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Notes on Lepturus (Gramineae) in Malesia PDF

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BLUMEA 47 (2002) 385-389 Notes on Lepturus(Gramineae) in Malesia R. Nowack & J.F. Veldkamp Nationaal Herbarium Nederland,UniversiteitLeiden branch, P.O. Box9514,2300RA Leiden,The Netherlands Summary Asurvey ofLepturusR.Br.(Gramineae)in Malesia is given.Monerma P.Beauv. is superfluous, iand hsenceMonermeaedanillegitiematetribsalnamec.OnenerwvairietybofL. repeens (G.Fdorst.) R..Br. Key words'.Gramineae,Lepturus,Monerma,Monermeae,Malesia. Introduction Lepturus R.Br.(Gramineae) is asmall genusofperhaps 8 species oftheshoresofthe IndianandPacificOceans. Foralong timeitwas associatedwithanumberofgrasses withsomewhatsimilarspikes, e.g. Aegilops L., HordeumL., Nardus L., Parapholis C.E.Hubb.,PhalarisL., RottboelliaL.f„ andwhatnowarecalledHainardiaGreuter andOropetiumTrin. (incl.Lepturella Stapf). Fromgeneralmorphological,anatomical(Hansen &Potztal, 1954),andnoweven moleculardata(Hilu& Alice,2000) ithasbecomeevidentthatitis theonly member (ifIschnurusBalf.f.isregarded as asynonym; cf.Clayton & Renvoize, 1986)ofthe Leptureae Holmb., close to the Chloridoideae-Cynodonteae Dumort. Hilu & Alice placed Lepturus as sister grouptotheChloridinaeJ.Presl. Ina combinedanalysis basedon morphology andleafanatomyVan denBorre & Watson (2000) includedLepturus in their 'Chloris group' next to Ischnurus. This affinity is supported by Lepturopetium Morat, which allegedly is a hybrid between Lepturus repens(G.Forst.)R.Br, andChlorisbarbataSw. However,thistheoryis not evidentfromthepositions ofthesegenerainVanden Borre& Watson'scladogram. It isalso underminedby thefact that both species have a very widedistribution, while Lepturopetium is only known fromNew Caledonia, theMarianas(Rota), Marshalls (Eniwetok), andCocos Keeling (West Islandor P.Panjang). TRIBALNAME AND NOMENCLATURALPOSITION OF MONERMA Beauvois(1812:110) notedunderLepturus R.Br,thatthisnamehadbeenusedforin- sects(LepturaL., 1758,Coleoptera, Longhorn beetles) andforthatreason hadtobe rejected. The only taxon he includedwas Rottboëllaespec. [G.?] Forst., and in his Index(p. 177)we findR. repens[G.?] Forst. as theonly Forster nameandtherere- ferredtoMonerma.Underthat name(Index, p. 168)M.monandraandM. repensare enumerated.Under thetreatment ofMonerma(p. 116) Rottb. repensis citedagain. 386 BLUMEA Vol. 47, No. 2, 2002 Obviously thiswasanintentionalrenamingofLepturus toMonerma.As namesunder theZoological andBotanicalCodes (Art. 54.1) arecompletely independent Leptura L. hasnopriority over Lepturus R.Br, (and may thisrulebe maintained!). Therefore,Monermais superfluous forand homotypic withLepturus. Hencethe tribalname MonermeaeC.E.Hubb. being derivedfrom anillegitimate nameis also illegitimate (Art. 19.5). ThegenusMonermainthesenseofEuropean florasistobecalledHainardiaGreuter andthetribetowhich itbelongs HainardieaeGreuter. Dumortier(1824) alsoknew about thebotanical/zoological homonymy andpro- posed the tribalname Lepiureae Dumort.taken fromLepiurus Dumort., aminorre- naming ofLepturus R.Br.Likewisethesenames are illegitimate.Forthesamereason Rafinesque (1819)proposed thenameLeptocercus (fideHitchcock, 1920).Foramore detaileddiscussionproandcontra this, seee.g.Scholz (1995)andGhandi(1996). LEPTURUS Themost widespread representative is thebeach grassLepturus repenswhichranges from theAfrican coasts ofthe IndianOcean to the Pacific. It growspreferably on coralsandsandlimestonerocks.Although itis distributedby theseaandonetherefore wouldexpect a continuousexchange ofgenetic informationwhereby its variability wouldberatherhomogeneous, itis quitevariable,andF. Brown (1931)andFosberg (1955)havedescribedafairnumberofinfraspecific taxa,someofwhichwouldalso occur in Malesia, e.g. var. subulatusFosberg (= var. y F.Br.) and var. occidentalis Fosberg. Fosberg basedhis taxa onpersonal fieldobservationsandfour quantitative characters,e.g.thicknessofthespike, length ofitsjoints, length ofjointplus glume, andshape ofthelatter'sapex. He concludedthat several regional varietiescouldbe segregated, though their geographic ranges are not very clear and their distinctive charactersoverlap invariousdirections. Anexaminationof the variability ofthespecies along Fosberg's lines was made forMalesianmaterial,amongwhichwere some paratypesof var. subulatusand var. occidentalis.Itturnedout tobeimpossible torecognise anydistinctentities, letalone taxa,as allanalyses showedacontinuouspatternofvariabilityandmix-upofFosberg's andothercharacters. However, two collectionsfrom two smallislands ontheWcoast ofNewGuinea (Pam Is., KofiauIs.(Halmaheira Sea)) wereimmediately conspicuous becauseofthe exceptional pubescence ofthestem,blades, andglumes in thisotherwise essentially glabrous species, andare hereregarded as representing anundescribedvariety. Lepturus R.Br. LepturusR.Br. (1810)207;I.Hansen & Potztal (1954)256; Fosberg (1955) 285. —Monerma P.Beauv. (1812) 116,168, 177,nom.superfl. —LepiurusDumort. (1824) 140,nom. superfl. —Leptocercus Raf. (1819) 190,nom.superfl. —L(epturus sect.EulepturusHack. (1889)78 nom. inval.)—Typespecies:Lepturusrepens (G.Forst.) R.Br. Stoloniferous annualsor perennials. Ligule a glabrous or ciliatemembrane.Blades flatorenfolded.Inflorescence asingle, terminalspike, disarticulating intocylindrical R.Nowack & J.F.Veldkamp: Notes onLepturusin Malesia 387 joints.Spikelets solitary, alternately inserted incavitiesalong thethickenedrhachis, dorsally compressed, 1-or2-flowered, falling atmaturity withtheadjacent internode. Terminalspikelet falling withthe uppermostlateralspikelet. Lowerglumes reduced to arudimentexcept intheterminalspikelet. Upper glumes abaxial, robust, several- nerved,closely covering the spikeletexcept inanthesis.Rhachillaprocess produced. Callus ofthe lemmashort,obtuse. Lemmas lanceolate,3-nerved, lateralsextended closeto theapex. Stamens 3.x =9. Distribution—8-12species on theshoresoftheIndianandPacific Oceans. KEY TO THETAXA la. Spikelets 8-22mmlong 2 b. Spikelets 3-5 mmlong 1.L. radicans 2a. Culms, sheaths,blades,peduncle, andupperglumes densely pubescent. Rhachis internodessparsely hairy 2b.L. repens var. pubescens b. Culms, sheaths,blades,peduncle, rhachis internodesandupperglumes glabrous 2a.L. repensvar. repens 1.Lepturus radicans(Steud.) A.Camus Lepturusradicans (Steud.)A.Camus(1923)87;Veldk. (1999)233. —Ophiurosradicans Steud. (July 1854)430. —Monerma radicans (Steud.)Hack. (1889)320. —Lectotype: Boivin 1980 (holoP;K), designatedbyC.E. Hubbard (1974). Mat-formingperennial. Culmstough,0.1-0.3m long,creeping to erect.Sheathsgla- brous.Ligule 0.1-0.5 mmlong. Leaves 3-12.5 cm by 3-8mm. Spikes 3-7cm by c. 2 mm. Spikelets lanceolate, 3-5 mm long. Lowerglumes absent; upper glumes 5-9-nerved,smooth,acuminateto caudate.Lemmas 2.5-4mmlong,(sub)glabrous. Anthers0.6-2mmlong. Caryopsis ellipsoid, 1-1.25mmlong. 2n = 18. Distribution—EAfrica, Madagascar, Comores,Seychelles, Mauritius,introduced inSriLanka;Malesia:PeninsularMalaysia (Selangor: KualaLumpur, Ryves s.n. Nov. 1992(K); JFVwent inJuly 1998totheroadsideopposite theConcordeHotel,where itwas collected,but couldnot findit). Habitat—Sands, grassland nearthe sea, roadsides,lawns,preferring someshade, upto500m altitudeinSriLanka. Uses — Saidtomake a good lawn grassin shadedareas. 2.Lepturus repens (G. Forst.) R.Br. Lepturusrepens(G.Forst.)R.Br.(1810)207.—Rottboellia repensG.Forst.(1786)9.—Monerma repens(G.Forst.)P.Beauv. (1812)117, 168,177.—Lectotype: Forsterdrawing20,engraving 14(holoBM),"Teoutea" (=Takaroa),designatedby F.R.Fosberg, in sched. (seenote 1). Lepturusaciculatus Steud.(inZoll. (June1854)57,nomen);(July 1854)357. —Type:Zollinger 1083 (holoP;perhapsinA,B,BM,BO, G,L,LE, W). [Lepturusrepens R.Br.var.y F.Br.(1931)90.—Voucher ('type'): E.H.Quayle 442 (BISH)]. — Lepturusrepens R.Br.var.subulatus Fosberg (1955)290. —Type:Fosberg 34173 (holoUS). Lepturusrepens R.Br. var. occidentalis Fosberg (1955)291. —Type: Backer 31067 (holoUS; BO,L,U). 388 BLUMEA Vol. 47, No. 2, 2002 a. var. repens Mat-forming glabrous perennial. Culms tough, 0.2-0.4 m long, creeping to erect. Ligule 0.3-0.5 mmlong. Leaves5-20 cm by 3-8 mm. Spikes 5-12cm by 1.2-2 mm. Spikelets 8-22 mm long. Rudimentoflower glumes triangular, 0.3-0.5mm long; upperglumes 7-11-nerved, scabrous, acuminateto long-caudate. Lemmas4- 5.1 mmlong, (sub)glabrous, sometimesawn-tipped.Anthers1.5-2mmlong. Caryopsis c. 2mmlong.2n - 42,54. Distribution—EAfrica, MascareneIs., Seychelles, SriLankato Japan, Australia (NTerritory,Queensland), Polynesia, Hawai'i;Malesia:widespreadinsuitableplaces. Habitat— Mat-forming onsandy androcky seashores.Preferably on coralsands. Alsofoundalong roadsandinsavannas (?). Usually ataboutsea-level,inFloresonce at 1000m altitude(Jaag 11810,L). Uses — Efficient sand-binder.Not usedfor fodderbecause ofitsharshness and inferior yield. Vernacularnames — Bunchgrass,Pacific Islandthintail. Notes — 1.The lectotypification isafteramanuscript by thelateDr.F.R. Fosberg kindly communicatedby Dr. D.H.Nicolson(US).Fosberg wrote: "Theseveralfrag- mentsofForstermaterialavailableareeithersterileor havesubulateglumes (=Lep- turus repens var. subulatusFosberg). Only the drawing in BMhastheacute glumes ofmy var. repens andis herechosenas thelectotypeto maintainthespecies namein thesenseofitsawnless aspect". Fosberg notedthatsyntypes inBM,K,P, S, UPS,and Whavetheawnsofvar. subulatus,whilethoseinFR, GOET, andUPS are sterile. 2. A collectionof thisspecies by R. BrownfromKupang, Timor,was distributed under the name L.filiformis. This seems to be a herbariumname and was never published by Brown. Lepturusfiliformis(Roth) Trin. (=Parapholisfiliformis (Roth) C.E.Hubb.)is a quitedifferentEurasianspecies. b. var. pubescens Nowack, var. nov. Plantacaespitosastoloniferaculmis ca.0.3 m longis. Differt avarietate typica culmis vaginisfoliis pedunculisetglumisdensepubescentibus.Spicaca. 10cmlonga1.5 mm lata internodiis 5-7 mm longis paene glabris.Glumae superiores acuminatae 12-15 mmlongae.—Typus: A.vanLeeuwen PAMt-3(holoL). Culmsc.0.3mlong; culms, sheaths,blades,peduncles, andglumes denselypubescent. Spikes c. 10cm by 1.5mm;internodes5-7 mmlong, sparsely hairy. Upperglumes acuminate, 12-15mmlong. Distribution— New Guinea,HalmaheiraSea,PamIs., KofiauIs, Habitat— Sandy beaches. Additional specimenseen: A. vanLeeuwen KOF (L),Halmaheira Sea,Kofiau Is. ACKNOWLEDGEMENTS Thisstudy wasmadeduringasabbatical attheNationaal Herbarium Nederland (L).Thehospitality wasverymuchappreciated.Specialthanks areduetoDr.D.H.Nicolson (US) formakingavailable the manuscriptbythe late Dr.F.R.Fosberg onthe typificationofFosternames. R.Nowack & J.F. Veldkamp: Notes onLepturus in Malesia 389 REFERENCES Beauvois,A.M.F.J.Palisot de. 1812. Essai d'une nouvelle Agrostographie: 110, 116,117, 168, 177.Fain,Paris. Brown,F. 1931.Floraofsoutheastern PolynesiaI. Monocotyledons.Bernice P.BishopMus.Bull. 84: 87-90,t. 12. Brown,R. 1810.Prodromus floraeNovae Hollandiae 1: 207. Johnson & Co.,London. Camus,A.A. 1923.Note surles genres 'Lepturus'R.Br,et 'Pholiurus' Trinius. Ann. Soc. Linn. Lyon69: 87-88. Clayton, W.D.& S.A. Renvoize. 1986. Genera graminum.Grasses ofthe world: 228-229. Her Majesty's Stationery Office,London. Dumortier,B.C.J. 1824,'1823'. Observations desGraminees du Belgique:90, 140.Casterman, Tournay. Forster,J.G.A. 1786.Florulae insularum australiumprodromus: 9.Dieterich,Gottingen. Fosberg,F.R. 1955.Pacific formsofLepturus R.Br.(Gramineae).Occas.Pap. Bernice P.Bishop Mus. 21:285- 294. Ghandi,K.N. 1996. Nomenclatural novelties for theWestern hemisphereplants. I. Harvard Pap. Bot. 8:63-66. Hackel,E. 1889.Andropogoneae.In:A.L.P.P. de Candolle,Monographiaephanerogamarum6: 320. Masson,Paris. Hansen,I. &E. Potztal. 1954. BeitragezurAnatomie und Systematik derLeptureae. Bot. Jahrb. Syst. 76:251-257. Hilu,K.W. &L.A. Alice.2000.Phylogeneticrelationshipsin subfamily Chloridoideae (Poaceae) based onmatK sequences: apreliminaryassessment.In: S.W.L.Jacobs & J.Everett, Grasses: systematics andevolution: 173-179. CSIROPublishing,Collingwood. Hitchcock,A.S. 1920.The genera ofgrasses ofthe United States,with specialreference to the economicspecies. USDABull. 772: 105. Hubbard,C.E. 1974.Lepturus. In:W.D.Clayton,S.M.Phillips&S.A.Renvoize,Floraoftropical EastAfrica.Gramineae,Part 2: 389. CrownAgents,London. Rafinesque,C.S. 1819.The genera ofNorth-American plants. Amer. MonthlyMag. & Crit.Rev. 4: 190. Scholz,H. 1995.Monerma P. Beauv. (Poaceae)-notanillegitimatename. Feddes Repert.Spec. Nov.RegniVeg. 106: 169-171. Steudel,E.G.June 1854.Gramineae. In:H. Zollinger,SystematischeVerzeichniss: 57. Kiesling, Zurich. Steudel,E.G. July-Nov. 1854.Synopsis plantarumglumacearum1: 357,430.Metzler,Stuttgart. Vanden Borre,A.&L.Watson. 2000.Ontheclassification oftheChloridoideae: resultsfrommor- phologicalandleafanatomicaldataanalyses.In:S.W.L.Jacobs&J.Everett,Grasses:systematics andevolution: 180-183. CSIROPublishing,Collingwood. Veldkamp,J.F. 1999.Newrecords ofGramineae for Malesia. Fl.Males. Bull. 12: 233

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