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A taxonomic revision of Indonesian Gelidiales (Rhodophyta) PDF

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BLUMEA 35 (1991) 347-380 A taxonomic revision of Indonesian Gelidiales (Rhodophyta) A.M. Hatta & W.F. Prud’homme+van+Reine Rijksherbarium/Hortus Botanicus,Leiden,TheNetherlands Summary Inthisstudy 12 taxaare treated,belongingtofourdifferent genera ofGelidiales (i.e. Gelidiella, Gelidium,Pterocladia, andPorphyroglossum) aswell asonespecies ofWurdemannia,agenus of unknown affinity. One new species,Gelidium amboniense,andanewforma,Gelidium latifolium forma elongatum,aredescribed. Holotypematerial ofGelidium bornetii Weber-van Bossein the Weber-van Bosse collection in Leiden canbesynonymizedwith Gelidiella lubrica (Kützing)Feld- mann& Hamel,while theoriginalsamplein the herbarium in Copenhagenonlycontains Gelidium pusillum (Stackhouse)Le Jolis varminusculumWeber-van Bosse. Inthekey tothe genera and species wealsoinclude the very similar genus CeratodictyonZanardini (=GelidiopsisSchmitz). Introduction Weber-van Bosse (1921) describedtwo groups withinthe genusGelidiumLa- mouroux(Gelidiales) forIndonesia, viz. Gelidiumwithoutand withinternalfila- ments.Thefirst group,ofwhichGelidiumpannosumBornetis arepresentative, has been separated fromthe genus Gelidiums.s. by FeldmannandHamel(1934) and hasbeen includedintheir new genus Gelidiella.Fan (1961) proposed anewfamily (Gelidiellaceae) forthis genus.ThesecondofWeber-vanBosse's groupsconsisted ofspecies nowplaced inthegeneraGelidiumandPterocladiaJ.Agardh. Inthe diagnosis ofthemonogeneric family Gelidiellaceae, Fan (1961) statedthat this family is lacking internalfilamentsas wellas asexual phase. However, recent investigations showthat internalrhizines as well as sexual reproduction mayoccur in this genus. Maggs and Guiry(1987) haveobserved internalrhizines in theirnew species Gelidiellacalcicola, although only in a limitedarea aroundtheattachment points. Acystocarpic plantofanunnamedGelidiellaspecies has beenfoundinIndia by RaoandTrivedi (1980). Todecidewhether specimens belong toPterocladiaor toGelidiumis usually not veryeasy. Many attempts havebeen madeto findcharacters separating thesegen- era. Thecharacters suggested include thelocationofinternalrhizines (Feldmann & Hamel, 1936; Loomis, 1949; Taylor, 1960),thestructureofthecystocarp (Feldmann 1) Recentaddress: Indonesian InstituteofSciences,CentreforOceanologicalResearch and Develop- ment,P.O.Box 44,Ambon, Indonesia. 348 BLUMEA VOL. 35,No. 2, 1991 Kam- Taka 13. Nusa Island; 4. Bay; Damar Jakarta 12. 3. Timor; Sumatra); 11. Mid Flores; of 10. coast Komodo; Islands. (S Arau 9. Kei Muara Sumba; 17. Ambon; 2. 8. Nias; Sumbawa; 16. 1. Strait; text. 7. the Galewo in Lombok; included 15. 6. Island; Madura; names Daram 5. geographicalJava); 14. Islands); Mid of of coast (Tiger Location (S Rate 1. bangan Bone Fig. A.M.Hatta& W.F. Prud'homme vanReinc: Revision ofIndonesian Gelidiales 349 & Hamel, 1936;Fan, 1961;Santelices, 1977),the apical structure (Rodriguez & Santelices, 1987, 1988), and the morphology and dimensionsofperipheral cells (Akatsuka, 1981, 1982) and of medullary cells (Loomis, 1949; Fan, 1961). How- ever, theposition ofinternalrhizines, the apicalstructure, andthemorphology and dimensionofcellscan only beusedtoseparatespecies andare not suitabletodefine genera.Differences inthe structureofthecystocarp (usually unilocularoroccasion- ally unequally bilocular, withonly onesurface provided withone ormoreostioles in Pterocladia, andalways bilocular, with oneormoreostiolesineach surfaceinGeli- dium)have generally been accepted as differentiating characters. New generade- scribed recently (PterocladiastrumAkatsuka 1986a,andOnikusa Akatsuka, 1986b) sharethecystocarp charactersofPterocladiaandGelidiumrespectively. LiteratureaboutthetaxonomyofIndonesianseaweeds isscanty andmostpapers aboutthese seaweeds discuss theireconomic value(Eisses, 1953; Zaneveld, 1955, 1959;Soegiarto, 1978). Intheyears 1984-1985the Indonesian-DutchSnellius-II Expedition to Indonesianwaters took placeandinthemonthsAugust andSeptember 1984manyseaweedshave been sampled. Thepurposeofthis study is to revise the informationabout IndonesianGelidialesas based on papers by Weber-van Bosse (1926,1928) and to incorporate all availableadditionalmaterial. One other genus showing morphological resemblanceto theGelidialeswill alsobe treated:Wurde- manniaHarvey (Wurdemanniaceae, afamilyofunknown affinity, possibly belong- ing to theorderGigartinales),while in thekey wealso include themuchresembling genusCeratodictyon Zanardini(probably Gracilariaceae, Gigartinales). MATERIAL AND METHODS Specimens coming fromthefollowing sources havebeenstudied: — specimens collectedduringtheDutchSiboga Expedition (1899-1900) and other specimens studiedby Mrs. Weber-vanBosse. Itis apity, however, thatmanyof the specimens studiedby Weber-vanBosse, apartfrom thoseofthe Siboga col- lection, are not preserved in theRijksherbarium or in theHerbariumBogoriense (BO)and seemtohave beenlost. — specimens collectedinIndonesiabefore1960andpreserved intheRijksherbarium, theNationalDutchHerbariumin Leiden. — specimens collectedduring theIndonesian-DutchSnellius-IIExpedition (Aug.- Sept. 1984). Ofthis Snellius-IIExpedition (SN-II) theA-seriesand theC-series (duplicates) arenowin theRijksherbarium, Leiden(L), theB- andD-series arein theCentreforOceanological Researchand Development - IndonesianInstituteof Sciences, in Jakarta(JAK) andinAmbon(AMB) respectively andtheE-series is in theBotanicalInstituteoftheUniversity ofGent,Belgium (GENT). For herbariarecordedintheIndexHerbariorumweuse thestandardizedabbrevia- tions. Ofall specimens theirexternal morphology as welas theiranatomyhasbeenstud- ied to name these specimens according tomodern classificationand nomenclature. Sectionsofherbarium specimens have been madeeither by hand(razor blades) or 350 BLUMEA VOL. 35, No. 2, 1991 by freezing microtome.Anilineblue,in aqueousdye, is usedto colourthesections, which are then mountedin an 80% solutionofKaro or in Aquamount. A camera lucidais usedinall anatomicaldrawings. All descriptions arebasedonIndonesianspecimens only,except whencharacters arerecordedfromtype specimens originating from othercountries.When in these descriptions layers of cellsare mentioned, thenumbersofcellscounted on straight linesfromthalluscentre tothallussurface are recorded. KEY TO THE GENERA AND SPECIES OF INDONESIAN GELIDIALES AND TAXA THAT RESEMBLE THEM la. Plants withoutinternalrhizines (no sexualreproduction known) 2 b. Plantswithinternalrhizines(sexual reproduction observedinsome species) 6 2a. Distinct apical cell present in all thalli Gelidiella (3) b. Thalliwithoutdistinctapicalcell 14 3a. Thallususually more than 25mm high,rigid, proximalpartsoferectaxes cy- lindrical, distalparts compressed, straight or curvedabaxially. Apical cell dist- inctly separated from the other cells 1. Gelidiella acerosa b. Thallusless than25mmhigh,cartilaginous, cylindricalorsomewhatcompressed throughout, unbranchedor branching (irregularly) pinnate. Apical cellnot dis- tinctly separated fromtheothercells 4 4a. Thallususually more than 10mm high, branching bipinnateor tripinnate 3. Gelidiella myrioclada b. Thallususually less than 10 mmhigh, theerectpart unbranchedor very little branched 5 5a. Only oneortwo layers ofinnercorticalcellsaroundonetierofmedullacells 4. Gelidiella pannosa b. Morethan twolayers ofinnercorticalcells andmedullacells not inonly one tier 2. Gelidiella lubrica 6a. Matureplants usuallyless than 1cm highandoftencrowdedtogether forming mats 7 b. Matureplants usually morethan 1 cm high 11 7a. Only onelayer ofinnercorticalcellsaroundonetierofmedullacells 5. Gelidiumamboniense b. Always more thanonelayerofinnercorticalcellspresent 8 8a. Erect thallusparts mostly unbranched;filiformorspatulate throughout 8. Gelidium pusillum var. minusculum b. Thalluswithbranched erectparts 9 9a. Erectpartscompressed throughout 10 b. Erectparts cylindrical or distallybecoming compressed. Branching irregular, oftentrifidand not in oneplane 9. Gelidium pusillum var. cylindricum 10a. Branchingoferectparts irregularor scarce. Always sterile.Juvenilesof 7. Gelidium pusillum var. pusillum b. Branchingof erectpartspectinate. Cystocarps unilocular, sometimesarranged in series 12. Pterocladia caloglossoides A.M.Hatta& W.F.Prud'homme vanRcinc: Revision of Indonesian Gelidiales 351 11a. Smallramulipresent, arranged in longitudinal rows onthe surfaceofthe flat- tened erectpartsofthe thallus 10. Porphyroglossum zollingeri b. Small ramuliabsentor not arranged inlongitudinal rows 12 12a. Erectparts withscarce orirregularbranching, proximal erectparts constricted, distallyflattenedthroughout (upto 1000|im inwidth) 7. Gclidium pusillum var. pusillum b. Erectparts pinnate orwithopposite branches 13 13a. Branchingoferectthallusopposite, oftenpinnate withpyramidal outline, rigid; apical cell sunken or exposed . . 6. Gelidiuntlatifolium forma elongatum b. Branching oferectthallus pinnate, notpyramidal inoutline, cartilaginous; api- calcell exposed 11. Pterocladia caerulescens 14a. Tetrasporangia zonately divided.Corticalcells and medullacellsin cross sec- tion are almost isodiametric 13. Wurdemannia miniata b. Tetrasporangia cruciatelvor tetrahedrally divided.Innermostcorticalcellslarger thancells ofouter cortexand thanmedullacells Ceratodictyon A detailedstudy ofIndonesianCeratodictyon willbepublished separately. GELIDIELLA EchinocaulonKiitzing (1843)405,nom. illeg., nonEchinocaulon Spach (1841)521 (a genus of floweringplants); Gelidiella Feldmann & Hamcl (1934)529,nom. nov. — Type: Echino- caulonspinellumKutzing(1843)406 =Gelidiella acerosa(ForsskAl) Feldmann & Hamcl. AcrocarpusKutzing (1843) 405,nom.illeg., non Wight(1839) 198 (a genusoffloweringplants). —Lectotype:Acrocarpus lubricus Kutzing = Gelidiellalubrica (Kutzing)Feldmann& Hamel. For description and discussion seeFan (1961: 340) and Maggs & Guiry(1987: 431-433). For Indonesianspecies see alsoTable 1. 1. Gelidiella acerosa (Forsskål) Feldmann& Hamel - Fig. 2 Fucus acerosusForsskdl (1775) 190.—Echinocaulon acerosumBprgesen (1932)5. —Gelidiella acerosaFeldmann & Hamcl (1934)533; Dawson (1954)422; Rao (1970)64, t. 1; Santelices (1977)63; Lawson& John (1982) 172,t.21,f.2.—Type: Mokka, Yemen (C, not seen). Fucus rigidus Vahl (1802)46, nom.illeg., non Turra (1780) 68.—Sphaerococcus rigidus C. Agardh (1822)285,nom. nov. —GelidiumrigidumGrevillc (1830)LVII.—Echinocaulon rigidum Kutzing(1868) 14,t.40,f. a-d.—GelidiopsisrigidaWeber-van Bosse (1904) 104; (1928)42.—Type: St.Croix,VirginIslands (notseen). Fucus spinaeformis Lamouroux (1805)77, t. 36,f. 3 & 4.—Gelidium spiniforme Lamouroux (1813) 129.—Type: Mauritius (notseen). Echinocaulon spinellum Kutzing (1843) 406, (1868) 14, t. 38,f. d& e. — Typ e: Mariancn, Gaudichaud inherb. Kunth (L941,27-215). Echinocaulon ramelliferumKQtzing(1868) 14,t.39,f. d-f.—Typ e: NewCaledonia,Vieillard 2061(L941,11-63;not L941,11-9,which isGracilia spec.). Gelidiopsisacerosavar. semipinnataWcbcr-van Bossc (1928)428 (Indonesianmaterial only, sec note 1).—Notypeselected. Gelidiellaacerosaformaminima Rao(1970)67, t. 1,f. c.—Ty p e: Adatra,Okha,India, in Cen- tral Saltand Marine Chemicals Institute,Bhaunagar,India (CSMRI- 4223 - D,notseen).(See note2). 352 BLUMEA VOL. 35, No. 2, 1991 Fig. 2.Gelidiella acerosa. a. Habit (arrow:slichidia);b. surfaceview ofthe apex ofanerect thallus; c. stichidium,densely covered with tetrasporangia; d. crosssection ofastichidium with tetraspo- rangia;e.surface view ofcortical cells; f&g.crosssections oferect thallus parts,f.with external andinner corticalcells,g.with small medulla cells surrounded by largeinnermost cortical cells (a, c-eKomodo Is., SN-II, 10850;bGalewo Strait,L940,355-78;f, gKomodo Is., SN-II, 10812). Mature plants tufted, 0.5-9 cm high; consisting ofcylindrical prostrate axes (diam. 100-320pm),attached tothe substratumby disk-like haptera issued atirreg- ular intervals and with erect,rigid, branched or unbranched, straight or abaxially curvedthallusparts, proximally cylindrical (diam. 100-400pm) and distally com- pressed (450-600 x 300-350pmin cross section) (fig. 2a). Branching pectinate orpinnate, opposite, alternate,orsecund; smallterminalramulifiliformorspatulate (fig. 2a). Apexofaxes andbranches conicalorattenuate;apical cell separated from theothercells butnot very conspicuous (fig. 2b). Externalcortical cellsin surface viewdiagonally arranged, thick-walledandrectangular incross section, 2-5 x4-8 pm,anticlinally elongate (fig. 2e, f);inner cortical cells in cross section rounded, diameterinwards grading from 3 to 26 pm (fig. 2f, g). Medullacells roundedin cross sectionand smallerthan innermostcorticalcells, diameter15-20pm(fig. 2g), A.M. Hatla & W.F.Prud'homme vanReine: Revision ofIndonesian Gelidiales 353 longitudinally elongate. Tetrasporangia inswollenconicalstichidialocatedin theter- minalramuli(fig. 2a, arrow); stichidia 150-180x 300-350pm and densely or sparsely, regularly or irregularly covered with tetrasporangia (fig. 2c). Proximal tetrasporangia usually in a moreadvanced stage than distalones, ovate insurface view(c. 20pmin width), oblong incross sectionofthe stichidia(30-42 x23-28 pm) (fig. 2d). Distribution. World: in warm temperate and tropical seas all over the world.Indonesia:commonin allIndonesianseas. Discussion. This species can be distinguished easily fromotherGelidiales by itsexternal morphology. Itoccurs inthelittoralzoneon wave-exposed shoresas well as in tidalpools, in the shallow sublittoraland even in water of30 m depth (Komodo Island, SN-II, 10980). The41 Indonesiansamples did not show morpho- logical discontinuities; variationin dimensionsis the only striking element.These dimensionsrangein our samples from8.5 mm high (Timor, L 941,27-253) up to 9 cm high plants (GalewoStrait, IrianJaya, L940,355-78). Notes. 1. Specimens recorded by Weber-van Bosse (1928) as belonging to var. semipinnata (Piccone & Grunow) Weber-vanBosse fallin therangeofvariation inmorphology and dimensionsofthis species andthereforecannot betreatedas a separateinfra-specific taxon. Isotype materialofGelidiumsemipinnatum Piccone & Grunowin Piccone(1884: 315) in L belongs to Gelidiellalubrica (Kiitzing) Feld- mann& Hamel. 2. Thedimensionsoftheplants ofGelidiellaacerosa formaminimaas describ- edbyRao(1970) are withintherangeofvariationofsamples fromIndonesia.There is noreason to considerthese specimens as belonging to a separate infra-specific taxon. 2. Gelidiella lubrica (Kützing) Feldmann & Hamel - Fig. 3 Acrocarpuslubricus Kiitzing (1843)405, t.60 II; (1849)761; (1868) t.32.—Gelidiellalubrica Feldmann & Hamel (1934)535, f. 3-5;Womersley & Bailey (1970)305.—Ty p e: Naples, Italy (L941,46-46and941,46-347). Gelidium semipinnatumPiccone &Grunow inPiccone(1884)315. —Gelidiopsisrigidavarsemi- pinnataWeber-van Bosse (1928)428.—Isotype: Assab Bay, Eritrea,Ethiopia, leg. A. Issel,onPatella (L941,46-291,seenote 1toG.acerosa). Gelidium bornetiiWeber-van Bosse (1926)107.— Gelidiellabornetii Feldmann & Hamel (1934) 535; Bprgesen (1938)210,f. 2; Dawson (1957a) 113, f. 21; Cribb(1983) 30, t. 7, f. 1.— L ectoty p e:Nusa Kambangan,Java,Indonesia,leg. Jensen (L941,27-296). Small (3-5 mm high), mat forming, saxicolous plants; consisting ofcylindri- calor somewhat compressed prostrate axes ( diam.90-127 pm), attached to the substratumby peg-like haptera issued atirregular intervals andopposite to theun- branchedor scarcely andirregularly branched, cartilaginous, filiformor somewhat compressed, erect thallusparts, diameter60-80pm(fig. 3a, b). Apex oferectpart attenuate;exposed apical cellnot separated fromtheothercells and quiteconspicu- ous. External corticalcells angularin surface view, diameter4-7pm,longitudinally andtransversely arranged in youngerparts, more irregular inolderparts (f. 3d). In cross section externalcortical cells quadrangular, diameter4-8 pm; innercortical 354 BLUMEA VOL. 35, No. 2, 1991 Fig. 3.Gelidiella lubrica.a&b. Details ofplants; c.stichidium;d. surface view ofexternal cortical cells;e& f. cross sections oferect parts,e. maturepart,f.youngpart(a,b, d,eNoimini,Timor,L 941,27-295;cTakaBoneRate,SN-II,11224-,flectotypeofGelidiella bornetii fromNusaKamban- gan, S coast Mid Java,L941, 27-296). cells and medullacells rounded, diameteroften inwardsgrading from7to 12pm (fig. 3e, f).Medullacells areelongate in longitudinal direction, diameteras ininter- nal corticalcells or smaller. Tetrasporangia in terminalstichidia whichare slightly swollenor not,cylindricalor lanceolateandsomewhatcompressed (250-650 x98- 115 pm), usually densely covered withtetrasporangia (mostly cruciate and often tetrahedral) which are arranged in transverse rows or inchevronsor are irregularly scattered(fig. 3c). Proximal tetrasporangia usually more advancedthan distalones, ovate in surface view(19-32 x 10-19pm). Distribution. World: in warm temperateandtropical seas: Mediterranean, India, Indonesia, MarshallIslands, SolomonIslands, GreatBarrierReef.Indonesia: NusaKambangan (S coast ofMidJava),NoiminiBay (Timor) andTakaBoneRate (Tiger Islands). Discussion. These small mat-forming algae are not easily separated from GelidiellapannosaandsmallGelidiumspecies. InGelidiellapannosathereis always only one tierofmedullacells andthetetrasporangia areoftenregularly arranged in chevrons ortransverse rows. In smallGelidium species thecorticalcells in surface A.M. Hatia & W.F. Prud'hommc vanRcine: Revision of Indonesian Gelidiales 355 view are smaller, moreroundedand irregularly arranged, in cross section internal rhizines canbe observed andthe tetrasporangia are arranged in irregular groups in thestichidium.The tetrasporangia inGelidiellalubricacanbearranged with4-6 in transverse rows orin chevrons, but in otherspecimens, and even inother stichidia onthe sameplant,arrangementoftetrasporangia can be irregular. Thisirregular ar- rangementoftetrasporangia hasbeen usedby Bprgesen (1938) andCribb (1983) to separateGelidiellabornetiifromG.tenuissima(= G.pannosa). However,lectotype materialofGelidiellabornetii,labelledby Weber-vanBosse, doesnot differenough fromG. lubricato be considereda separate species. Moreover,the description by Weber-vanBosse (1926) differs fromthelectotype in dimensions, form,andarran- gementofthecortical cells. Two microslides in L, labelled‘Gelidiumbornetii’by Weber-vanBosse, also conformto the description ofIndonesianspecimens ofG. lubricaandagain are not exactly inaccordance withher description ofGelidiella(as Gelidium)bornetii.Erectparts ofG. lubricacanbe filiform, cylindrical or somewhat compressed and thisagain is not areliablecharactertoseparatespecies. A sample in C,labelled'Gelidiumbornetiin. sp., NusaKambangan' consists ofapiece ofcoral on whichonly specimens ofGelidiumpusillumvar. minusculumcouldbe detected by us. Thedescription by Cribb(1983: 3)ofGelidiella species is in agreementwith our conceptofG. lubrica. Thetype material ofG. lubrica is larger(up to 3cm high) and moreprofusely branched than the Indonesianspecimens. However, the irregulararrangementof tetrasporangia as wellas theirarrangementin transverserows occurs in these speci- mensandtheanatomyofthalliis not differentfromthatofIndonesianspecimens. Note.Kutzing distributedin 1836driedspecimens collectedby himduring his journey toItaly. In thismaterial, called'Actien',heusedthe nomennudumSphaero- coccus lubricusforspecimens nowtobenamedGelidiellalubrica. 3. Gelidiellamyrioclada (Børgesen) Feldmann& Hamel - Fig. 4 Echinocaulon myriocladumB0rgescn (1934)5, f.4& 5. — GelidiellamyriocladaFeldmann & Hamel (1934) 533; Bprgcscn (1935)44; Dawson (1954)422, f. 33d; Santelices (1977)66, f. IF-H.—Type: Malabar Hill,Bombay,India,B0rgesen 5235(4slides in C, one, fig. 5a in B0rgescn 1934,has been selectedasthelectotype). Maturesaxicolousplants tufted,12-22mm high;consisting ofcylindrical pros- trateaxes (diam. 75-135 pm) attached tothe substratumby peg-like haptera (75— 450 pmin length) and with erect,branched,cartilaginous thallusparts, proximally cylindrical (diam. 80-140pm) and distally ovate throughout incross section (100- 180pm broad). Branching bipinnate or tripinnate,branchesproximally constricted; lateralterminalramuli 1-3.5mm long, filiformorspatulate (fig. 4a). Apex ofaxes and branches attenuate; exposed apical cell not separated from theother cells and quite obvious. External cortical cells angular in surface view, often withrounded edges and not arranged in distinct rows (fig. 4c), in cross sections quadrangular, compressed (3-7 x3-10pm); first layer ofinnercortex similarto theouter cortex, inwards followedby threelayers of ovate cells, 7-10pm wide (fig. 4d). Medulla cells in 4-5 layers, ovate in cross section, 10-14pm wide, longitudinally elon- gated. Tetrasporangia incompressed lanceolatestichidia, locatedinthelateralramuli; 356 BLUMEA VOL. 35. No.2, 1991 Fig. 4. Gelidiella myrioclada. a. Habit;b. stichidia with tetrasporangiaarranged in lateral rows; c. surface view ofexternal corticalcells; d. cross section oferect thallus part (a, c, dTaka Bone Rate,SN-II, 11262;b lectotype, Malabar,India). stichidia380-765x 125-155p.m, withclosely setroundedtetrasporangia,arranged in transverse rows (fig. 4b). Diameterofthe tetrasporangia 18-20|im in surface view(description ofthestichidiais takenfromthelectotype). Distribution.World: tropicalIndo-Pacific Oceans:Kenya, India, Vietnam, Indonesia, Hawaii. Indonesia:TakaBoneRate (Tiger Islands; SN-II, 11262). Discussion. Differences between the sterile specimens from Taka Bone Rate andotherIndo-Pacific specimens concern bipinnate subopposite branching and smallerspecimens in Vietnam(Dawson, 1954)and smallercellsin cross sectionin Hawaiianspecimens (Santelices, 1977).The structureofthe quitesimilarly branched GelidiellamachrisianaDawson (1957b: 17,f.4B) studiedin thetype(from Puerto Rico, Dawson 16745, in LAM) proved to differconsistently from G. myrioclada, especially as forthedimensionsandsituationsofthecells. 4. Gelidiellapannosa (Feldmann) Feldmann& Hamel- Fig. 5 Gelidium pannosum Bornet (1892)267, non G. pannosum Grunow, which is aCeratodictyon; Wcber-van Bossc (1921)233,f. 68 t. 7, f.3(p.p). —Echinocaulon(?) pannosum Feldmann (1931b) 12.—Gelidiellapannosa Feldmann & Hamel (1934)534,f. 1 &2; Cribb(1983)31,t. 6, f.2;Egcrod(1971) 127,f.29-31.—Gelidiellatenuissima Feldmann & Hamel (1936) 226,

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