Table Of ContentBLUMEA 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