rhodymenia coccocarpa, a new combination …mwynne/pdffiles/wynne.pdf · the red alga phyllophora...

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RHODYMENIA COCCOCARPA, A NEW COMBINATION BASED ON PHYLLOPHORA COCCOCARPA MONT. Michael J.Wynne University of Michigan Herbarium 3600 Varsity Drive Ann Arbor, Michigan 4808-2287 ABSTRACT. The new combination Rhodymenia coccocarpa (Mont.) M. J.Wynne, based on Phyllophora coccocarpa Mont., is proposed; it takes precedence over the nomen novum Rhodymenia subantarctica Ricker. The red alga Phyllophora coccocarpa was described by Montagne (852b) on the basis of a d’Urville collection from the Strait of Magellan. The alga was described as a stipitate frond, with a bare, terete or broadened stipe expanding into flattened cuneate blades proximally and having linear bifid enervate apical portions, with rotund lobes. Cystocarps were said to be on the distal segments, sessile and globose, with the center (in the dried state) depressed, acorn-shaped and ostiolate. According to Montagne, part of his earlier concept (Montagne 845) of Rhodymenia palmetta (J.V. Lamour.) Grev. [current name: Rhodymenia pseudopalmata (J.V. Lamour.) P. C. Silva] belonged within his new species Phyllophora cococcarpa. Montagne (854) later repeated his description of P. coccocarpa and also provided a single figure of its habit. In all of his accounts of this species, Montagne (852b, 854, 856) described his material as fertile, referring to cystocarps [“conceptaculis”]; indeed that is what he depicted in the habit illustration (Montagne, 854, pl. 6, fig. ), which is repro- duced here as Fig. .Yet, according to Hariot (889) Montagne’s Herbarium (in PC) contains a single non-reproductive specimen of original material of Phyllocarpa coc- cocarpa, which Hariot treated as Epymenia membranacea Harv., currently known as Rhodymenia obtusa (Grev.)Womersley (Womersley 996). I asked Dr. Bruno de Reviers to check on authentic material of Phyllocarpa coc- cocarpa in PC. He kindly informed me that he located in the Montagne herbarium two elements belonging to the protologue: a single specimen of the holotype (PC 0060225; MA 4824) (Fig. 2) and an original and unpublished sketch of the anatomy (MS 434, p. 4809) (Fig. 3); scanned images of these elements were provided to me. He did not find any additional specimen in any place in PC. It is clear that the holotype corresponds perfectly to Montagne’s (854) plate 6, fig. , except that the speci- men does not show any cystocarps. Figure 4 is a closer view of distal regions of the holotype showing the absence of cystocarps. This observation confirms that Hariot was correct but renders doubtful Montagne’s description of the cystocarps. It is pos- sible that Montagne had another, small, fertile specimen that he used for adding the observations on cystocarps, but there is no evidence of such material. The sketch (Fig. 3), which is very faint, is accompanied by the following nota- tions: “Phyllophora coccocarpa Montg.”, “Rhodymenia palmetta olim”, “Magellan”, “D’Urville” and “Hb. de C. Montagne.”The sketch is of a portion of a cross section, showing cortical, subcortical and medullary cells, an organization that conforms to the anatomy of Rhodymenia. In addition to the more usual pattern of dichotomous to irregular branching and marginal proliferation, a distinctive mode of producing new branches from a thickened 24 Contr. Univ. Michigan Herb. 25: 24–247. 2007.

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Page 1: RhODYmENIa cOccOcaRpa, a NEW cOmBINaTION …mwynne/pdffiles/Wynne.pdf · The red alga Phyllophora coccocarpa was ... He kindly informed me that he located in the Montagne herbarium

2007 WYNNE:RHODYMENIACOCCOCARPA 24�

RhODYmENIa cOccOcaRpa, a NEW cOmBINaTION BaSED ON phYllOphORa cOccOcaRpa mONT.

MichaelJ.WynneUniversityofMichiganHerbarium

3600VarsityDriveAnnArbor,Michigan48�08-2287

AbstrAct. ThenewcombinationRhodymenia coccocarpa(Mont.)M.J.Wynne,basedonPhyllophoracoccocarpaMont., isproposed; ittakesprecedenceoverthenomennovumRhodymenia subantarcticaRicker.

TheredalgaPhyllophoracoccocarpawasdescribedbyMontagne(�852b)onthebasisofad’UrvillecollectionfromtheStraitofMagellan.Thealgawasdescribedasastipitatefrond,withabare,tereteorbroadenedstipeexpandingintoflattenedcuneate blades proximally and having linear bifid enervate apical portions, withrotundlobes.Cystocarpsweresaidtobeonthedistalsegments,sessileandglobose,withthecenter(inthedriedstate)depressed,acorn-shapedandostiolate.AccordingtoMontagne,partofhisearlierconcept(Montagne�845)ofRhodymeniapalmetta(J.V.Lamour.)Grev.[currentname:Rhodymeniapseudopalmata(J.V.Lamour.)P.C.Silva]belongedwithinhisnewspeciesPhyllophoracococcarpa.Montagne(�854)laterrepeatedhisdescriptionofP.coccocarpaandalsoprovidedasinglefigureofitshabit.Inallofhisaccountsofthisspecies,Montagne(�852b,�854,�856)describedhismaterialasfertile,referringtocystocarps[“conceptaculis”];indeedthatiswhathedepictedinthehabitillustration(Montagne,�854,pl.�6,fig.�),whichisrepro-ducedhereasFig.�.Yet,accordingtoHariot(�889)Montagne’sHerbarium(inPC)containsasinglenon-reproductivespecimenoforiginalmaterialofPhyllocarpacoc-cocarpa,whichHariottreatedasEpymeniamembranaceaHarv.,currentlyknownasRhodymeniaobtusa(Grev.)Womersley(Womersley�996).

IaskedDr.BrunodeRevierstocheckonauthenticmaterialofPhyllocarpacoc-cocarpainPC.HekindlyinformedmethathelocatedintheMontagneherbariumtwoelementsbelongingtotheprotologue:asinglespecimenoftheholotype(PC0060225;MA4824)(Fig.2)andanoriginalandunpublishedsketchoftheanatomy(MS434,p.4809)(Fig.3);scannedimagesoftheseelementswereprovidedtome.HedidnotfindanyadditionalspecimeninanyplaceinPC.Itisclearthattheholotypecorrespondsperfectly toMontagne’s (�854)plate�6,fig.�,except that the speci-mendoesnotshowanycystocarps.Figure4isacloserviewofdistalregionsoftheholotypeshowingtheabsenceofcystocarps.ThisobservationconfirmsthatHariotwascorrectbutrendersdoubtfulMontagne’sdescriptionofthecystocarps.Itispos-siblethatMontagnehadanother,small,fertilespecimenthatheusedforaddingtheobservationsoncystocarps,butthereisnoevidenceofsuchmaterial.

Thesketch(Fig.3),whichisveryfaint,isaccompaniedbythefollowingnota-tions:“PhyllophoracoccocarpaMontg.”,“Rhodymeniapalmettaolim”,“Magellan”,“D’Urville”and“Hb.deC.Montagne.”Thesketchisofaportionofacrosssection,showingcortical,subcorticalandmedullarycells,anorganizationthatconformstotheanatomyofRhodymenia.

Inadditiontothemoreusualpatternofdichotomoustoirregularbranchingandmarginalproliferation,adistinctivemodeofproducingnewbranchesfromathickened

24�

Contr.Univ.MichiganHerb.25:24�–247.2007.

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242 CONTR.UNIVERSITYOFMICHIGANHERBARIUM VOLUME25

Fig.�.ReproductionofMontagne’s(�854)pl.�6,fig.�,ofPhyllophoracoccocarpa[=Rhodymeniacoccocarpa(Mont.)M.J.Wynne].

midribregionofproximalportionsofbladescanbeobservedintheholotype(bothinthespecimenandthefigure)andinspecimensassignedtothisspeciesbyTaylor(�939).These collections, in MICH, are the following: chiLe. PuntaArenas Ma-gallanesProv.,4Feb�927,Waldo L. Schmitt 315;fALKLANdIsLANds.StanleyHarbor,EastFalklandIsland,withoutdate,Waldo L. Schmitt 98.ThisoccasionalproductionofnewbranchesfromthefaceofaparentbladewaspointedoutbyTaylor(�939).

AnannotationlabelisattachedtotheholotypeofPhyllocarpacoccocarpa,datedDecember �3, �985 with the determination“Dendrymenia skottsbergii” by MariaEmilianaRamírez.ThisnameisbasedonRhodymeniaskottsbergiiDawson(�94�),aspeciestransferredtoDendrymeniabyLevring(�960);however,suchadetermi-nationisnotadmissable,becausethenameP.coccocarpapredatesR.skottsbergii.Rhodymeniaskottsbergiiisacurrentlyrecognizedspecies,whichisendemictothe

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2007 WYNNE:RHODYMENIACOCCOCARPA 243

temperatewestcoastofSouthAmerica(Hoffmann&Santelices�997).Ithasadis-tinctivemannerofproducinglateralfronds,whicharisespirallywithinthefunnel-shapedregionatthebaseoftheparentfrond.Newbladescontinuetobesympodiallydevelopedwithanamplexicaulstructureintheregionofeachnewbranch(Dawson�94�).AccordingtoHoffmanandSantelices(�997),R.skottsbergiiisuniqueintheorderRhodymenialesinhavingthisdevelopmentalmode.Clearly,thedeterminationoftheholotypeofPhyllophoracoccocarpaasRhodymenia[Dendrymenia]skottsbergiiisincorrect.

When Skottsberg (�923) distinguished his new genus Dendrymenia fromRhodymeniaGrev.,hebasedthegenusonthesinglespeciesD.flabellifolia(Bory)Skottsberg[basionym:Sphaerococcus flabellifoliusBory(�828)].AspointedoutbyDawson(�94�),ithasbeenrecognizedthatthematerialSkottsberg“hadinhand”wasnottaxonomicallyidenticaltoBory’sspecies,andhepublishedthenewspeciesRhodymeniaskottsbergii,with the type fromConcepcionBay,Chile.At thesametime Dawson (�94�) demonstrated that genuine Rhodymenia flabellifolia (Bory)Montagne,whichalsooccursonthewestcoastofsouthernSouthAmerica,lackstheamplexicaulcharacterpresentinR.skottsbergii.Althoughsomeauthors(e.g.,Taylor,�945�947;Kylin�956;Levring�960;Santelices�989)haverecognizedDendryme-niaasdistinctfromRhodymeniaonthebasisofitssympodialdevelopment,otherauthors(e.g.,Dawson�94�;Sparling�957;Ramírez&Santelices�99�;Hoffmann&Santelices�997;Saundersetal.�999)havenot.Thecriticalpointisthatthetypeofagenus,namely,Dendrymenia,isaspecies(Art.�0.�oftheICBN),and,asnotedabove,D.flabellifolia[Sphaerococcus flabellifoliusBory] lacksthecriticaldiagnostictraitofDendrymeniaasconceivedbySkottsberg,thesympodialdevelopment;therefore,DendrymeniamustberegardedascongenericwithRhodymenia.

Fig.2.HolotypespecimenofPhyllocarpa coccocarpaMont.;PC0060225.

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244 CONTR.UNIVERSITYOFMICHIGANHERBARIUM VOLUME25

Fig.3.Originalsketch(inPC;MS434,p.4809).

PhyllophoracuneifoliaJ.D.Hooker&Harvey(�845)wasdescribedwithatypelocalityintheFalklandIslands.OneofthetwooriginalHookerspecimens(KinBM)wasdepictedbyCotton(�9�5,pl.7).AccordingtoRicker(�987),thelectotypeforPhyllophoracuneifolia[KinBM]wascollectedbyJ.D.Hookeron�April�842fromSt.SalvadorBay,EastFalklandIslands(selectedbyA.GeppandJ.H.Price).WhilereportingonnewcollectionsofthisspeciesfromtheFalklandIslandsandsouthernChile,Taylor(�939)transferredP.cuneifoliatoRhodymenia,butinsodoing,hecre-atedalaterhomonymofR.cuneifoliaOkamura(�934),thelatternamenowtreatedasGracilariacuneifolia(Okamura)I.K.Lee&Kurogi(�977),aspeciesrecognizedfromJapan,China,andKorea(Yoshida�998;Zhang&Xia�994;Kimetal.2006).Inthebeliefthatthenameofataxonomicsynonymwasnotavailable,Ricker(�987)proposedthenewnameRhodymeniasubantarctica,andthatnamehasbeenusedincontemporaryliterature(Wiencke&Clayton2002;Graeveetal.,2002).Thenameofasynonym,however,isavailable,namely,PhyllophoracoccocarpaMontagne(�852b),andthetransfertoRhodymeniaisproposedhere.Thisdisposition,whichwasfirstproposedbyKylinandSkottsberg(�9�9),hasbeenacceptedbyPapenfuss(�964),Ramírez&Santelices(�99�),andbyAlgaeBase(Guiry2006).

Rhodymenia coccocarpa(Mont.)M.J.Wynnecomb.nov.PhyllophoracoccocarpaMont.Ann.Sci.Nat.,Bot.sér.3,�8:3�8.�852.Membranifoliumcoccocarpum(Mont.)Kuntze,Rev.gen.pl.�89�:903.

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2007 WYNNE:RHODYMENIACOCCOCARPA 245

PhyllophoracuneifoliaHook.f.&Harvey,LondonJ.Bot.4:260.�845.Rhodyme-niacuneifolia(Hook.f.&Harvey)W.R.TaylorPapersMich.Acad.Sci.,Arts&Letters24:�47,�939,nonRhodymeniacuneifoliaOkamura,�934.Rho-dymeniasubantarcticaRicker,TaxonomyandBiogeographyofMacquarieIslandSeaweeds:2�5.�987.

InadditiontoMontagne’sillustration(�854,pl.�6,fig.�),Phyllophoracocco-carpahasbeendepictedbyCotton(�9�5,pl.7),Kylin&Skottsberg(�9�9,fig.�0,as“Rhodymeniacorallina”),Ricker(�887,fig.90,asRhodymeniasubantarctica),andWienckeandClayton(2002,pl.�6,figs4-5,asRhodymeniasubantarctica).

Thisspecieshasbeenreported(sometimesunderthenamesEpymeniamembra-naceaandRhodymeniacorallina)fromtheStraitofMagellanandTierradelFuego(Piccone�886;Hariot�889;Skottsberg�923),southernChile(Hariot�895;Kylin&Skottsberg�9�9;Taylor�939;Levrig�960),EasterIslandandJuanFenandezIsland(Levring�94�;Etcheverry�960), southernArgentina (Hariot �895), theFalklandIslands (Hooker & Harvey �845; Hooker �846–�847; Cotton �9�5;Taylor �939),Îles Kerguelen (Hooker �846–�847; Dickie �876; Askenasy �888; Zinova �973),MacquarieIsland(Ricker�987),andAntarctica(namely,KingGeorgeIslandandPalmerStationoftheAntarcticPeninsula)(Claytonetal.�997;Wiencke&Clayton2002).Taylor(�939)explainedthatwhileSkottsberg’s(�923)conceptofRhodyme-niacorallinaconformedtoPhyllophoracoccocarpa,Kylin’s(�93�:2�,pl.8,fig.20)conceptdidnot.

ACKNOWLEDGMENTS

IthankDr.BrunodeReviersoftheMuséumNationald’HistorieNaturelle,Paris,forlocatingtheholotypeofPhyllocarpacoccocarpaandsendingmeaCompactDiscwithscannedimagesfromthistype.Healsoprovidedahelpfulreviewofthemanuscript.IamgratefultoDavidBayforassemblingtheplatesfromtheimages.

Fig.4.Detailofdistalportionofholotype.

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246 CONTR.UNIVERSITYOFMICHIGANHERBARIUM VOLUME25

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