Further Taxonomic Studies on the Families Calymperaceae (Musci
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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