. Branchleafapices. 108-109. Upperlaminalcells. 110. Basallaminalcells.(103,105,107, 110from NY00970201,type of M.fasciculare ;102,106,108,from PC0105915,type of M.fasciculare var. angustifolium ;104from BM000868312,type of Dasymitriumborbonicum ;109 from FH, type of M.fasciculare var. javense ).
Macromitriumfasciculare Mitt.var. fasciculare , J.Proc.Linn. Soc.,Bot., Suppl. 1:51. 1859, syn. nov. Type citation.Hab.InCeylon adHorton Plainset NeweraEllia, Gardner! (No. 225,229,230). Type specimen.SriLanka, Horton Plains,Feb1846, Gardner230 (NY 00970201! – lectotype,selected here ;isotype –BM000919522!). Macromitriumfasciculare var. javense M.Fleisch., MusciBuitenzorg 2: 432.1904, syn. nov. Type citation.Exsiccata: M.FLEISCHER, Musc.Archip. Ind.,No. 281 (1902). Type specimen.Java, Fleischerexicc.281 (FH 00258463!– lectotype,selected here ). Macromitriumfasciculare var. angustifolium P.delaVarde, Rev.Bryol. Lichénol. 19:152.1950. Type citation.Sommetorientaldumassif duMarojéjyà l’ouest de lahauteManantenina.Gneiss etquartzite. Alt.1.850–2.100 m. Type specimen.Madagascar, Humbert s.n. (PC0105915! – lectotype,selected here ). Dasymitriumborbonicum Besch., Ann. Sci. Nat.Bot.,Sér. 6,9:355. 1880, syn. nov. Type citation.La Réunion:plaine desChicots,BORY (herb.COSSON); BOIVIN, 1849 (in herb.Mus.Par.);plaine desPalmistes,G.DE L’ISLE, 1875, 60 J.Wilbraham&L.Ellis nos.200 et286; pasde Belcombe,P.LÉPERVANCHE, 1877 (hb.Mus.Par.,sub Macromitrio schizomitrio Nob.priùs). Madagascar(N.O.):PERVILLÉ, 1849, no. 827. Type specimen.Réunion Island, G.d’Isle 286 (BM000868311! – lectotype, selected here ).
Discussion –Macromitriummicrostomum haslanceolatebranchleaveswithacute toacuminateapices(Fig. 102,103). Cellsin the upperlaminaareirregularly rounded toshortlyoblong and smooth(Fig. 109),whilst thoseinthe basallamina arelinearand smooth,occupying the lower1/3-1/2 of the leaf(Fig. 110). Sporophytesin thisspeciesareisosporous withelongate,smoothsetaeand capsulesthatarepuckered atthe mouth,the calyptraisessentiallyhairless.This pantropicalspecieshasbeen recognised underalarge numberof localnamesand additionalresearchinto Macromitrium islikelytoincreasethislist of synonyms still further. Inthe protologuefor Macromitriumfasciculare ,Mitten (1859) compares hisplantwith M.reinwardtii Schwägr.,aname whichislaterplaced in synonymy with M.microstomum byVitt &Ramsay(1985). Mitten considered his M.fasciculare tobesimilartothe speciesnowknownas M.microstomum ,but distinguished bythe “ structurafoliorum,cellulisminoribus numerosioribus infra mediumfolii descendentibus etcalyptraparcepilosadisprepans ”. Examination of the lectotype materialof M.fasciculare ( Gardner230,NY)found the leafshape and laminalcellstobeagood matchforthoseofthe lectotype of M.microstomum ( Spence&Neill s.n.,E). The upperlaminalcellsof thisspeciesaresomewhat irregularin shape,range in diameterfrom 3.5–8 µ m,and vary slightlyin the thickness of theircell walls.Calyptraeintype materialof M.fasciculare ( Gardner 229 ,NY)areessentiallyglabrous,though anoccasionalhairisvisible. Isotype materialof M.fasiculare ( Gardner230,BM)hasalsobeen determined by Shuiliang Guo(ShanghaiNormalUniversity)as M.microstomum (pers.comm.). Macromitriumfasciculare hastworecognised varieties:var. angustifolium P.delaVarde described from Madagascarand var. javense M.Fleisch. described from Java.The type materialofvar. angustifolium ( Humbert s.n.,PC0105915) represents anunusualformof M.microstomum .Thisspecimen possessesbranch leaveswhicharenarrowerthanistypicallyfound in M.microstomum and has upperlaminalcellswhicharemoredistinctlyelongate(Fig. 108). However,these distinguishing featuresof var. angustifolium aresomewhatplasticwithin M.microstomum and though extreme in thisplantdo notjustifyrecognition atan infraspecificlevel. The type materialof M.fasciculare var. javense ( Fleischer exsicc.281,FH ) wasfound tobeinseparable from collectionsof the type variety. Fleischer(1904) hadconsidered var. javense distinctin possessing smallerupper laminalcellsthanthe type variety,but theseactuallyfall well within the range of sizesobserved in M.microstomum . Dasymitriumborbonicum wastreated asasynonymof Macromitrium serpens bySim (1926),though he did notsupport hissynonymywithcited specimens.Syntype specimensof D.borbonicum in Bescherelle’sherbarium,BM ( Lépervanche s.n.; Boivin s.n.; G.d’Isle 286 ; Pervillé s.n.),areall referable to Macromitriummicrostomum ,aspeciesclearlydistinctfrom M.serpens .Crosby et al. (1983)alsolist D.borbonicum asasynonymof M.serpens ,but again, evidentialspecimensarenotcited. Part of the type materialof Dasymitrium borbonicum (Réunion, Lépervanche s.n.,BM)isalsothe basisforBescherelle’s herbariumname Macromitriumschizomitrium .Thismaterialwasexamined by Fleischer(1904),who placed M.schizomitrium , nom. nud. in synonymywith M.fasciculare [= M.microstomum ].The available evidencesuggests that Calymperaceaeand Orthotrichaceaeinthe Mascarene Islands61
Figs111-116. Macromitriumserpens (Hook. et Grev.) Brid. 111-112. Branchleaves. 113-114. Branchleafapices. 115. Upperlaminalcells. 116. Cross-section of upperlamina.(111, 114,115 from BM000873888,isotype of M.serpens ;112,113,116from NY00518241,type of M.astroideum ).
D.borbonicum should beplaced in synonymywith M.microstomum .Owing to the past confusion of thesetaxa, distribution recordsof Dasymitriumborbonicum from Réunion Island and Madagascar,havebeen mistakenlyallocated to M.serpens .However,asboth M.microstomum and M.serpens occur on these islands,thisdoesnotaffecttheirrecorded distributions. Macromitriummicrostomum waspreviouslyknowntooccur in Bolivia (Churchill,2005),Brazil,DominicanRepublic, Cuba, SouthPacificIslands, Hawaii,StHelena, SouthAfrica, Malawi,SriLanka, Java, NewGuinea, Eastern Australia, Tasmaniaand NewZealand (Wilbraham,2007). Itwasknownunder the name M.fasciculare from Tanzania, Madagascar,Réunion Island (O’Shea, 2006),Philippines(Tan,1991),Java, Sulawesi,NewGuinea(Eddy,1996)and China, HainanProvince(Redfearn etal. 1996). Macromitriummicrostomum is commonlyencountered in montane forests on Réunion,and hasbeen collected from 1000–2300 ma.s.l.,growing asanepiphyteorrarelyon rocks.
Selection of specimensexamined Macromitriummicrostomum (Hook. et Grev.) Schwägr. TANZANIA.IushotoDistrict,East UsambaraMts,25March1975, Townsend 75/476 (Priv. Herb.Townsend). MADAGASCAR: Pervillé s.n.(BM000868313,syntype of Dasymitrium borbonicum ). RÉUNION ISLAND. Lépervanche s.n. (BM000868312,syntype of D.borbonicum )and Boivin s.n. (BM000868314,syntype of D.borbonicum ). Forêtde Plaine desPalmistes,PremierVillage,sentiervers laCascade Biberon,13March1997, Arts 62 J.Wilbraham&L.Ellis
RÉU25/21 (BR-BRYO 318205-45). Plaine desCafres,sentierGR-R2vers le Piton des Neiges,entrePiton MareaboueetPiton Tortue,21September1997, Arts RÉU33/106 (BR-BRYO 318212-52). Forêtde NotreDame de La Paix,Sentierbotanique,25September 1997, Arts RÉU48/55 (BR-BRYO 318223-63). CirquedeMafate,footpathGR R1from the Pasde Col duBoeuf(Grand Îlet)toLa Nouvelle,3October1997, Arts RÉU61/41 (BR-BRYO 318237-77). Forêtde Belouve,forest path,9November1998, Arts RÉU94/57 (BR-BRYO 318263-06). Plaine desCafres,SourceReilhac, west side of piton de laSource, 10November1998, Arts RÉU96/48 (BR-BRYO 318267-10). Forêtde Tevelave,Sentierdes Tamarins,18 November1998, Arts RÉU116/05 (MEISE, BR-BRYO-318293-36). Cirquede Cilaos,BrasSec, SentierdesCalumets northof’le Bonnetde Pretre’,4March2000, Arts RÉU171/62 (BR-BRYO 318337-80). SRI LANKA: NuwaraEliya, July1845, Gardner 225 (37)(NY 00970199,syntype of Macromitriumfasciculare ). Horton Plains,February 1846, Gardner229 (NY 00970200,syntype of M.fasciculare ). Macromitriumserpens (Hook. et Grev.) Brid., Bryol. Univ. 1:736.1826. Basionym . Orthotrichumserpens Hook. et Grev., Edinburgh J.Sci. 1:119, pl. 5. 1824. Type citation. Trunksand branchesof treesin woodsatSylvasStation,near George’sPlain,AnteniquaLand,Cape of Good Hope. W.J.Burchell,Esq. Type specimen. SouthAfrica, Burchell s.n.(E– lectotype, vide vanRooy&vanWyk, 1992,notseen;isolectotype –BM000870451!). Macromitriumastroideum Mitt., Philos.Trans. 168(extravol.):390.1879, syn. nov. Type citation. Rare. Onlyatthe top of OysterRivervalley.Trailing over boulders. Type specimen. Rodriguez, Balfour s.n.(NY00518241! – lectotype, selected here ;isotype –BM000873888!) Discussion –Macromitriumastroideum hasbeen considered endemictothe island of Rodriguez(Mitten,1879) though itisfound heretobeasynonymofthe more widespread M.serpens whichischaracterised bythe possession of along,creeping primary stem withshort secondary branches; lanceolatebranchleaveswithfragile apices(Figs111,112)and basallaminalcellsthatareoften tuberculate. The upper laminaisformed bymultipapillosecellsbut hasanunusualstructureinpossessing irregularlydistributed unicellularprotruberancesfrom bothdorsaland ventral surfaces(Fig. 116). The type materialof M.astroideum (Rodriguez, Balfour s.n., BM, NY)possessesall of thesecriticalfeatures,and isindistinguishable from collectionsof M.serpens . Macromitriumserpens hasbeen recorded from Madagascar,Malawi,Mozambique,Réunion Island,SouthAfrica and Tanzania (O’Shea, 2006)and waspreviouslyunrecognised from Rodriguez.Itisarareplant on Réunion.
Selection of specimensexamined Macromitriumserpens (Hook. et Grev.) Brid. SOUTH AFRICA: Cape Province,StormsRiverForest,1August 1952, Taylor468 (BM 000873891). Cape,Knysna, Eddy&Sims7098 (BM000878010). RÉUNION ISLAND: AboveLa Montagne,atthe junction toSt.Bernard,5November1998, Arts RÉU82/19 (BR-BRYO 318248-88). Macromitriumlanceolatum Broth. Macromitriumlanceolatum wasnewlyrecorded forRéunion Island by Een (1978). However,thisrecordhasbeen overlooked in recentchecklists forthis island (Ah-Peng &Bardat,2005;Ellis&Wilbraham,2007). Thisspeciesisknown from Mauritius and from asingle collection from Réunion ( Rodriguezs.n., S-B63459). Macromitriumlanceolatum ismorphologicallyvery closetothe more widespread M.orthostichum Nees ex Schwägr. Macromitriumorthostichum possessesovateleaveswithstronglybulging upperlaminalcellswhile Calymperaceaeand Orthotrichaceaeinthe Mascarene Islands63
M.lanceolatum differs bythe lanceolateacutebranchleaveswithflatterupper laminalcells.Additionalspecimensof M.lanceolatum need tobeexamined before the taxonomicstatus of thisspeciescanbedetermined. Bothtaxabelong tothe distinct Macromitrium subg. Cometium ,whichisdiscussed byWilbraham(2007).
Specimensexamined Macromitriumlanceolatum Broth. RÉUNION: Rodriguess.n. (S–B63459). Macromitriumorthostichum Nees ex Schwägr. RÉUNION: Plaine desPalmistes,sous le village La Plaine desPalmistes,28December 1973, DeSloover17.826 (BR-BRYO 273490-47). ForêtDomaniale de BrasPanon,atthe “Sitetouristiqued’Eden”,aformerthee plantation,12November1998, Arts RÉU102/162 (BR-BRYO 318275-18). Macromitriumbelangeri Müll. Hal. Macromitriumbelangeri isconsidered endemictoRéunion Island. The type material(Réunion Island, Belanger4 ,in herb.C.Müller)wasprobably destroyed withMüller’sherbariuminBerlin,and extantmaterialassociated with Müller’sprotologueof M.belangeri (Müller,1862)hasnotbeen located. However,hisoriginaldescription of the speciessuggests acloseaffinity with M.sulcatum .Inparticular,Müller(1862)refers tothe denticulateleafapexand tuberculatebasalcellstypicalof M.sulcatum .Itisvery likelythesetwonamesare synonymous. Acknowledgements. Wearegratefultothe curators atBR, G, H, MO, NY, PC and Sforthe loanofspecimensand toR.D.Porleyfordonating part of hiscollection of Syrrhopodon flexifolius subsp. reunionenis forexamination. Wealsothank the Université de laRéunion and La Réunion ParcNationalforhosting the Bryophytesof Mascarene Islands–First TaxonomicWorkshop and Field Meeting on Réunion Island. Some initial datafrom the fieldworkareincluded in thispaper.
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APPENDIX
List of taxain Calymperes , Syrrhopodon and Macromitrium knownfrom Réunion Island. Modified from Ah-Peng &Bardat(2005) and Ellis&Wilbraham(2008).
Bold print =currentlyaccepted name E= Taxaconsidered endemictoRéunion Island
CALYMPERES (Calymperaceae) Calymperesafzelii Sw. C.erosum Müll. Hal. C.graeffeanum Müll. Hal. C.hispidum Renauld &Cardot C.palisotii Schwägr. C.pallidum Mitt.(newtoRéunion) C.taitense (Sull.) Mitt.subsp. pachyloma (Hampe) L.T.Ellis C.tenerum Müll. Hal.
SYRRHOPODON (Calymperaceae) Syrrhopodon africanus (Mitt.) Parissubsp. africanus S.apertifolius Besch. S.armatus Mitt.subsp .insularum (Bizot&Onr.) Orbán&W.D.Reese S.dimorphophyllus L.T.Ellis(newtoRéunion) S.flexifolius Mitt.subsp. reunionensis L.T.Ellis (E) S.hispidocostatus Renauld &Cardotsubsp. hispidocostatus S.hispidocostatus Renauld &Cardotsubsp. artsii L.T.Ellis (E) S.gardneri (Hook.) Schwägr. S.gaudichaudii Mont. S.involutus Schwägr. S.mahensis Besch. S.mauritianus Müll. Hal. ex Ångstr. S.parasiticus (Brid.) Paris S.pottioides Orbán S.vardei L.T.Ellis(newtoRéunion) 66 J.Wilbraham&L.Ellis
MACROMITRIUM (Orthotrichaceae) Dasymitriumborbonicum Besch. =Macromitriummicrostomum Macromitriumastroideum Mitt.=M.serpens M.chloromitrium (Besch.) Wilbraham M.fasciculare Mitt.var. fasciculare=M.microstomum M.fasciculare Mitt.var. angustifolium P.delaVarde =M.microstomum M.fasciculare Mitt.var. javense M.Fleisch. =M.microstomum M.fimbriatum (P.Beauv.) Schwägr . M.fimbriatum (P.Beauv.) Schwägr.var. chloromitrium Besch. ≡ M.chloromitrium M.lanceolatum Broth. M.mauritianum Schwägr. M.mauritianum Schwägr.var. viride Broth. = M.mauritianum M.microstomum (Hook. &Grev.) Schwägr . (newtoReunion) M.orthostichum Nees ex Schwägr. M.pallidum (P.Beauv.) Wijk &Margad . M.rhizomatosum Müll. Hal. exBesch. = M.mauritianum M.serpens (Hook. &Grev.) Brid. M.subpungens Hampe ex Müll.Hal. = M.mauritianum M.subpungens Hampe ex Müll. Hal. var. madagassum Cardot.=M.mauritianum M.sulcatum (Hook.) Brid. M.voeltzkowii Broth. = M.pallidum
Taxanewlyexcluded from the moss floraof Réunion Island: Calymperestaitense (Sull.) Mitt.subsp. taitense Syrrhopodon albidus Thwaites&Mitt.subsp. integrifolius (E.B.Bartram) L.T. Ellis(= S.prolifer var. seychellarum Orbán) S.asper Mitt. S.crenulatus Mitt. S.prolifer Schwägr. Macromitriumbelangeri Müll. Hal.