Transcript
Page 1: mYRIOGRammE mElaNESIENSIS aND m. hETEROSTROma ...mwynne/pdffiles/NYeurtetal.pdf · Collection and analysis. Material was collected by SCUBA; part was stored in 5% buffered Formalin

2007 N’YEURT,WYNNE&PAYRI:TWONEWSPECIESOFMYRIOGRAMME 2�3

mYRIOGRammE mElaNESIENSIS aND m. hETEROSTROma (DElESSERIacEaE, RhODOphYTa), TWO NEW SpEcIES

FROm ThE SOlOmON ISlaNDS aND VaNUaTU

AntoineD.R.N’YeurtUniversitédelaPolynésiefrançaise

B.P.6570Faa’a,98702Tahiti,FrenchPolynesia

MichaelJ.WynneUniversityofMichiganHerbarium,3600VarsityDrive

AnnArbor,Michigan48�08-2287

ClaudeE.PayriUMR7�38Systématique,Adaptation,Evolution

IRD-Nouméa–BPA5,98848NouméaCedex,NewCaledonia

AbstrAct. TwonewspeciesoftheredalgalgenusMyriogramme,M. melanesiensisandM.hetero-stroma,aredescribedfromsubtidalhabitatsintheSolomonIslandsandVanuatu,SouthPacific.Myrio-gramme melanesiensisischaracterizedbyatristromaticthalluswithlayerscomposedofsubrectangularcellsofapproximatelyequalheight,smoothmarginslackinganyspinesorproliferations,andmultipleovoid toelongate two-layered, submarginal tetrasporangial sori.Myriogramme heterostroma isdistin-guishedbyitstri-topentastromaticthallus,withasinglelargelayerofclearmedullarycellsandsmallerpigmentedcorticalcells,andtetrasporangiaoccurringinsingle,large,mediansori.

PreviouspublishedrecordsofthemarinealgalfloraoftheSolomonandVanuatuIslandsarefewandscattered.FromtheSolomonIslands,themostcomprehensiveworktodateisthatofWomersleyandBailey(�970),listingsome233species.ToourknowledgetherearenopreviouslypublishedrecordsofthemarinealgaeofVanu-atu,althoughsomesevenspecieswereseenbythefirstauthorincollectionsoftheNaturalHistoryMuseum,London(BM);thisVanuatumaterialispoorlydocumentedandveryold.AfewanonymousandundatedcollectionsofHalimedafromVanuatuarerecordedinHillis-Colinvaux’smonograph(Hillis�959).

TheSolomonIslandsoccupyatotal landareaof27,556km2spreadover992islandsalignedovermorethan�,500kminthesouthwesternPacificOcean,encom-passingcoralreefsamongstthosewiththehighestglobalbiodiversity(Lovelletal.2004;Payrietal.2004).Vanuatu isanarchipelagoofsome93 islandswithatotallandareaofover�4765km²alignedover800km,located�00kmtothesoutheastoftheSolomonIslands.RecentcollectionsfromtheSolomonIslandsandEspirituSanto,thelargestislandofVanuatu,sampledbythethirdauthorfromJunetoJuly2004andAugust2006,respectively,haveyieldedmanynewrecordsfrompreviouslyunexploredhabitats(Payrietal.2004,unpubl.).

ThegenusMyriogrammewaserectedbyKylin(�924)andcurrentlycomprisessome28species.Inthispaperwereporttwonew,locallyabundant,subtidalbenthicmarinespeciesbasedoncollectionsfromtheSolomonIslandsandVanuatu.

2�3

Contr.Univ.MichiganHerb.25:2�3–224.2007.

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2�4 CONTR.UNIVERSITYOFMICHIGANHERBARIUM VOLUME25

MATERIALSANDMETHODS

Collection and analysis.MaterialwascollectedbySCUBA;partwasstoredin5%bufferedFormalininseawater,andtherestwasdriedasherbariumspecimens.Driedmaterialwasre-hydratedinweakdetergentsolutionpriortosectioningwithafreezingmicrotome.Sectionswerestainedusing�%anilinebluein60%clearcornsyrup,andmadepermanentifnecessarybyaddingmore60%clearcornsyrupandallowingtodry.

Photography.MacrophotographsweretakenwithaNikonE-995digitalcam-era(NikonCorporation,Tokyo,Japan).PhotomicrographswereobtainedusinganOlympusBH2compoundmicroscopefittedwithanOlympusC-5050digitalcamera(OlympusOpticalCo.Ltd.,Tokyo,Japan),andtheresultingfilesprocessedintofig-uresbycomputersoftware.

Herbarium specimens.VoucherspecimenshavebeendepositedintheherbariaofPC,NOU-IRD(PhycologicalHerbarium,InstitutdeRecherchepourleDével-oppement,Nouméa,NewCaledonia),andMICH.HerbariumabbreviationsareinaccordancewithHolmgrenetal.(�990).Accessionnumbersprecededbytheletter‘S’refertomicroscopeslidecollections.

RESULTS

cerAmiALes: deLesseriAceAe

myriogramme melanesiensis N’Yeurt,M.J.Wynne&Payri,sp.nov.—type: soLomoN isLANds.NggelaSule,MbuganaIsland,09°��.055'S,�60°��.982'E,–�0m,��Jul2004,C. E. Payri (holotype:PC0062772,tetrasporangial;isotype:NOU-IRDS483,tetrasporangial). Figs.�–8.

Thallus 5–�5 mm altus, roseus, irregulariter lobatus. Laminae 75–80 µm incrassitudine,tristromaticae,stratiscellularumcrassitudinecircaaequalibus.Cellulaerectangulares-subquadrataeinsectionetransversali,circa25×50µm.Margineslami-narumlaeves;partesdistalesclavatae-rotundatae,sinedentibus;partesproximalesligulatae vel irregulariter dichotomae. Rhizoidea marginalia 90–�00 µm diametroet �.0–�.2 mm longa, ex 6–7 filis multicelliaribus fasciculatis constantia, secusperipheriamthallidispersa.Chloroplastiperipheralesetnumerosi.Tetrasporangiadivisacrucialiter,60–90µmdiametro,insorisdispersisovalibusvelelongatistaeni-formibusque,monostromaticisveldistromaticis,0.5–�.0mmlatiset�.8–4.0mmlongis,inpartibusdistalibuslaminaedispositis.

Thallus5–�5mmhigh,pinkish red, irregularly lobed.Blades75–80µm thick,tristromatic,withcelllayersofapproximatelyequalthickness.Cellsrectangulartosubquadrateintransversesection,ca.25×50µm.Marginsofbladessmooth,distal

FIGS.�–8.Myriogramme melanesiensis (PC0062772exceptwhereotherwise stated).�.Habitofholotype;scale=�0mm.2.ProximalregionofolderbladefromVanuatu(C. E. Payri(NOU-IRD�396),showing‘nerve’region(n)[thecross-likerunneronthenerveisanepiphyticbryozoan];scale=200µm.3.Surfaceviewofcorticalcells;scale=50µm.4.Transversesectionofthethallus,showingthemedullarylayer(m)ofthesameheightascorticallayers(c);scale=25µm.5.Detailofmulticellularmarginalrhizoids(arrow);scale=�00µm.6.Detailoffertilesubmarginalregionofblade,showingelongate,ribbon-liketoovoidtetrasporangialsori(arrows);scale=700µm.7.Detailoftetrasporangia;scale=50µm.8.Crosssectionofatwo-layeredtetrasporangialsorus(t);scale=50µm.

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portions clavate–rounded, without dentations; proximal portions strap–shapedto irregularlydichotomous.Amedian‘nerve’ region400–600µmwidepresent inproximalareasof some larger,older thalli, consistingof largecells70–�00µm indiameter.Marginalrhizoids90–�00µmindiameter,�.0–�.2mmlong,composedof6–7bundledmulticellularfilaments,scatteredalongperipheryofthallus.Chloroplastsperipheralandnumerous.Tetrasporangiacruciatelydivided,60–90µmindiameter,inscatteredovaltoribbon-like,elongate�-to2-layeredsori0.5–�.0mmwide,�.8–4.0mmlong,indistalportionsofblade.

Distribution.SolomonIslandsandVanuatu;atdepthsof3–35moncoastalreefslopes,nestedamidstbranchedcorals.

pArAtype. Solomon Islands.Makaira,SanCristobalIsland,20Jul2004,–�2m,C. E. Payri(NOU-IRDS482).

AdditioNAL specimeNs exAmiNed.Solomon Islands. HelePassage,VangunuIsland,04Jul2004,–3m,C. E. Payri(NOU-IRD478,477);KoloLagoon,NewGeorgiaIsland,8Jul2004,–20m,C. E. Payri(NOU-IRD474,475;SUVA-A);NuguMatthewsShoal,Guadalcanal,�0Jul2004,–35m,C. E. Payri(NOU-IRD48�);NggelaIsland,�2Jul2004,–0.8m,C. E. Payri(NOU-IRD476). Vanuatu.EspirituSanto,24Aug2006, C. E. Payri(NOU-IRD�396,tetrasporic).

ThespecificepithetreferstoMelanesia,theregionwherethisspecieshasbeencollected.

myriogramme heterostroma N’Yeurt,M.J.Wynne&Payri,sp.nov.—type: soLomoN isLANds.TanavulaPoint,NggelaSule,09°02.804'S,�60°03.7��'E,��Jul2004,–�0m,C. E. Payri(holotype:PC0062773,tetrasporangial). Figs9–�6.

Thallus 3–�2 mm altus, iridescens, roseus, irregulariter lobatus vel ligulatusanastomosibus frequentibus. Laminae 70–80 µm in crassitudine, ex 3–5 stratiscellularum inaequaliter altis constantes. Stratum medullosum singulare, excellulis magnis subquadratis hyalinis constans; cortex ex �–2 stratis cellularumpigmentiferarumgradatimminorum5–�8µmdiametroconstans.Cellulaeextimaeparvae, subsphaericae, 5–7 µm diametro, dense pigmentiferae. Cellulae corticalesinterioressparsae,7.5–�8.0µmdiametro,gangliiformes,numerosisconjunctionibussecundariis instructae. Margines laminarum laeves; partes distales lobatae, sinedentibus. Rhizoidea marginalia nulla; thallus affixus per haptera dispersa latamarginalia.Chloroplastiperipheralesetnumerosi.Tetrasporangiadivisacrucialiter,�8–20µmdiametro,insorismagnis,distromaticis,singularibus,medianis�500–�800µmdiametro,inpartibusdistalibuslaminaedispositis.

Thallus3–�2mmhigh,slightlyiridescentpinkishredincolor,irregularlylobedtostrap-shaped,withfrequentanastomoses.Blades70–80µmthick,composedof3–5celllayers,ofunequalheight.Medullarylayersingle,oflargesubquadratehyalinecells40–42µmindiameter;cortexcomposedof�–2layersofprogressivelysmallerpigmentedcells5–�8µmindiameter.Outermostcellssmall,subsphaerical,5–7µmindiameter,denselypigmented.Innercorticalcellssparse,7.5–�8.0µmindiameter,ganglionic, with multiple secondary pit connections. Margins of laminas smooth,distal portions lobed, without dentations. Marginal rhizoids absent; attachmentvia scattered broad marginal haptera. Chloroplasts peripheral and numerous.Tetrasporangiacruciatelydivided,�8–20µmindiameter,inlarge,single,median2-layeredsori�500–�800µmindiameter,indistalportionsofblade.

Distribution.Solomon Islands; atdepthsof6 to20monvertical reef slopes,nestedamidstbranchedcoral.

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2007 N’YEURT,WYNNE&PAYRI:TWONEWSPECIESOFMYRIOGRAMME 2�7

FIGS.9–�6.Myriogramme heterostroma(PC0062773).9.Habitofholotype;scale=5mm.�0.Detailofthallus,showingdenselayerofsmalloutercorticalcellsandbladeanastomosis(arrow);scale=�mm.��.Surfaceviewofcorticalcells;scale=20µm.�2.Detailofamarginalattachmentpad;scale=20µm.�3,�4.Transversesectionsintwodifferentregionsofthethallus,showingalarge,clearsinglemedullarylayer(m)andamuchsmaller,pigmentedcorticallayers(c);scale=�0µm.�5.Detailofalargemediantetrasporangialsorus(s);scale=200µm.�6.Detailoftetrasporangia(arrow);scale=20µm.

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2�8 CONTR.UNIVERSITYOFMICHIGANHERBARIUM VOLUME25

pArAtype.Solomon Islands.AnutaPainaIsland,Malaita,�8Jul2004,C. E. Payri(NOU-IRDS484,tetrasporangial).

AdditioNAL specimeNs exAmiNed.Solomon Islands.RendovaIsland,NewGeorgia,–20m,5Jul2004, C. E. Payri(SUVA-A);Honiara,GuadalcanalIsland,9Jul2004,–20m,C. E. Payri(NOU-IRD480).

Thespecificepithetreferstothedifferentsizesofthecellsofthemedullarylayerandthecorticallayers.

DISCUSSION

Probably more than any other genus in the Delesseriaceae, Myriogrammehaslongservedasa“catch-all“genus,towhichmanydisparatespecieshavebeenassigned.AsWynne(�983)stated,theill-definedcircumscriptionofthisgenusresult-edfromKylin’s(�924)basingmuchofhisaccountofMyriogrammeonM.minutaKylin,aMediterraneanspecies,yetdesignatingM.livida(J.D.Hook.&Harv.)Kylin,aspeciesfromtheFalklandIslands,asthetypeofthegenus.Kylin(�924)assignedatotalof�8speciestohisnewgenus,andtheformsofthethalliexpressedinthesespecies show much variation, including very small to large, robust habit, veinlessbladestobladeswithpronouncednerves,andgreatvariabilityinthepositionoftet-rasporangialsori.Whereknown,thecystocarpsinspeciesofMyriogrammeweresaidtoproducecarposporangiainchains.Wynne(�983)notedthatcriticalexaminationofthetypespeciesisnecessaryforapropercircumscriptionofMyriogramme.SuchastudyofM.lividawasundertakenbyHommersandandFredericq(�997a),andfol-low-uppublications(Hommersand&Fredericq�997b;Linetal.200�)contributedto elucidating the characters of sometimes similar-appearing genera. In addition,overtheyearsanumberofspeciesformerlyassignedtoMyriogrammehavebeentransferred other genera, as the circumscription of Myriogramme became morepreciselydrawn.SuchattritionhasbeencarriedoutbyZinova(�98�),whobasedHaraldiophyllumonMyriogrammebonnemaisoniiKylinandHideophyllumonM.yezoensisYamada&Tokida(Yamada�935).MaggsandHommersand(�993)trans-ferredM.minutaandHaraldiophyllumheterocarpum[=Myriogrammeversicolor]toErnstandFeldmann’s(�957)genusDrachiella.Myriogrammeerosa(Harvey)KylinwastransferredtoHaraldiophyllumbyMillarandHuisman(�996).WhenLinetal.(2004)erectedAugophyllum,theyincludedinitMyriogrammemarginifructaR.E.Norris&M.J.Wynne(�987).Womersley(2003)basedhisnewgenusNitospinosaonMyriogrammepristoidea(Harv.)Kylin.Despitethisongoing“cleansing”ofthegenus,therestillremainanumberofpoorlyknownspeciesassignedtoMyriogramme.

Thetri-topentastromaticnatureofthebladesandtherarepresenceofabasalnerveinspecimensofM. melanesiensisandM.heterostromaareamongthefeaturesthatledustoconcludethatthesetworelativelycommonlyfoundalgaeintheSolo-monandVanuatuIslandsbelongtoundescribedspecies.WeareabletoexcludethegreatmajorityofspeciesofMyriogramme(andsomerelatedgenera)fairlyeasily,inparticular,thosespecieswithbladesbearingpronouncednervesthroughouttheirthalli.InM. melanesiensis,thepresenceofamedian“nerve”regionappearstobearelativelyrarefeaturestronglydependentonageandmaturityofthethallusandismorepronouncedinproximalareasofsomelargerplants.Themajorityofplantsinagivenpopulationdonotshowanyclearnervesorveins.WehavecomparedthetwonewspecieswithagroupofsomewhatsimilarspeciesinTable�,wherewelistimportantmorphologicaltraits.

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2007 N’YEURT,WYNNE&PAYRI:TWONEWSPECIESOFMYRIOGRAMME 2�9

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220 CONTR.UNIVERSITYOFMICHIGANHERBARIUM VOLUME25

Myriogrammealliacea(P.Crouan&H.Crouan)Athanasiadis,aspeciesoccur-ring on the coast of Brittany, France (Crouan & Crouan �85�), has a distinctiverose-winecolorinwell-preservedsamples.Thebladeshavesmoothmarginsandaretristromaticforalmostalloftheirlength.Thecellsincrosssectionareofuniformsize.Tetrasporangiaarearrangedinsmallsorioverthecentralpartofthethallus(Magne�959);however,thalliaretaller(3–6cm)thanthoseinthenewspecies,andtheyreleaseoftheodorofonionswhenthalliareimmersedinfreshwaterandthenemersed,adistinctivefeatureofM.alliacea(Crouan&Crouan�85�).

AnotherspeciesofMyriogrammewithatristromaticorganizationisM.goaensisV.Krishnamurthy&Varadarajan(inKrishnamurthy�992),described fromIndia.Onlytetrasporangiatethalliweredescribed,andthetetrasporangiaweresaidtobeinscattered,smallsori(Krishnamurthy&Varadarajan�990),buttheirFig.�3depictsabladewithonlyasinglelargesorus.Also,althoughthethalliweresaidtobe“stipitate,deeplydividedintosmall,sinuate,lineartolanceolateblades,”thebladesinFig.�3appeartobesimple.ThepresenceofmarginalteethlikeprojectionswithtransverselydividingapicalcellswouldseparatethisIndianspeciesfrombothnewspeciesfromtheSolomonIslandsandVanuatu.

AnotherspeciesfromIndiaisMyriogrammequiloneensisAnilKumar&Pan-ikkar(�993).Thalliareessentiallydistromatic throughout,and theblademarginsbearshortspine-likeprojectionsaswellastuftsofrhizoidalinitialsforattachmentpurposes,whichdistinguisheditfromthenewspecies.

Myriogrammerepens,knownfromthePacificcoastofNorthAmerica(Hollen-berg�945)hasarhizomebearingsimplebladeslessthan�0mminheightandwithverysmall,scatteredsorioftetrasporangia.Thesefeaturesaswellasthemonostro-maticnatureofthebladesdistinguishitfromthenewspecies.

Myriogramme prostrata was originally described as Haraldia prostrata fromPacificMexico(Dawsonetal.�960),butwaslaterreportedfromthetropical-sub-tropicalwesternAtlantic(Wynne�990;Littler&Littler2000;Ballantineetal.2004),theSouthPacific(Skelton&South2002;South&Skelton2003).LittlerandLittler(2000)describedthebladesasexhibitinga“blue-greensheen,”thussomewhatiri-descent likeM. heterostroma.Myriogrammeprostratawasalsoreported,butwithaquery,fromKwazulu-Natal,SouthAfrica(DeClercketal.2005),butthatplant,withamottled,stronglyiridescentaspectandwithnumerousmarginalteeth,seemsunlikegenuineM.prostrata,whichisbasicallyaprostratealgawithfrequentmarginalrhizoidsforattachment.Amonostromatic,marginallydentatebladearisingfromaprostrateaxis,withmarginalrhizoids,andbearinglargescatteredtetrasporangialsoriwasreportedandillustratedfromFijiintheSouthPacific,atfirstasMyriogrammesp.(Southetal.�993)and laterasM.prostrata (South&Skelton2003),andalsofromSamoa(Skelton&South2002).Theoccasionalsmallmarginaldentations,themonostromaticnatureoftheblades,aswellasthepresenceofscatteredlargetet-rasporangialsoriconfinedtomarginalregionsalldistinguishM.prostratafromthenewspecies.

Myriogrammecartilaginea(Harvey)Womersleyisalittle-knownspeciesoccur-ringinWesternAustralia(Harvey�855),butcystocarpicplantshavenotyetbeenobserved and the assignment to Myriogramme uncertain (Womersley 2003). Itsthalliare4–8cmtallandwithabundantirregularlyalternatebranching,oftenwithcrispate margins and a cartilaginous texture.The blades become tristromatic andlaterbecomepolystromaticinthecentralregions.Thesefeatureseasilyseparateitfromthenewspecies.

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2007 N’YEURT,WYNNE&PAYRI:TWONEWSPECIESOFMYRIOGRAMME 22�

Myriogramme variegata Yamada, with a type locality of Sagami Bay, Japan(Yamada�944),hasalsobeenreportedfromCalifornia(Abbott&Hollenberg�976).Itconsistsofadiscoidbasegivingrisetostipitatesimple,orrarelydivided,smallbladesreachingonly�.5cminheight.Thebladesaremonostromaticandweresaidtohavemarginsbothentireandminutelydentate.Tetrasporangialsoriaresmallandproducedontheleafypartsoftheblade(Yamada�944).ThemonostromaticnatureofthebladeandtheminutemarginalteethdistinguishM.variegatafromthenewspecies.

MyriogrammedistromaticaBoudouresquewasbasedonmaterialintheThuret-BornetHerbariuminParisbearingthemanuscriptname“Nitophyllumdistromati-cumRodriguez.”ThetypewasaRodriguezcollectionofMay,�897,fromasubmarinecaveatadepthof90moffPortMahon,Minorca,BalearicIslands,inthewesternMediterranean.ThisspecieshasalsobeenreportedfromdeepwatersoffthecoastofNorthCarolinaandSouthCarolina(Schneider&Searles�99�).Thesmall(to�5mmacross)prostratebladesareconsistentlydistromatic,withtheventrallayerofcellslargerthanthoseofthesurfacelayer.Multicellularrhizoidsarefrequentontheundersurfaceandattachthebladetothesubstratum.Reproductionisnotknowninthisspecies.ThesingleparietalfenestratechloroplastpercellclearlyindicatesthatthisspeciesdoesnotbelongtoMyriogramme(Hommersand&Fredericq�997a).

NitophyllumtristromaticumRodriguezexMazza(Mazza�903)bearsdiscussioninthatBoudouresqueetal.(�984)intendedtotransferittoMyriogramme,buttheircombination was not validly published (Greuter et al. 2000).This Mediterraneanspecies,describedfromMinorca,BalearicIslands(Mazza�903),hasatristromaticorganization,andthereisacentrallayerofrelativelylargecellsandcorticallayersofsmallercells,asoccursinM. heterostroma.AccordingtoGómezGarretaetal.(200�),thecellsbearasingleparietal,platelikechloroplastpercell,afeaturethatprecludesplacingthisspecieswithineitherNitophyllumorMyriogramme.

Schizoserisbombayensis(Børgesen)WomersleyisanothersmallDelesseriaceanspecies with a wide range in tropical and warm temperate waters of the PacificOcean,includingtheSouthPacific(Millar&Kraft�993;Abbott�999;Littler&Lit-tler2003;Lobban&N’Yeurt2006).Thebladesinthisspeciesareusually�–2(–5)cmhigh, irregularly todichotomouslybranched,withmedianmacroscopicnervesusuallycoursingfromthebasetodistalregionsofthethallus(Abbott�999;Womer-sley2003).Thebladesaremonostromaticinmostregionsofthethallusbetweenthenerves(Børgesen�93�;Segawa�94�,asMyriogrammesubdichotoma;Dawson�950,asS.pygmaea)butmaybecomedistromaticinolderregions(Børgesen�93�).Thechloroplastshavebeenshowntobedissectedandribbon-like(Lobban&N’Yeurt2006),andthespecieshasneverbeenreportedasiridescent.Thesedifferencesdis-tinguishthisspeciesfromthenewspeciesofMyriogramme.

AnotherDelesseriaceanalgawithsuperficialresemblancetoMyriogrammemela-nesiensisandM. heterostroma isDrachiellaminuta (Kylin)Maggs&Hommersand.BasedonacollectionfromthevicinityofNaples,Italy(Kylin�924),thisspecieswasdescribedasnon-stipitatesmallblades,�–2cmtall,irregularlysinuateorlobed,distallymonostromatic,buttristromaticbelow,andlackingveins.Tetrasporangialsoriaresmallandscatteredinthedistalportionsoftheblades,andthetetrasporangiaareproducedintwolayers.Magne(�956,�957)provideddetailedobservationsonthedistinctivecellularorganizationofthisspeciesanditsdistinctiveprocessofmanychloroplastsundergoingafusionprocesstoformasinglelobedchloroplast.MaggsandHommer-sand(�993)describedspecimenstobeofrareoccurrenceinsouthwesternEngland

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222 CONTR.UNIVERSITYOFMICHIGANHERBARIUM VOLUME25

andtheChannelIslands,wheretheyformthalli5cmhighand5cmwide.Thethallibecomedecumbentandattimesshowastrongblueiridescence.Thebladeincrosssectionshowsthecelllayerstobeofuniformsize.Tetrasporangiaareproducedinsoridirectlyontheprimarybladeinthisspecies,quiteunliketheproductionofsmallspecializedtetrasporophyllsinDrachiellaspectabilisErnst&Feldmann(�957),thetypeofthegenusDrachiella.ThisledWynne(�994)toexpressdoubtoftheplace-mentofM.minutainDrachiella.Itseemsthattheprimaryjustificationforthepro-posedtransferbyMaggsandHommersand(�993)isthepresence,asinD.spectabilis,ofonelargeribbon-liketoconvolutedchloroplastpercellratherthanmanysmalldiscoidchloroplastspercell.ThiscautionledGómezGarretaetal.(200�)toretainM.minutainMyriogramme.Thisspeciesisdiffersfrombothnewspeciesinitsdif-ferentcellularorganization.

Myriogramme melanesiensis and M. heterostroma increase our knowledge ofalgaldiversityinthisregionoftheworld,andenrichtheDelesseriaceaefromtheSolomonIslands,forwhichWomersleyandBailey(�970)hadnotreportedthegenusMyriogrammeamongthesixtaxalistedforthisfamily.

Key to the species of myrioGrAmme from the soLomoN isLANds ANd vANuAtu

�. Medullarylayerofapproximatelysamedimensionsascorticallayers;attachmentviamarginalrhizoids;tetrasporangiainmultipleirregularmarginalsori. M. melanesiensis

�. Medullarylayermuchlargerthancorticallayers;attachmentviamarginalhaptera;tetrasporangiainlarge,singlemediansori. M. heterostroma

ACKNOWLEDGMENTS

ADRNandCEP thank theCoralReef Initiative for theSouthPacific (CRISP)programme forfinancialsupporttowardsthisstudy,andCEPisgratefultothecrewoftheIRDoceanographicvessel‘ALIS’andthediversfromIRD.WethankMr.GregoryLasneforthephotographsofherbariumsheets,andCraigW.Schneiderforhishelpfulreviewofthemanuscript.

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