12
MYRIOGRAMME MELANESIENSIS AND M. HETEROSTROMA (DELESSERIACEAE, RHODOPHYTA), TWO NEW SPECIES FROM THE SOLOMON ISLANDS AND VANUATU Antoine D. R. N’Yeurt Université de la Polynésie française B. P. 6570 Faa’a, 98702 Tahiti, French Polynesia Michael J.Wynne University of Michigan Herbarium, 3600 Varsity Drive Ann Arbor, Michigan 4808-2287 Claude E. Payri UMR 738 Systématique, Adaptation, Evolution IRD-Nouméa–BPA5, 98848 Nouméa Cedex, New Caledonia ABSTRACT. Two new species of the red algal genus Myriogramme, M. melanesiensis and M. hetero- stroma, are described from subtidal habitats in the Solomon Islands and Vanuatu, South Pacific. Myrio- gramme melanesiensis is characterized by a tristromatic thallus with layers composed of subrectangular cells of approximately equal height, smooth margins lacking any spines or proliferations, and multiple ovoid to elongate two-layered, submarginal tetrasporangial sori. Myriogramme heterostroma is distin- guished by its tri- to pentastromatic thallus, with a single large layer of clear medullary cells and smaller pigmented cortical cells, and tetrasporangia occurring in single, large, median sori. Previous published records of the marine algal flora of the Solomon and Vanuatu Islands are few and scattered. From the Solomon Islands, the most comprehensive work to date is that of Womersley and Bailey (970), listing some 233 species.To our knowledge there are no previously published records of the marine algae of Vanu- atu, although some seven species were seen by the first author in collections of the Natural History Museum, London (BM); this Vanuatu material is poorly documented and very old. A few anonymous and undated collections of Halimeda from Vanuatu are recorded in Hillis-Colinvaux’s monograph (Hillis 959). The Solomon Islands occupy a total land area of 27,556 km 2 spread over 992 islands aligned over more than ,500 km in the southwestern Pacific Ocean, encom- passing coral reefs amongst those with the highest global biodiversity (Lovell et al. 2004; Payri et al. 2004). Vanuatu is an archipelago of some 93 islands with a total land area of over 4 765 km² aligned over 800 km, located 00 km to the southeast of the Solomon Islands. Recent collections from the Solomon Islands and Espiritu Santo, the largest island of Vanuatu, sampled by the third author from June to July 2004 and August 2006, respectively, have yielded many new records from previously unexplored habitats (Payri et al. 2004, unpubl.). The genus Myriogramme was erected by Kylin (924) and currently comprises some 28 species. In this paper we report two new, locally abundant, subtidal benthic marine species based on collections from the Solomon Islands and Vanuatu. 23 Contr. Univ. Michigan Herb. 25: 23–224. 2007.

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

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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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2007 N’YEURT,WYNNE&PAYRI:TWONEWSPECIESOFMYRIOGRAMME 2�5

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2�6 CONTR.UNIVERSITYOFMICHIGANHERBARIUM VOLUME25

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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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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