Belg. Journ. Bot. 130 (2): 177-197 (1998)

THE SIPHONOCLADALES SENSU EGEROD (CHLOROPHYTA)FROM PAPUA NEW GUINEA AND INDONESIA

(SNELLIUS-II EXPEDITION)

F. LELIAERT, E. COPPEJANS & O. DE CLERCK

Vakgroep M orfologie, Systematiek en Ecologie ; Labor atorium Plantkunde, Universiteit Gent ,K.L. Ledeganckstraat 35, B-9000 Gent , Belgium

SUMM ARY. - Identification keys, descriptions and illustrations are provided for thegenera and species of A nadyo mene (3 spp.), Micro dictyon (3 spp.), Boergesenia (l sp.),Boodlea (2 spp.) , Struvea (1 sp.), Phyllodictyon (2 spp .), Dictyosphaeria (3 spp.), Valonia(4 spp.) and Ventricaria (1 sp.) based on the specimens collected in Papua New Guineaand Indonesia from 1980 to 1994. Four new records for the region were found: Microdictyonokamurae (collected in the Banda and Flores Sea , Indonesia), Microdictyon palmeri andPhyllodictyon gardineri (collected along the N coast of Papua New Guinea), andDictyosphaeria ocellata (collected along the S coast of Papua New Guinea).

RESUME. - Les Siphonocladales sensu Egerod (Chlorophyta) de Papouasie Nouvelle­Guinee et d'Indonesie (Expedition Snellius-II) - Des clefs d'identification, des descriptionset des illustrations sont donnees pour 1es especes appartenant aux genres Anadyomene (3spp.) , Microdictyon (3 spp.), Boergesenia (1 sp.), Boodlea (2 spp.), Struvea (1 sp.),Phyllodictyon (2 spp.), Dictyosphaeria (3 spp .), Valonia (4 spp.) et Ventricaria (1 sp.) , baseessur les specimens recoltes en Papouasie Nouvelle-Guinee et en Indonesie de 1980 a 1994.Quatre nouvelles especes sont mentionnees pour la region: Microdictyon okamurae (dans1es mers de Banda et de Flores, Indonesie), Microdictyon palmeri et Phyllodictyon gardineri(Ie long de la cote Nord de 1a Papouasie Nouvelle-Guinee) et Dictyosphaeria ocellata (Ielong de la cote Sud de la Papouasie Nouvelle-Guinee),

INTRODUCTION

Research on marine macroalgae in PapuaNew Guinea has been carried out by Eric Cop­pejans since 1980. Collecting was done along theNand S coast (mainly the Madang Province andthe Port Moresby region, respectively), result ingin detailed studies of some groups (COPPEJANS& MEINESZ 1988, COPPEJANS 1992) or in anno­tated check-lists (COPPEJANS et al. 1994, Cop-

PEJANS et al. 1995, MILLAR et al. 1998). Sipho­nocladales are mentioned in BROU NS & REUS1985, R EUS 1985, KING 1990, ENOMOTO & OHBA1992, OHBA & ENOMOTO 1992, COPPEJANS et al.1994.

Unlike Papua New Guinea, Indonesia hasa long history of macroalgal research. The dutchSiboga expedition at the end of the 19th centurywas an early milestone, and led to Weber-vanBosse's 'Liste des algues du Siboga', which is still

178 BELG IAN JOURNAL OF BO TANY 130

a useful guide tod ay (WEBER-VAN BOSSE 1913).During th e 'o cea nographic Snellius-Il ex peditio n'(1984) macroalgal collections were made in theBand a and Flores Sea (COPPEJANS &P RUO'HOMM E VAN REINE 1989a, 1989b , I992a,1992b). During the Burginesia-IIl pr oject (1988­1990) macroalgae were collected in th e Sp er­monde Archipelago, SW Sulawes i (VERHEIJ &P RUO' HOMME VAN REI NE 1993).

. The Siphonocladales-C ladophorales com­plex has recently been the su bjec t of severalstudies co nsidering the sta tus of both orders. T hegenera dealt with in thi s study all have a sipho­nocladous morphology (e.g. inflated cells, net- orbladelike structure) and were tradition ally placedin the Siphonocladales. The Sipho no cladales werebelieved to have been derived fro m certain mem­bers of the Cladophorales, which have a m oresimple morphology. M ore recently, there hasbeen disagreement on th e status of th e ordersSiphono clad ales and Cladophorales. From ma­cro-morphological, ult rastructural and mo lecul arevidence it appears th at neither th e Siphonocla­dale s nor the Cladophorales are mon ophyleticand th at there is no reason for separa ting the twoord ers (VAN DEN HOEK 1982, OLSEN-STOJKOVICH1985, 1986, BAKKERet al. 1994).

Disagreement abo ut th e delimitat ion s of th efamilies has resulted in a diversi ty of classificationsin the past (BORGES EN 1940, 1946, EGEROD 1952,TAYLOR 1960, WOMERSLEY & BAI LEY 1970,WOMERSLEY 1984, LEW IS 1987, LEWIS & NORRIS1987, S ILVA et at. 1987, 1996, SARTONI 1992,SOUTH & KASAHARA 1992).

The aim of this study is an inventory anddescription of th e species belonging to the Sipho­nocladales sensu Egero d, resulting in an identi­fication key fo r th e regio n.

MATER IAL AN D M ETHODS

Field work was carried out in Mada ng Province(N coast of Papua New Guinea) from J une to August1980, in Jul y-Augu st 1986, fro m June to August 1988,in July-August 1990 (map with collecting sites inCOI'PEJANS 1992). In the Port Mo resby area seaweedswere collected in Ju ly 1986 and in July-August 1994(map with collecting sites in COPPEJANS et al. 1995).

During the Snellius-Il expedition collections of

macroalgae were made in September 1984 in the Band aand Flores Sea, Ind onesia (map with collecting sitesin COPPEJANS & PRUD'HOMMEVA N R EI NE 1992).

Sampling was done by wad ing in the intertid alzone at low tide and by snor keling and SC UBA-d ivingin the subtida l areas. Mos t collected specimens weredried and pressed as herb arium reference material.Some are preserved in seawater/formalin solution.

The species descriptions are based on persona lobservations ; drawings were made by camer a lucida .All specimens deposited in the Herbarium Universita tisGa ndave nsis (GEND were examined (Co pp. & PvR .= Co ppejans & Prud'homme van Reine ; HEC =Herb arium Eric Co ppejans; ODC = Olivier DeClerck ; Sn-I l = Snellius-Il expedition ; UNS W =University of New South Wales). Macromorphologicaland microscopic cha racters were exa mined in bothdried specimens and rehydrated herbarium frag ments(solutio n of 4% formalin and 5% glycerin in seawater) .In genera where cell dimensions are important foridentification at species level, 20 cells of each specimenwere measured. Major publ icat ions used for speciesidentification are mentioned under the species' name,after the type locality. Ecological data were ob tainedfrom the herbarium labels. Geographical d istribut ionis based on checklists for the region (Ind ian Ocean :SILVA et at. 1996 ; Taiwan: LEWIS & NORRI S 1987 ;Philippines: SILVA et at. 1987 ; Indonesia & Philip­pines: TAYLOR 1966 ; Micronesia : TSUDA 1977, 1981 ;So lomon Isles : WOMERSLEY & BAI LEY 1970 ; N­Australia : LEWIS 1987 ; Fiji: SOUTH & KASAHARA1992 and SOUTH et at. 1993 ; N Pa pua New Guinea :ENOMOTO & OHBA 1992). For supplementary data ongeographical distri bution see SILVA et al. (1996).

RESULT S

IDENTIFICATION KEY TO THE GENERA OF THESIPHONOCLAOALES FROM PAPUA NEW G UINEA

AND INDONESIA

I. a. Tha llus com posed of one or mo re blades, cellsof the blade essentially branching in one plane ,contiguo us or net-like . . 2

b. Thallus cushion-like, composed of a network ofintert wined cells or co mposed of one or morelarge, inflated cells or pseudoparenchymatous ...5

2. a. Blades fanshaped (flabellate), meshes of the bladefilled with small interstitial cells ; margin of thelamina consisting of small spherical cells . .

................................ Anadyomeneb. Blades netlike, meshes of the blade not filled with

interstitia l cells . . 3

SIPHONOCLADALES FROM PAPUA NEW GUINEA AND INDONESIA 179

3. a. Blades not stipitate, shape indefinite , margin ofthe blade consisting of free branching cells ;anastomosis by smooth annulate or crenulatewall thickenings at the tip of terminal cells

.................. ..Microdictyonb. Blades stipita te, elliptical or triangular ; anasto-

mosis by tenacula . . 44. a. Stipe conspicuous, with basal annular constric-

tion s, blades triangular to elliptical Stru veab. Stipe without basal annular constrictions, blades

elliptical to irregular . . Phyllodictyon5. a. Thallus composed of a network of branching

cells in three dimensions ; anastomosis by tena­cula . . Boodlea

b. Thallus composed of one or more inflated,macroscopically visible cells or pseudoparenchy­m~0~6

6. a. Thallus pseudoparenchymatous ; solid and but­ton-shaped, or hollow and spherical or cup-shaped . . Dictyosphaeria

b. Thallus not pseudoparenchymatous, composedof one or more inflated cells 7

7. a. Thallus composed of compressed or looselyinterwoven branching cells, forming hemisphe-rical domes or irregular cushions Valonia

b. Thallus composed of one or more unbranchedaseptate cells 8

8. a. Cells large, clavate , frequently curved , with basalannular constrictions, isolated or clustered; pa le-green Boergesenia

b. Thallus a single cell without basal annularconstrictions, never branched, extremely tough ;dark blue-green . . Ventricaria

TAXONOMIC TREATM ENT

Anadyomene Lamouroux

Unistratose fans haped blade or cluster ofblades formed by polychotomous branching ofuniseriate veins (vein segments between polycho­tomies consisting of one to several vein cells),interstitial spaces completely or partially filledwith smaller (interstitial) cells ; attachment to thesubstrate by rhizoids, which may intertwine toform a stipe (LITTLER & LITTLER 1991: 101).

Identification key

I. a. Interstitial cells parallel , partially overlapping thevein cells . . A. stellata

b. Interstitial cells randomly placed, partially orcompletely overlapping the vein cells 2

2. a. Vein cells of the whole thallus completely coveredby overlapping interstitial cells A . brownii

b. Interstitial cells only overlapping the vein cellsat the base of the thallus . . A . plicata

Anadyomene brownii (1. E. Gray) J . Agardh(1887 : 127) (Fig . 6)

WEBER-VAN BOSSE (1913 : 75).Calomena brownii J. E. Gray (1866: 46), [type

locality : Australia]

Thallus composed of undulate or plicateblades. Fronds eperforate, with lobed margins,1-4.5 em tall, anchored at the base by looselyentangled rhizoids forming a short stipe . Veinsbranching polychotomously (4-6 branches) at theapices of segments , but lateral- oranches alsopresent. Vein segments composed of 4-6 cells inthe central and marginal parts of the blade, of1-7 cells in the basal part. Vein cells subeylindrical,in the central part of the blade up to 5.5 mmlong, (2) 75-225 urn. Interstitial spaces filled withsmall , randomly arranged, polygonal cells, over­lapping the vein cells completely in the entireblade , except for the marginal part. Blade marginconsisting of small spherical cells, (2) 20-60 urn.

Ecology: infralittoral fringe and infralittoral,epipsammic or on coral rubble.

Distribution : Philippines, Indonesia , nor­thern Australia, Solomon Islands

Specimens examined: Indonesia, Banda Sea,Maisel Islands : Sn-ll 10111 C: 07.ix.1984. NEcoast of Sumba, E of Melolo : Sn-II 10516 E :13.ix.1984. SW Salayer: Sn-ll 11520 E:lO.x.1984. E of Komodo Island, Selat Linta : Sn­II 10838: 18.ix.1984. SW Salayer: Sn-Il 11559C, E : lO.x.1984.

Anadyomene plicata C. Agardh (1823 : 400)(Figs . 1-3)

[type locality : "Ravak" [Rauki], Island Wai­geo, Moluccas, Indonesia], WEBER-VA N BOSSE(1913 : 75, figs. 16, 17), SARTONI (1992 : 292,figs. 2, 3)

Thallus composed of rosulate, dark greenblades. Fronds stiff, eperforate, flat when young,undulate or plicate when mature, up to 7 em tall ,anchored at the base by loosely entangled rhizoidsforming a short stipe. Veins macroscopicallyvisible, vein segments composed of 1 or 2 cells,

180 BELGIAN JOURNAL OF BOTANY 130

branching polychotomously (4-7 (-8) branches) atsegment apices; angle of polychotomies 10°-35°.Vein cells subcylindrical, (1-) 3.8 (-5.5) mm long,o 75-150 (-190) urn, L jW up to 22. Lateralbranches randomly distributed along the veinsegments. Interstitial spaces filled with small,randomly arranged, polygonal or elongated cells(interstitial cells), partly overlapping the vein cells.Vein segments in the basal part of the bladecovered by rhizoidal outgrowths of the interstitialcells. Blade margin consisting of small sphericalcells (0 38-75 urn),

Ecology : infralittoral fringe or infralittoral,epi1ithic on coral boulders.

Distribution: Somalia, Seychelles, Malaysia,Indonesia, northern Australia, Papua New Gui­nea, Vietnam, Philippines.

Specimens examined : Papua New Guinea,Madang Province, Laing Island: HEC 4225:26.v.1980, HEC 4323 : 09.vi.1980, HEC 4477 :14.vii.1980, HEC 4737: 28.viii.1980, HEC 6471 :15.viii.l986. Madang lagoon area: HEC 7513 :20.vi.l988. Boisa Island : HEC 7713: 07.vii.l988.Cape Iris, Beliau, Astrolabe Bay: HEC 7994 :26.vii.1988. Murukinam: HEC 8082: 15.vi.1988.Demasa Island: UNSW 20919: iv.l988. reefbetween Sinub Island and Wongat Island: Copp.& PvR. 13240 B : 18.vii.1990. Ruo Island: Copp.& PvR. 13259 B : 19.vii.l990. Neptunus Point:Copp. & PvR. 13298 B: 21.vii.1990. Bagabag,Christmas Bay: Copp. & PvR. 13529 B :21.vii.1990. Boisa Island , Hansa Bay: Copp. &PvR. 13780 B : 20.viii.l990. Port Moresby area,Motupore Island: HEC 6312 : 05.vii.l986. PortMoresby area, Lion Island: HEC 10215 :22.vii.1994.

Remark: the specimens of A. brownii fromIndonesia and of A . plicata from Papua NewGuinea only differ by the fact that the blades ofA. plicata are not completely corticated. The levelof cortication however could be an ecologicaladaptation to less sheltered conditions (increasingcortication with increasing level of exposure).

Anadyomene stellata (Wulfen) C. Agardh (1823 :400) (Figs. 4, 5)

WEBER-VAN BOSSE (1913: 74), TAYLOR(1960: 125), LITTLER & LITTLER (1991 : 112-114).

Viva stellata Wulfen in Jacquin (1786 : 351) [typelocality : Adriatic Sea].

Anadyomene jlabellata Lamouroux (1816 : 366),[type locality: "Dans la Mousse de Corsedes pharmaciens" (Alsidium helminthochor­tos)].

Anadyomene cutleriae J. E. Gray (1866 : 48),[type locality : Bermuda].

Anadyomene stellata var. floridana J. E. Gray(1866: 48), [type locality: Key West, Flo­rida].

Anadyomene stellata var. luxurians De Toni(1889: 369), [type locality not specified]

Thallus composed of rosulate blades . Frondsstiff, undulate, eperforate, with lobed margins,1 em tall. Veins branching polychotomously (4­7 branches at segment apices), with interstitiallateral cells. Vein segments composed of 1-7 cells;vein cells subcylindrical to clavate, up to 1.5 mmlong , (75-135 urn, L jW : 7-14. Interstitial spacesfilled with small, cylindrical cells (0 20-130 urn),parallel, partly overlapping the vein cells. Blademargin consisting of small spherical cells (40­75 urn).

Ecology: no data on the herbarium label.Distribution: Atlantic Ocean; Indian

Ocean: Tanzania, Kenya, Madagascar, India, SriLanka, Andaman Islands, Indonesia; PacificOcean: Philippines. (Pan-(sub)tropical).

Specimens examined: Indonesia, Banda Sea,Tukang Besi Islands, Kaledupa reef: Sn-II 10254E : 09.ix.l984.

The synonymy of A. jlabellata was proposedby AGARDH (1823 : 400), the synonymy of A.cutleriae and the two varieties of A. stellata, var.floridana and var. luxurians was proposed byLITTLER & LITTLER (1991 : 113, 114).

Microdictyon Decaisne

Non stipitate, net-like blades formed bypolychotomous, opposite or alternate branchingof cells in one plane (primary cells form the mainveins and are distinctly larger than the secundarycells which form the 'branchlets'); anastomosisby smooth annulate or crenuiate wail thickeningsat the tip of terminal cells.

Identification key

I. a. Anastomosis by means of crenulate or hair-like

6$Cf?'~~x

F IGs. 1-3. - A nadyo mene p /icata C. Agardh . I : Main bra nching pattern of blade (interstitial cells have been omitt ed) ;(Copp. & PvR . 13259) (2 mm). 2 : Detail of blade with vein and interstitial cells ; (HEC 7713) (500 urn). 3 : Cros s sectionthrough centr al part of the blade, large vein cells and small interstiti al cells ; (HEC 4225) (200 urn).

FIGs. 4, 5. - Anadyomene stellata (Wulfen) C. Agardh ; (Sn-II 10254). 4 : Detail of central part of de blad e, veinand interstitial cells ; (200 urn), 5 : Detail of basal part of the blade, vein cells part ially covered by rhizoidal outgrowthsof interstitial cells ; (200 urn).

FI G. 6. - A nadyomene brownii (1. E. Gray) J. Agardh : Cross section through centra l part of the blade, large veincells covered completely by small interstitial cells ; (5n-II 11559) (200 urn),

182 BELGIAN J O URNA L OF BOTA N Y 130

outgrowths at the distal part s of apical cells ;colour dark green M. okamurae

b. Anastomosis by means of smoo th, annulateoutgrowths at the distal part s of apical cells ;colour pale green .. . 2

2. a. Branching opposite or stellate, all meshes ofsimilar size M . palmeri

b. Branching alternate to unilateral, meshes of twodistinct sizes (large and small) .... M. vanbosseae

Microdictyon okarnurae Setchell (1925 : 107)(Figs. 7, 8)

[type locality : Ryukyu, Japan], Setchell(1929 : 553), Taylor (1950 : 41).

Thallus composed of netlike blades, attachedby aseptate rhizoids ; blades dark green, indefinitein shape, up to 15 em tall. Branching oppositeor flabellate ; cells subcylindrical, 150-675 urnlong, 0 150-300 urn, L/W : 1-4 ; apical cells freeor anasto mosing, 150-490 urn long, 0 75-190 urn,Anastomosis accomplished by a crenulate thicke­ned membrane at the tip of an apical cell. Meshesdelimited by anastomosing cells or branch estriangular to pentangular, largest diameter 150­2200 urn , Blades not adhering firmly to paperwhen dried.

Ecology: infralitt oral (-1/ -30 m), epilithic onreefs or epipsammic in mangroves and seagrassmeadows. No morphological differences wereobserved between specimens growing at differentdepths.

Distribution : Seychelles, Alda bra Islands,Taiwan, Philippines, Indonesia, North Australia,Micronesia.

Specimens examined : Indonesia, Banda Sea,Maisel Islands : Sn-II 10153E : 07.ix.l984. BandaSea, Tukang Besi Islands, Kaledupa reef: Sn-II101 88 E : 08.ix.1984, Sn-II 10219 E : 08.ix.l984,Sn-II 10246 E : 09.ix.l984. W coast of Binongk o :Sn-II 10295 E: lO.ix.1984. NE Taka Bone Rate(Tiger Island), Taka Garlarang atoll : Sn-II 11305E: 27.ix.l 984, Sn-II 11372 C : 27.ix.l 984. SWSalayer, N of Pulau Bukuluang Sn-II 11644 C,E : 01.x.l984.

1. '::icrouictyuil pal neri Sctchell (1925 : 106)(Figs. 9, 10)

SETCHELL (1929 : 535).[type locality : Guadalupe Island , in the

Pacific Ocean, off the west coast of Mexico].

Thallus comp osed of netlike blades, attachedby long, aseptate rhizoids ; light green, delicate,orbicular to reniform, up to 7 em tall, withslightly lobed margins . Basal cells cylindrical toclavate, 350-750 urn long, 0 150 um, branchingstellate. Primary cells subcylindrical, 150-375 urnlong, 0 100-I50 urn, L / W : 1-2, branching fla­bellate, opposite or alternate. Secondary cells,150-225 urn long, 0 75-150 um ; L/W : 1-2.5,branching opposite or alterna te. Apical cellsanastomosing or free, 75-160 urn long, 0 40-70urn. Anastomosis accomplished by a smoo thannulate wall thickening at the tip of a cell.Meshes delimited by anastomosing cells orbranches tr iangular to pentangular, largest dia­meter 100-300 urn, Blades adhering firmly topaper when dried.

Ecology : specimens Copp. & PvR. 13485B and 13653 B : shallow infralittoral, epilithic oncoral bould ers ; specimen HEC 4690 : -35 m, oncoral rubbl e.

Distribut ion : Guadalupe Island (type loca­lity).

Specimens examined: Papua New Guinea,Madang Province, Hansa Bay, The Pinn acle:HEC 4690: 17.viii.l980. Bagabag, NW point ofChristmas Bay : Copp. & PvR. 13485 B:02.viii.l990. Bagabag, Badilu village : Copp. &PvR. 13653 B : 08.viii.l990.

Microdictyon vanbosseae Setchell f. explanatumSetchell (1926 : 153) (Figs. I I , 12)

[type locality : Lirung-reef, Salibabu Island ,Indonesia] , SETCHELL (1929 : 543).

Thallus composed of netlike blades. Bladeslight green, delicate, with two distinct types ofmeshes : one, more num erous, angular, delimitedby three or four cells, the others less numerous,larger, delimited by numerous cells. Branchingunilateral, opposite or flabellate. Cells subcylin­drical, 110-375 urn long, 0 50-150 um ; apicalcells anastomosing or free I 10-260 urn long, 045-75 urn, L/W: 1-3. Anastomo sis accompli shedby a smooth annulate wall thickening at the tipof a cell. Small meshes with greatest diameter 90­900 um, large meshes with greatest diameter I 10­5000 urn. Blades adhering firmly to paper whendried.

Ecology : infralittoral, epilithic.

S I P HONOCLAD ALES FRO M PA P UA NEW GUIN EA A ND IN D O NESI A 183

FIGs. 7, 8. - M icrodictyon okamurae Setchell ; (Sn-II 10246). 7 : Branching patt crn ; (500 urn). 8 : Anastomosingcells ; (100 urn).

FIGs. 9, 10. - Microdictyon palm eri Setchell ; (Co pp. & PvR. 13485). 9 : Branching pattern in basal part of theblade ; (500 urn), 10 : Anastomosing cell; (100 urn).

FI Gs. II, 12. - Micr odictyon vanbosseae Setch ell f. explan atum Setchcll (Sn-II 10887). II: Branching pattern form ingsmall and large meshes ; (500 urn), 12: Anastomosing cells ; (100 urn).

184 BELGIAN JOURN AL O F BOTANY 130

Distribution : Indonesia.Specimen examined: Indonesia, Komodo

Island , NE-cape : Sn-II 10887 E : 26.x.l984.

Boergesenia J. Feldmann

See species description (monospecific genus).

Boergcsenia forbesii (Harvey) J . Feldmann (1938 :1503) (Fig. 13)

JAAS UND (1976 : 15, fig), SARTONI (1992 :306; fig. 7b).Valonia forbesii Harvey (1860: 333), [syntype

localities: Ryukyu-retto, Japan ; Sri Lan­ka], Taylor (1950 : 41).

Pseudovalonia forb esii (Harvey) Iyengar (1938 :132) (invalid).

Thallus a light (yellowish) green clavate cellwith basal annular constrictions, up to 3.5 emtall, up to 1.6 em in diameter, frequently curved(at least at the basis). Cells connected with eachother by a septate stoloniferous rhizoidal system.Thalli isolated or clustered.

Ecology : midlittoral, epilithic or on coralrubble in sheltered habitats of the reef, mostlyin shallow reef pools, sometimes on sand-coveredrock. Also observed in seagrass meadows(Bnouxs & HEUS 1986: 38).

Distribution: pantropical.Specimens examined : Papua New Guinea,

Madang Province, Neptune Point: HEC 7620 :27.vi.1988. Barol Point: HEC 7845 : l 8.vii.l 988.Gumbi Bay : HEC 7911 : 25.vii.l988. Bagabag,NW point of Christmas Bay: Copp . & PvR .13506 B : 02.viii. 1990. Nagada Harbour in frontof Gossem Island: Copp. & PvR. 13598 B :07.viii.1990. Mugil Harbour : Copp. & PvR .13831 B : 22.viii.1990. Port Moresby area, Mo ­tupore Island: H EC 6334: 06.vii.l986.

Boodlea Murray et De Toni

Thallus a spongy cushion composed of a 3­dimensional network of branching cells, attach­ment by rhizoids ; anastomosis by tenacula.

Idcn ificatio key

I. a. Long rhizoids frequently arising from the distalside of any cell of the thallus B. vanb osseae

b. Rhizoids , if present, only at the base of thethallus . ... B. composita

Boodlea com posita (Harvey) Brand (1904 : 187)(Figs. 14-20)

B0RGESEN (1940 : 21), B0RGESEN (1946:15), TAYLOR (1950 : 44), EGEROD (1952 : 362),EGEROD (1975: 50), SARTONI (1992: 306).Conferva composita Harvey (1834 : 157), [type

locality : Mauritius].Cladoph ora composita (Harvey) Kiitzing (1849 :

415).A egagropila composita (Harvey) Kiitzing (1854 :

14).Boodlea siamensis Reinbold (1901: 191), [type

locality : Ko Kahd at , Ko Chang Archipe­lago, Thailand] , Taylor (1950 : 41).

? Boodlea coacta Dickie (1876: 451), [type lo­cality: "Osima Harbour" (O-shima, Wa­kayama Prefecture), Japan].

Thallus cushion -like or spongy, light green ,composed of a 3-dimensional network of bran­ching cells, up to 13 cm in diameter, light green;marginal part of the thallus sometimes "Struvea /Phyllodictyon"-like ; young thalli someti mes bear­ing a stipe. Branching in three dimensions, upto the sixth order, regularly opposite to alternate.Primary cells 180-4500 urn long, <0 (110-) 180­640 urn, L/W: 1-5 ; intermediate cells 75-l~00

urn long , <0 (20-) 40-300 urn, L/W : 1-10 ; apicalcells 70-500 (-2700) urn long, <0 (20-) 40-150 urnanastomosing or free; L/ W of all cells stronglyvariable within a specimen. Anastomosis accom­plished by tenacula, less frequently by a cren~late

thickened membrane at the tip of an unmodifiedcell.

Ecology : various habitats: infralittoral, epi­lithic, epipsammic or epiphytic (e.g. on Halimedamacroloba) on coral reefs or in seagrass mea­dows ; or loose-lying, sometimes even free-float-ing. .

Distribution: tropical and subtropical Indo­Pacific.

Specim ens examined: Papua New Guinea,Madang Province, Laing Island: HEC 4431 :25.vi.l980, HEC 6451 : l5.viii .1986, HEC 6535 :

'")'") ... IQ86 HEr (,(,74 .18.viii.1986, HEC 6581 : ~~.V111.. , ~_ .

lO.vii.1986, HEC 7638 : 29.vi.1988, HEC 7690 :05.vii.1988. Boisa, S coast: HEC 7735 :07.vii.1988. Hansa Bay, Durangit reef : HEC7847 : 17.vii.1988. Hatzfeldthafen : HEC 7903 :

SIPHONOCLADALES FROM PAPUA NE W GU INEA AND INDONESIA 185

13

-~-6-------17------<::::::(~"S/-'V1 -11j~~'ts\~~

19

FIG. 13. - Boergesenia forbesii (Ha rvey) J . Feldmann ; (H EC 7845) ( I mm).FIGs. 14-20. - Boodlea composita (Harvey) Brand. 14. Branchin g pattern in the marginal part of a thallus; (Copp.

& PvR. 1335 1) (500 urn) . IS : Branching pattern in the central part of the thallus ; (Copp. & PvR . 13133) (500 urn). 16­19: Detail s of branching patterns and anastomosing cells ; (HEC 8078) (200 urn). 20 : Anastomosing cell ; (Co pp. & PvR.13133) (50 urn),

FIGs. 21, 22. - Boodlea vanbosseae Reinbold ; (Sn-ll 1011 3). 21 : Branching cells with long rhizoids ( I mm) . 22.Cell tip with tenaculum ; (100 urn).

186 BELGIAN JOURNA L O F BOTANY 130

21.vii.1988. Gumbi Bay: HEC 7934 : 25.vii.l 988,HEC 7943: 25.vii.l988. W Malam al Island :HEC 8067 : 07.viii.1988. Kranket Island: HEC8078: 07.viii.1988, Copp. & PvR. 13133 B, B' :13.vii.1990. SW of Wongat Island: Copp. & PvR .13157 B : 13.vii.1990. Island N of Dem asa Island :Copp. & PvR . 13170 B : 15.vii.1990. BetweenSinub and Wongat Island : Copp. & PvR . 13237B : 18.vii.1990. Ru o Island : Copp. & PvR . 13262B: 19.vii.1990. Sarang Harbour: Copp. & PvR .13351 B : 25.vii.1990. Bagabag, NW point ofChristmas Bay: Copp. & PvR. 13509 B :02.viii.1990. Nagada Harbour: Copp. & PvR .13134 B : 13.vii.l990. N of Pig (Tab) Island, innerslope of reef : Copp. & PvR. 13588 B :05.viii.l990. Port Moresby area, Motupore Is­land : HEC 6339, HEC 6347, HEC 6354 :06.vii.l986, HEC 10203: 21.vii.l 994. Horse Sh oeReef: HEC 10340 : 30.vii.1994. Loloata Island:ODC 250: 28.vii.1994.

Indonesia, Banda Sea, Tukang Besi Islands,harbour of Taipabu, NW coast Binongko: Sn­II 10329 E, Sn-II 10330 E, Sn-II 10351 E:10.ix.l984. E of Komodo Island, Selat Linta : Sn­II 10842 C, Sn-II 10843 C: 18.ix.1984. S Tomea :Sn-II 11165 C : 07.ix.l984. NE Taka Bone Rate(Tiger Island ), E coast Ta rupa Kecil : Sn-II 11278E, 11279 E : 24j 26.ix.l 984.

Boodlea vanbosseae Reinbold (1905 : 148)(Figs. 21, 22)

[synt ype localities : spread over Ind one sia],GEpp & GEI'P (1909 : 375), R EI NBOLD in WEBER­VAN BosSE (1913 : 70), B0RGESEN (1936: 63),EGEROD (1975 : 52), SARTONI (1992 : 307).

Thallus cushion-like , subspherical to irregu­lar, up to 8 em in diameter, composed of a 3­dimensional network of branching cells. Bran ­ching in three dimension s, opposite or unilateral.Primary cells 360-2200 J.1m long, <0 150-300 J.1m ;intermediate cells 75-1100 J.1m long, <0 50-225 J.1m ,LjW : 1-7 ; apical cells 100-525 (3600) J.1m long,<0 35-225 J.1m ; short apical cells sometimes witha tenaculum and anastomosing. Long rhizoidsar ising from the distal end uf an y cell of thethallus, 1800-6300 J.1m long, <0 40-70 J.1m, witha crenulate thickened membrane at the tip.

Ecology : infralittoral frin ge or infr allitor al,

epilithic or epipsa mmic, on coral reefs or III

seagrass meadows.Distribution : tropical Indo-Pacific.Specimens examined : Indonesia, Banda Sea,

M aisel Islands: Sn-II 10113 E: 07.09.1984, Sn ­II 10117 E : 07-09-1984. Tukang Besi Islands,Kaledupa reef: Sn-II 10260 E : 09.ix.1984. Tu­kang Besi Islands, W coast of Binongko : Sn -II10292 E : lO.ix.1984. Maisel Islands: Sn-II 10383E: 04j07.ix.l984, Sn-II 10390 E: 07.ix .1984, Sn­II 10404 E : 05.ix.1984, Sn-II 10421 E: 05.ix. 1984.NE coast of Sumba, E of Melol o : Sn-II 10442:14.ix.1984. NE of Taka Bone Rate (Tiger Island),S of Tarupa Kecil : Sn -II 11216 C : 13/l7.x.1984,Sn-II 11242 C : 13jl7.x.1984.

Phyllod ictyon J. E. Gray

Same as Stru vea but laterals produced bycentripetal wall ingrowths dividing parent cellsinto approximately equal halves (K RA FT &WYNNE 1996 : 140).

Identification key (including Stru vea)

1. a. Thallus with a conspicuous stipe with basa lannular constriction s, blades tr iangular to ellip-tical . . S. elegans

b. St ipe, if present , without basal annular constric-tions, blades elliptical to irregular . ..... 2

2. a. Apical cells up to 450 urn long, blades less than4 cm long, often ana stomosing P. anastomosans

b. Apical cells up to 7000 urn long, blades up toI I ern long, not anastomosing ... . ... P. gardineri

Phyllodictyon anastomosans (Harvey) Kraft &Wynne (1996 : 131, fig. 16-25) (Figs. 23, 24)

Cladophora ? anastomosans Harvey (1859: pI.CI) , [type locality : Fremantle, WesternAustralia].

Pterodictyon anastom osans (H arvey) J . E. Gray(1866 : 70).

Struvea anastomosans (H arvey) Piccone & Gru­now ex Piccone (1884: 20), TAYLOR (1928 :73), B0RGESEN(1952: 7-8), EGEROD (1952:359-361), CHAPMAN (1961 : 93), EGEROD(1971 : 123), EGERO D (1975 : SO), uART07'~ r

(1992 : 317-319).Struvea delicatula Kutzing (1866 : 1), [type lo­

calit y : New Caledonia].

SIPHONO C LADALES F RO M PAPU A N E W GU IN EA AND INDONESIA 187

Struvea tenuis Zanardini (1878 : 39), [type loca­lity : New Guinea].

Struvea multipartita Pilger (1920), [type locality:not found in literature].

Thallus composed of stipitate, netl ike, oval,ovate or irregular blades, light green ; anchoredby multicellular, branched rhizoids. Stipe some­times branched, up to 4 em long, (2) 190-360 (­630) urn, without annular contrictions. Blade upto 3.4 mm long and 2.5 mm wide, branching inthe blade opposite, 3-,4- or 5-pinnate. Lower cellsof the main axis 180-750 urn long, (2) 180-1000(2200) urn ; primary branchlets upwardly curved ;cells of the first and higher order branchlets (75-)150-600 urn long , (2) 100-340 um ; apical cells ofthe secundary and higher order branchlets anas­tomosing or free, 150-450 (-1250) urn long, (2) 75­120 urn. Anastomosis accomplished by tenacula,either between cells of the same blade or betweencells of adjacent blades resulting in cushion-likethalli .

Ecology: infralittoral (-0.5 / -40 m), epilithicon coral boulders or epiphytic on Galaxaura.

Distribution : pantropical (at least partly asa result of so many taxa synonymized with it(KRAFT & WYNNE 1996 : 139)).

Specimens examined: Papua New Gu inea ,Madang Province, Laing Island: HEC 4492 :15.vii.l980, HEC 6670: lO.vii.l986, HEC 6675 :10.vii. 1986. Boisa, S coast : HEC 7748:07.vii.1988. N of Wongat Island: Copp. & PvR.13466 B: 0 I.viii.1990. Bagabag, NW point ofChristmas Bay: Copp. & PvR. 13508 B :02.viii.l990. Bagabag, New Years Bay: Copp. &PvR.13541 B 02.viii.1990 . N of P ig (Tab) Island:Copp. & PvR . 13586 B : 05.viii.l990.

Remark : the variability within this specieswas discu ssed by seve ral authors (EGEROD 1975 :50-51).

Synonymies : Struvea delicatula was treatedas a taxonomic synonym by M URRAY & BOODLE(1888), Struvea tenuis was considered as a taxo­nomic synonym by CRIBB (1960), Struvea mul­tipartita was treated as a taxonomic synonym bySTEENTOFT (1967).

Phyllodictyon gardineri (A. Gepp & E. Gepp)Kraft & Wynne (1996 : 139) (Figs. 25-27)

Stru vea gardiner i A. Gepp & E. Gepp (1908 : 166­167) [type locality : Cargados Carajos,North of Mauritius], A. Gepp & E. Gepp(1909 : 376-377) .

Thallus composed of stipitate, netlike blades,anchored by multicellular, br an ched rhizoids,light green. Stipe unbranched, without annularcon strictions, 40-50 mm long, (2) 700 urn, Bladesoval, up to 11 em high and 6.5 ern broad, L/W: 1.7-2.5, branching regularly opposite in youngblades, becoming less regular in mature ones, 3­pinnate. Lower cells of the main axis 1.8-3.5 mmlong, (2) 550-700 um ; cells of th e primary bran­chlets 1.5-7.0 mm long, (2) 300-450 urn, L /W :up to 30 or even more ; apic al cells-free o ranastomosing, 2.7-7 mm long, (2) 300 urn , L/W :up to 30 or more. Anas tomosis by tenacula.

Ecology : infralittoral (-15/ -30 m), epipsam­rmc.

Distribution : up to now exclusively knownfrom the type locality.

Specimens examined: Papua New Guinea,Madang Province, Laing Island: HEC 4548 :22.vii.1980, HEC 4696: 18.viii.1980 , HEC 7760 :13.vii.l988.

Remark: the large blade-surface could beinterpreted as an adaptation to low light intensitiesat thi s depth.

Struvea Sonder

Thallus composed of one or more netl ike,stipitate blades. Branching of the blade-cells op­posite, essentially in one plane ; anastomosis bytenacula. Laterals produced by segregative celldivision (KRAFT & WYNNE 1996 : 140).

Struvea elegans B0 RGES EN (1912 : 264) (Figs. 28,29)

[synt ype locality : Virgin Islands], B0RGESEN(1913: 51-54), TAY LOR (1928 : 74), KRAFr &WYNNE (1996 : 139).

Thallus composed of stipitate, netlike blades,anchored by multicellular, branched rhi zoids,dark green. Stipe branched or unbranched, upto 5.5 em long, (2) 800-1100 um ; one or morebasal annular constrictions present ; blades trian­gular to oval-ovate, up to 26 mm long and 16mm wide , L/W : 1.5-3 ; branching regularly

188 BELGIAN JOURNAL OF BOTANY 130

opposite, 2- to 3-pinnate. Lower cells of the mainaxis 600-1000 (-1440) urn long, 0 450-675 um ;cells of the primary and secundary branch1ets225-600 (-1350) urn long, 0 150-370 um ; apicalcells free or anastomosing, 110-300 urn long, 0300-1200 urn, Apical cells of the secondary bran­chlets frequently anastomosing with the upper orlower primary branchlets resulting in a zigzagpattern. Anastomosis accomplished by tenacula.

Ecology : infralittoral (depth -1 j -45 m), epi­lithic on coral substrate. No morphological dif­ferences between specimens growing at differentdepths.

Distribution : Atlantic Ocean, CaribbeanSea: Virgin Islands ; Gulf of Mexico : Florida ;Indian Ocean : Seychelles, Reunion.

Specimens examined: Papua New Guinea,Madang Province, Laing Island: HEC 4395 :22.vi.l980, HEC 4413: 24.vi.1980, HEC 6450 :15.viii.l986, HEC 6460 : 15.viii.l986. Boisa, Scoast : HEC 7734: 07.vii.l988. N of Pig (Tab)Island: Copp. & PvR. 13587 B, B' : 05.viii.l990.Port Moresby area, Motupore Island : HEC6348: 06.vii.1986, HEC 10236 : 23.vii.l994. Lo­loata Island : HEC 10366: 3l.vii.l 994, HEC10437: 05.viii.1994.

Valoniaceae

Dictyosphaeria Decaisne

Thallus pseudoparenchymatous, subsphericalor flattened, solid and tough or hollow and stiff­brittle, or monostromatic, composed of vesicularor polygonal cells which are held together bysmall hapteroidal cells; trabeculae (spine-shapedoutgrowths) sometimes present on the inner cellwalls. Cells divide by segregative cell division.

Identification key

1. a. Thallus hollow when young, cup-like when older,trabeculae absent . . .. . D. cavernosa

b. Thallus massive, trabeculae present on the innerrellw~ 2

2. a. Thallus dorsoventrally compressed, cells polygo-nal D. versluvsii

b. Thallu s not compressed , composed of an irre­gular aggregation of spherical cells ... D. ocellata

Dictyosphaeria cavernosa (Forsskal) Borgesen(1932 : 2) (Figs. 30-33)

B0RGESEN (1940: 12), B0RGESEN (1946 :13), TAYLOR (1950: 43), EGEROD (1952: 350),TAYLOR (1960: 116), JAASUND (1976: 15),SARTONI (1992: 319).VIva cavernosa Forsskal (1775 : 187), [syntype

localities: "Gornfodae" [AI-Qunfudhah],Saudi Arabia ; Mokha, Yemen].

Valonia favulosa C. Agardh (1823 : 432), [typelocality : "Ravak" [Rauki], Waigeo Island,Moluccas, Indonesie],

Dictyosphaeriafavulosa (c. Agardh) Decaisne exEndlicher (1843: 18), WEBER-VAN BOSSE(1913: 63)

Thallus a pseudoparenchymatous cushion,3-8 em across , very stiff-brittle, composed ofpolygonal cells, dark green ; young thalli sub­spherical and hollow, becoming convoluted andruptured when old , the 'roofs' disappearing, re­sulting in cup-like structures. Cells polygonal, 01-2 (-3.5) mm ; cohering by minute bifurcatehapteroidal cells, 0 40-60 urn, arranged along thecontactlines. No trabeculae present.

Ecology: infralittoral fringe or infralittoral(down to -30 m), epilithic on horizontal surfacesof coral boulders ; some specimens epipsammicin seagrass meadows or lagoons.

Distribution: Red Sea, tropical Indian, Pa­cific and western Atlantic Ocean .

Specimens examined : Papua New Guine a,Madang Province, Gumbi Bay : HEC 7912 :25.vii.1988. Beliau Island, W Bay : Copp. & PvR.13076 B : 1O.vii.1990. Bagabag, NW point ofChristmas Bay : Copp. & PvR. 13490 B:02.viii.1990.

Indonesia, Banda Sea, Maisel Island: Sn­II 10112 E: 07.ix.1984, Sn-ll 10376 E: 04/07.ix.1984. Banda Sea, Tukang Besi Islands,Kaledupa reef: Sn-ll 10190 E: 06j08.ix.l984.Banda Sea, Tukang Besi Islands, S coast ofTomea: Sn-ll 11193 C: 07.ix.1984. E of KomodoIsland, Selat Linta: Sn-ll 10800 E: 18.ix.1984,Sn-ll 10832 E: 18.ix.1984. NE Taka Bone Rate(Tiger Island), S of Tarupa Kecil: Sn-ll 11235C: 13/ 17.x.1984.

Dictyosphaeria ocellata (Howe) Olsen-Stojkovich(1985 : 62) (Figs. 34-36)

Valonia ocellata Howe (1920 : 603), [type locality:

SIPH O N O CL A D AL ES FRO M PA P UA NEW G UI N EA AND INDON ESI A 189

27

29

25

F IGs. 23, 24. - Phyllodictyon anastomosans (Harvey) Kraft & Wynne ; (HEC 6675). 23 : Stipita te blade ; ( I mm).24 : Anastomosing cells ; (100 11m).

F IGS. 25-27. - Phyllodictyon gardineri A. Gepp & E. Gepp ; (HEC 7760). 25 : Detail of blade ; (I em ). 26 : Margin alpart of the blade ; (I mm) . 27 : Cell tip with tenaculum ; (100 11m).

FIGS. 28, 29. - Struvea elegans Bergesen ; (H EC 6460). 28 : Stipitat e blade; (2 mm). 29 : Cell tip with tenaculum ;(100 11m).

190 BELGI A N JO URN AL OF BO TA NY 130

Watling's Island, Bahama Islands] , TAYLOR(1928 : 75), CHAPM AN(1961 : 98), CORDERO(1977 : 25), Ln TLER et at. (1989 : 60)

Thallus forming irregular clumps, 1-9 emacross, composed of subspherical aggregations ofspherical cells, anchored by thick basal rhizoids,dark green. Cells 0 2-4 mm, cohering by minutehapteroidal cells, 40-50 urn long, 0 20-100 urn,ar ranged along the contactlines. Trabeculae onthe inner cell walls with blunt tips, 150-500 urnlong, 0 20 urn,

Ecology: high infralittoral, epilithic on ver­tical coral boulders; one specimen on Hypnea­mats.

Distribution : Atlantic Ocean : CaribbeanSea; Pacific Ocean : Philippines.

Specimens examined : Papua New Guinea,Motupore Island: HEC 6359 : 06.vii.l9 86, HEC10206 : 21.vii.l 994.

Dictyosphaeria versluysii Weber-van Bosse (1905 :104) (Figs. 37-39)

[syntype locality : various localities in Indo­nesia], GEPP & GEPP (1908 : 168), GEPP & GEPP(1909 : 378), WEBER-VAN BOSSE (1913: 64),EGEROD (1952 : 351 , 354), EGEROD (1974 : 140),J AASUND(1976: 15), SARTONI (1992 : 319).Dictyosphaeria vanbosseae Bergesen (1912 : 256),

[type locality : Cane Bay, St. Croix, VirginIslands].

Dictyosphaeria bok otensis Yamada (1925 : 81),[type locality : Pescadores Islands, For­mosa].

Dictyosphaeria australis Setchell (1926 : 79), [typelocality : not found in literature].

Dictyosphaeria setchellii Bergesen (1940: 12),[type locality : Mauritius]

Thallus solid, extremely tough, more or lessflattened, frequently several specimens laterallyimbricated like a jig-saw, greyish green ; pseudo­parenchymatous, 1-3 em across, 1-2 em tall,composed of polygonal cells 0 0.5-2 mm. Ad­jacent cells cohering by minute unifurcate orbifurcate hapteroidal cells 3U- 11U urn long, 10 2u­50 um ; intercellular spaces bridged by longerhapteroidal cells. Spinulose trabeculae on theinner cell walls, 120-150 urn long, 0 6-10 urn.

Ecology: infralittoral fringe or infralittoral(down to - 30 m), epilithic on horizontal surfacesof coral boulders ; some specimens epipsammicin seagrass meadows (Thalassodendron).

Distribution : tropical Indo-Pacific.Specimens examined: Papua New Guinea,

Madang Province, Laing Island : HEC 4254 :28.v.l980. SW of Wongat Island: Copp. & PvR.13153 B : 13.vii.1990. between Tausch and SekIsland: Copp. & PvR. 13286 B: 20.vii.1990.Bagabag , New Years Bay : Copp. & PvR. 13540B: 02.viii.l990. Bay at Malala Village : Copp. &PvR. 13740 B: 18.viii.1990. Mugil Harbour:Copp. & PvR. 13813 B : 22.viii.90. Port Moresbyarea , Motupore Island: HEC 6673: lO.vii.l986,HEC 10264 : 26.vii.l994.

Ind onesia, Banda Sea, Tukang Besi Island s,Kaledup a reef: Sn-I1 10248 E : 09.ix.1984. MaiselIslands : Sn-I1 10389 E: 04 & 07.ix.l984, Sn-II10402 E : 05.ix.1984. NE coast of Sumba, E ofMelolo: Sn-I1 10443 C: 12jl4.ix.1984, Sn-II10456 E: 121I4.ix.l 984, Sn-I1 10518 E:13.ix.1984. N of Sumbawa, bay of Sangar : Sn­II 11073 E: 21.ix, 29/31. x.1984. NE Taka BoneRate (Tiger Island) , S of Tampa Kecil: Sn-II11234 E: 13jl4.x.1984. NE Taka Bone Rate(Tiger Island), Taka Garlarang atoll : Sn-II 11307C: 27.ix.1984.

Valonia C. Agardh

Thallus composed of branched vesicular,coenocytic cells, forming hemispherical domes, orirregular cushions ; adjacent cells held togetherby tenacula.

Identification key (including Ventricaria ventri­cosa )

I. a. Thallus consisting of a single spherical to ovatecell or an aggregation of such cells, tenaculaabsent . .. 2

b. Thallus with elongate, clavate cells, tenacula~~m .. . 3

2. a. Thallus a single cell, never branched, dark bluishgreen, cell wall refract ive and shiny .

........... Ventricaria ventricosab. Thallus generally with daughter cells, sometimes

unbranched, grass green to dark green, cell wallsnot refractive Valonia ma crophysa

3. a. Thallus prost rate (cells 'creeping'), cells irregularly

30~32---

[\ 33~_.J

34

d35

37 ~~~ ~ ~)oo}"..'"

39!'Io~Q7

~~38- &iff »»

'-----'

41

~O

43 go()

'8a44o

CJoo

o

FIGS. 30-33. - Dictyosph aeria cavernosa (Fo rsskal) Borgesen ; (Co pp. & Pvr. 13076). 30: Young, subspherical andhollow thallus ; (I em). 31 : Older , convoluted and ruptured thallus ; ( I em). 32 : Hapteroidal cells arranged a long thecontactlines ; (200 urn), 33 : Hapteroidal cells; (50 urn).

F IGS . 34-36. - Dictyosphaeria ocellata (Howe) Olsen-Stojkovich ; (HEC 1026). 34 : Young thallus ; (5 mm) . 35 :Hapteroidal cells ; (50 urn), 36 : Trabeculae ; (50 urn).

FIG S . 37-39. - Dictyosphaeria versluysi i Weber-van Bosse ; (HEC 6673). 37 : Hapteroidal cells arra nged in rows;(200 urn). 38 : Hapteroidal cells ; (50 urn). 39 : Trabeculae; (50 urn).

FIGS . 40, 41. - Valonia aegagropila C. Agardh; (Sn-II 10250). 40 : Young thallus with basal rhizoids ; ( I mm) . 41 :Tenacula; (200 urn).

F IGS . 42-44 . - Valonia fa stigiata Harvey ex J. Agardh; (HEC 7782). 42 : Branching cells ; ( I em). 43 : Circular clusterof tenacula ; (500 urn). 44 : Tenaculum ; (200 urn).

FIG.45. - Valonia utricularis (Roth) C. Agardh : Branching cells ; (HEC 7478) ( I em).

192 BELGIAN JOURNALOF BOTANY 130

shaped, often arcuate ; branching distal or lateral........ V. ut ricular is

b. Thallus hemispherical (cells radially placed) ;branching mainly distal . . 4

4. a. Branching regular, di- to polychotomous, celllength exceeding 5 mm, tenacula grouped incircles . . V.f ast igiata

b. Branching irregular, di- to polychotornous, celllength smaller than 3 mm, tenacula randomly

. arranged on the cell surface V. aegagropila

Valonia aegagropila C. Agardh (1823 : 429)(Figs. 40, 41)

[lectotype locality : Venetia, Italia fideEGEROD (1952 : 348) and OLSEN & WEST (1988 :106)], TAYLOR (1928 : 74), TAYLOR (1950: 41),CRIBB (1960), CHAPMAN (1961: 98), TAYLOR(1966 : 347).

Thallus hemispherical or cushion-like, com­posed of radially placed, irregularly branchingcells, dark green. Cells vesicular, clavate or ar­cuate, 2.5-11 mm long, (0 1-2.5 mm . Mainbranching irregular (not markedly di- or poly­chotomous) with numerous lateral, smaller branch­lets. Adjacent cells cohering by tenacula, ran- 'domly arr anged along th e lateral cell walls.

Ecology: in seagrass meadows.Distribution: cosmopolitan in warm seas.Specimens examined : Indonesia, Banda Sea,

Tuka ng Besi Islands, Kaledupa reef : Sn-II 10250C : 09.ix.1984.

Valonia fastigiata Harvey ex J . Agardh (1887 :101) (Figs. 42-44)

[syntype localities : Sri Lanka ; Tonga], GEPP& GEPP (1909 : 379), B0RGESEN (1946: 13),J AASUND (1976 : 15), SARTONI (1992: 321)

Tha llus a hemispherical or flatt ened solidcushion , dark green, 5-15 em across, composedof densely packed, erect, radially arranged cells ;anchored by small, asepta te rhizo ids. Mai n bran­ching regularly di- to polychotom ous at the distalend ; small lateral branchlets can also be for med.Cells subcylindrical to clavate , (5-) 8-15 (-22) mmlong, (0 2.5-5 rnm, Lj W: 2-3.5 (-4.5). Adjacentcells cohe ring by circular clusters of tenacula.Septa without trabeculae.

Ecology : infralitt oral fringe or infralittoral(down to -30 m), epilithic on coral substr ate orepipsammic in seagrass meadows.

Distribution : Somalia, Tanzania, Madagas­car, Mauritius, Reunion, Cargados Carajos, Sayade Malha Bank, Seychelles, Chagos Archipelago ,Sr i Lanka, Burma, Singapore, Indonesia, Malay­sia, Papua New Guinea, northern Australia,Micronesia, Fiji, Philippines, Taiwan .

Specimens examined: Papua New Guinea ,Madang P rovince, Mugil Harbour & VidariIsland: HEC 7602 :' 27.vi.l988. Laing Island:HEC 7782 : 13.vii.l988. Hatzfeldh afen : Co pp. &PvR. 13333 B: 21.vii.l 990. N of Pig (Tab)Island : Copp. & PvR. 13667 B : 05.viii.1990.Port Moresby area, Motupore Island: HEC10232: 23.vii.1994. South Patch Reef: HEC10295 : 28.vii.l994.

Indonesia, E of Komodo Island , Selat Linta :Sn-II 10810 E : 18.ix.1984. SW Salayer, E ofPu lau Gu ang : Sn-II 11395 E : 06.x.l984.

Valonia macrophysa Kut zing (1843 : 307)

[type locality : Hvar Island , Yugoslavia],TAYLOR (1928 : 75), B0 RGESEN (1913: 29),CHAPMAN (1961 : 97).

Thallus a single or a cluster of vesicular,clavate to pyriform or subspherical, dark greencells, anchored by basal rhizoids. Primary cells2-5.5 em tall , (0 1-2.5 em; apical or lateralbranchlets smaller (7-17 mm long, (0 3-6 mm ) ;cells not cohering by tenacula.

Ecology : infral ittor al on reef.Specimens exa mined : Indonesia, Band a Sea,

Tukang Besi Islands, Kaledupa reef : Sn-II 10196C : 06j08.ix.l984.

Valonia utricularis (Roth) C. Agardh (1823:431) (Fig. 45)

TAYLOR (1928: 75), TAYLOR (1950: 41),B0 RGESEN (1932: 1), B0 RGESEN (1940 : II ),EGEROD (1974 : 139), EGEROD (1975 : 45).Conferva utricularis Roth (1797 : 160), [type

locality : Mediterranea n Sea].

Thallus for ming succulent, stiff-britt le, com­pact or diffuse repent mats, 7-14 em across,composed of vesicular cells, dark green. Branch ingirregular, di- polychotomous, distal or la teral ,Cells clavate, arcuate or irregularly shaped (3-)5-20 (-30) mm long, (0 (3-) 4-6 (-8) rnm, L jW :1-4. Adj acent cells laterally cohering by tenaculawhich are randomly arranged along the cell walls.

SIPHONOC LA D A LES F R O M PAPUA NEW GUINEA AND INDONESIA 193

DISCUSSION

This study reports five new record s for theregion: Microdictyon okamurae (collected indifferent locations in the Banda and Flores Sea),Microdictyon palmeri and Phyllodictyon gardi­neri (from Madang Province, N Papua NewGuinea) , and Dictyosphaeria ocellata (from Mo­tupore Island, S Papua New Guinea).

Mi crodictyon palmeri, Microdictyon van­bosseae, Phyllodictyon gardineri and Struveaelegans are unique occurrences compared withadjacent tropical areas : Taiwan (LEWIS & NORRIS1987), Ph ilippines (SILVA et al. 1987), Micronesia(TSUDA 1977, 1981), Solomon Isles (WOM ERSLEY& BAILEY 1970), N-Australia (LEWIS 1987), Fiji(SOUTH & KASAHARA 1992, SOUTH et al.1993).Species richness is lower than in the Philippines,about the same as in N-Australia, Micronesia andTaiwan , and higher than in the Solomon Islesand Fiji.

About half of the species is also record edin the western Indian Ocean (Anadyomene pli­cata, A. stellata, Boergesenia forbesii, Boodleacomp osita, Dictyosphaeria cavernosa, D. versluy­sii, Microdictyon okamurae, Phyllodictyon anas-

Ecology : infralitoral (depth 1-35m), epilithic Ecology: infralittoral (down to -20 m), epi-on coral substrate or epipsammic in seagrass lithic on coral rubble or corals . Also observedmeadows. No morphological differences were in seagrass meadows (HEIJS & BROUNS 1986 :observed between specimens growing at different 38).depths . Distribution: pantropical to subtropical.

Distribution : Caribbean Sea, Bahamas, Flo- Specimens examined : Papua New Guinea,rida, Bermuda, Madeira, Spain, Mediterranean Madang Province, Laing Island: HEC 4235 :Sea, Bahrain, Kenya, Madagascar, Reunion, Mau- 27.v.1980, HEC 7477 : 17.vi.l988. Kranket Is­ritius, Seychelles, Pakistan, India, Maldive Is- land: HEC 7567 : 23.vi.1988. Gumbi Bay: HEClands, Chagos Archipelago, Sri Lanka, Thailand, 7932: 25.vii.l988. W of Malamal Island : HECNicobar Islands, Malaysia, Indonesia, Papua 8076 : 07-viii-1988. Bagabag, NW point of Christ­New Guinea, Micronesia, Fiji, Philippines, Tai- mas Bay: Copp. & PvR. 13491 B. Nagadawan. Harbour : 13136 B: 13.vii.l990. Port Moresby

Specimens examined: Papua New Guinea, area, Motupore Island: HEC 6288: 04.vii.l986.Madang£ro-'yince, _Laing_Island_ : _HEC_423~_:_LQloataJsland_:_HEC_I0329_:29 .yii.19_9~._ _27.v.l980, HEC 4455 : 27.vi.l980, HEC 7478: Indonesia, Banda Sea, Tukang Besi Islands,17.vi.1988. Gumbi Bay : HEC 7916 : 25.vii.l988. Kaledupa reef : Sn-II 10247 E : 09.ix.1984, Sn-Bagabag, NW point of Christmas Bay: Copp. II 10211 E: 06/08.ix.1984, Sn-II 10218 E : 06/& PvR. 13511 B : 02.viii.l990 . Mugil Harbour : 08.ix.1984. W-coast Binongko : Sn-II 10274 E:Copp. & PvR. 13812 B : 22.viii.1990. Port Mo- 1O.ix.84. Maisel Islands , W of Kaurangka : Sn-resby area, Motupore Island : HEC 6298: II 10391 E: 05.ix.l984. Maisel Islands: Sn-II05.vii.1986. 10367 E : 04&07.iv.1984.

Indonesia, NE coast of Sumba, E of Melolo :Sn-II 10565 E : I2fl4.x.l984. E of KomodoIsland , Selat Linta: Sn-II 10723 E : 22.x.I984,Sn-II 10852 E: 18.ix.1984. Komodo Island , NEcape: Sn-ll 10892 C: 26.x.1984. NE Taka BoneRate (Tiger Island) , S of Tarupa Kecil : Sn-II11239 C : 13&17.x.1984.

Ventricaria Olsen & West

A monospecific genus.

Ventricaria ventricosa (1. Agardh) Olsen & West(1988: 104)

SARTONI (1992 : 323, fig. 14E).Valonia ventricosa J . Agardh (1887: 96), [lecto­

type locality : Guadeloupe, West Ind ies fideOLSEN & WEST (1988 : 104)]. WEBER-VANBOSSE (1913: 60), TAYLOR (1928 : 75),B0RGESEN (1940 : 11, 1948 : 20), EGEROD(1952: 347, 348), CHAPM AN (1961 : 95),TAYLOR (1966: 347), J AASUND (1976: 13).

Thallus dark bluish green, consisting of asingle, unbranched, aseptate , coenocytic, vesicular,obovate to clavate cell, (2) (1-) 2-6 (-10) em.Attachment by aseptate rhizoids at the base ofthe cell. Older specimens strongly covered bycrustose Corallinaceae, sponges, ...

194 BELGIAN JOURNAL OF BOTANY 130

tomosans, Valonia aegagropila, V. fastigiata, V.macrophysa, V. utricularis, Ventricaria ventri­cosa). However, this is not surprising as theycomprise mainly pan-(sub)tropical species.

The genera Boodlea and Phyllodictyon aredifficult to distinguish. Life history and molecularevidence show these two genera to be very closelyrelated (BODENBENDER & SCHNETTER 1990,KOOISTRA et al. 1993) or even congeneric(EoERoD 1975). In a phenetic study, a morpho­logical continuum between specimens of Boodleacomposita and Phyllodictyon anastomosans couldbe observed from a principal component analysis.Phyllodictyon gardineri and Struvea elegans (usedas outgroup in the analysis) were clustered se­parately (LELIAERT 1996: 79-86). For speciesidentification under the current classification acombination of classical characters and charactersderived from this study (branching order, cellulardimensions) is recommended.

Difficulties were also encountered in iden­tifying specimens of Valonia. Many workers havecommented on the lack of clarity of speciesconcepts within the genus (OLSEN & WEST 1988 :103).

This work is meant to give the initial impetusto a broader study of the Siphonocladales-Cla­dophorales complex (SCC) in the tropical IndianOcean and the Western Pacific. Aims of ourfuture research are the understanding of evolu­tionary relationships within the SCC and biogeo­graphic diversification of this complex, using bothmorphological and molecular data.

ACKNOWLEDGMENTS

We are very grateful to M. Jebb (ChristensenResearch Institute), J. M. Ouin (Laing Island BiologicalStation) and J. Rewald (Motupore Island) for theirhelp with the accommodation and the organisation ofthe field-work. The second author acknowledges forhis participation in the Snellius-II expedition. C. DeMaire is acknowledged for mounting the specimensand for their original labelling. This research wassponsored by the Fund for Scientific Research, Flanders(FW.O. , Belgium : projects nos. 2.9006.86, 2.9001.90,2.9008.90.).

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