sign in sign up
<<
Home , Bryopsis plumosa, Seychelles

Belg. Journ. Bot. 129 (1) : 47-65 (1996)

THE CODIALES (EXCLUDING CODIUM) () FROM , AND

Eric COPPEJANS & Caroline VAN DEN HEEDE Vakgroep Morfologie, Systematiek en Ecologie ; Laboratorium Plantkunde Universiteit Gent, Ledeganckstráat 35, B-9000 Gent, Belgiurh

ABSTRACT. - Descriptions and illustrations are provided of one species of Boodleopsis, seven taxa of and one species of Trichosolen, based on specimens collected in Kenya, Tanzania and the Seychelles from 1985 to 1996 : Boodleopsis pusilla, Bryopsis hypnoides, B. indica, B. pennata var. pennata, B. pennata var. leprieurü, B. pennata var. secunda, B. plumosa, B. sp. and Trichosolen sp. Bryopsis indica is a new record for Tanzania, Bryopsis hypnoides is new for the Seychelles and Trichosolen sp. is new for Kenya. RÉSUMÉ. Les Codiales (excl. Codium) (Chlorophyta) du Kenya, de Tanzanie et des Seychelles. - Les especes suivantes, récoltées au Kenya, en Tanzanie et aux Seychelles entre 1985 et 1996, sont décrites et illustrées : Boodleopsis pusilla, Bryopsis hypnoides, B. indica, B. pennata var. pennata, B. pennata var. leprieurü, B. pennata var. secunda, B. plumosa, B. sp. et Trichosolen sp. Bryopsis indica est une nouvelle espece pour la Tanzanie, Bryopsis hypnoides est nouvelle pour les Seychelles et Trichosolen sp. est nouvelle pour le Kenya.

INTRODUCTION MATERIALS AND METHODS

The aim of the present study is the inventory Material was collected along the Kenyan coast by and description of the Codiales along the East phycologists of the University of Gent (), African coast, resulting in a useful identification during 9 field trips over a period of 8 years (see VAN key. These data will be incorporated in a flora of DEN HEEDE & COPPEJANS 1996). The specimens fram the Chlorophyta from the western lndian Ocean, were gathered in July-August 1993 by Cop­ which is in preparation. Such a marine macro­ pejans & Van den heede and in August 1994 by Coppe­ jans & De Clerck. Those from Mafia lsland, Pemba algal flora, comparable to that of LAWSON & lsland and Mbudya lsland (Tanzania) were collected JOHN (1987) for the West African coast, is still in January 1996 by Coppejans & De Clerck. Qne non-existent, although indispensable as a basis month (December-January) of collecting around the for futher ecological field work, monitoring and Seychelles took place during the Netherlands lndian managing coastal ecosystems. Qcean Expedition 1992-1993 (COPPEJANS el al. 1994) (map 1). 48 BELGIAN JOURNAL OF BOTANY 129

INDIAN OCEAN

... : Mahé

SEYCHELLES

COMOROS::> 500 km

MAP l. Map of the study .

Sampling was done by wading in the intertidal The anatomical analysis (number of measurements : zone at low tide and by in the subtidal areas. main axis n = 10 per specimen, pinnulae n = 30) was During the Seychelles Expedition, SCUBA-diving carried out on all the collected specimens with a light (down to 25 m) and dredging from the ship resulted in microscope; drawings were made by camera lucida. specimens from deeper waters (COPPEJANS el al. 1994). ldentification was mainly based on the publications of Ecological data and vegetation transects or species lists JAASUND (1976), MOORJANI & SIMPSON (1988), B0R­ were noted in situ on a plexiglass plateo Sorne specimens GESEN (1936, 1937, 1938, 1948, 1957), GEPP & GEPP were preserved in 4% formalinj seawater for further (1911), DURAIRATNAM (1961), EGEROD (1974, 1975), study, but the majar part of the collected specimens and WOMERSLEY (1984). For more complete references was dried and pressed as herbarium reference material. on biogeographical distribution in the lndian Ocean The main collection is deposited in the herbarium of see SILVA el al. (1996). the University of Gent (GENT, Belgium, here indicated by HEC = Herbarium Eric Coppejans), except for the RESULTS Seychelles-expedition which is mainly in the Rijks­ herbarium Leiden (L, the Netherlands, here indicated by SEY). Duplicates were left in the respective research IDENTIFICATION KEY TO THE GENERA OF THE institutes (the Kenya Marine and Fisheries Research CODIALES FROM KENYA, ZANZIBAR AND THE lnstitute in Mombasa, the lnstitute of Marine Science SEYCHELLES (WESTERN ) in Zanzibar and the Fisheries in Mahé). The study of 1.a. Thallus erect, p1umose, extremely supple, compo­ the material was carried out by VAN DEN HEEDE (1994). sed of a single, branched siphonous filament ... 2 Macromorphological features (incl. growth pattern : b. Thallus composed of branched, intricated sipho- repent, ascending, erect ; branching pattern) were exa­ nous fIlaments 3 mined in both dried and preserved specimens. Ana­ tomical characters were studied in preserved specimens 2.a. Gametes formed in morphologically unmodified when available, or in resoaked herbarium fragments. ramuli Bryopsis CODIALES FROM THE WESTERN INDIAN OCEAN 49

b. Gametes farmed in specialized gametangia, sepa- sediments and therefore becoming covered by rated from the ramuli by a basal septum . substrate and colourless or brownish, forming ...... T'richosolen dense mats, partly erect and green. 3.a. Filaments loosely intricated, forming creeping Anatomy: colourless lower filaments (75-) mats, medulla and cortical utricles absent .. 90-95 (-115) Jlm in diameter; branching rare, ...... Boodleopsis dichotomous, no marked constrictions between b. Filaments intertwined and compact resulting in branching points. Green erect fIlaments (23-) 40 a spongy thallus, consisting of a medulla and a (-50) Jlm in diameter; branching di-(rarely tri-) cortex formed by utricles Codium chotomous, rather dense, slightly swollen at the dichotomies, (15-) 19 (-25) Jlm in diameter at IDENTIFICATION KEY TO THE SPECIES OF BRYOPSIS interdichotomal and infradichotomal constric­ FROM KENYA, ZANZIBAR AND THE SEYCHELLES tions. Plastids oval to rounded, 2,8-4,6 !-lm long, (WESTERN INDIAN OCEAN) each with a single pyrenoid. Gametangia oval, l.a. Ramuli of last arder in all directions resulting in rounded or obovate, (155-) 173 (-193) /-lm long three-dimensional apical parts ...... B. hypnoides and (125-) 145 (-195) Jlm wide, borne on a long b. Ramuli of last arder not in all directions, but in or short pedicel [(7-) 40 (-100) Jlm long] on the a single plane, distichous or unilateral, resulting unconstricted repent filaments. in apical plumules ; the thallus itself can be three- Ecology: collected from upper to lower dimensional 2 levels of the intertidal zone, epipelic, epipsammic 2.a. Outline of plumule triangular; pinnules on 2 dou- or epiphytic on mangrove aerial roots. ble, opposite rows B. plumosa b. Outline of the plumule not triangular but linear, Geographic distribution : originally described obovate to ovate, pinnules on 2 single or double, fram and by COLLlNS (1909b : opposite rows 3 431) as Dichotomosiphon pusillus. Cosmopolitan 3.a. Plants small, 1-3 cm, plumule oblongate to ovate, in (sub-)tropical seas. 1-3 mm broad and 2-8 mm long ...... B. indica Atlantic Ocean: North Carolina (SCHNEIDER & b. Plants larger, plumule linear-lanceolate, some- SEARLES 1991), Bermuda (TAYLOR 1960), Florida times bent 4 (DAWES 1974), Bahamas (TAYLOR 1960), Brazil 4.a. Main axis generally curved, ramuli unilateral or (TAYLOR 1960), (TAYLOR 1960), Jamaica partly distichous, but then the curvature of the (TAYLOR 1960, CHAPMAN 1961), Puerto Rico ramuli from the most dense row parallel with the (TAYLOR 1960), (TAYLOR 1960), West main axis, those from the sparsely set row in- (LAWSON & JOHN 1987). curved B. pennata var. secunda Indian Ocean: Kenya (COPPEJANS & GALLlN b. Main axis mostly straight, ramuli clearly disti­ 1989, COPPEJANS et al. 1992), Tanzania (LAW­ chous, on 2 opposite (sometimes double) rows S SON 1980), (POCOCK 1958), Seychel­ S.a. Plumule not interrupted by bare parts .. les (KALUGINA-GUTNIK A.et al. 1992), ...... B. pennata var. pennata (SHAMEEL & TANAKA 1992), (CHIHARA b. Plumule interrupted by bare parts without ra- & TANAKA 1988). muli B. pennata var. leprieurii Pacific Ocean: Philippines (FORTES & TRONO 1979, SILVA et al. 1987), Queensland, DESCRIPTION OF THE SPECIES (CRIBB 1954).

BOODLEOPSIS Gepp & Gepp Specimens examined: Kenya: HEC 6741 : 08/07/1987 - Gazi ; HEC 8607 : 17/09/1990 ­ Boodleopsis pusilla (Collins) Taylor, Joly & Ber­ Mombasa, Bamburi Bay; HEC 8762: 13/09/ natowicz Fig. 1, 5 1991 - Kanamai. Syn. : Dichotomosiphon pusillus Collins The dimensions of our specimens agree best Morphology: thallus composed of inter­ with those given by SCHNEIDER & SEARLES twined siphonous filaments, partly creeping, fixing (1991 : 89) for material from South East America 50 BELGIAN JOURNAL OF BOTANY 129

(lower fI1aments ± 90 tlm in diameter, erect 135) ; cosmopolitan in warm and temperate re­ fI1aments 23-48 tlm in diameter) ; they are coarser gions. than the West African specimens described by Atlantic Ocean : Bermuda, North Carolina, Flo­ LAWSON & JOHN (1987: 86). No other species rida, Bahamas, Cayman (TAYLDR 1960), of Boodleopsis has been mentioned in the area Jamaica (CHAPMAN 1961), studied. (HARVEY 1849, BURROWS 1991), (DEBRAY 1899, CHALDN 1905, FELDMANN 1937, COPPE­ BRYOPSIS Lamouroux JANS 1995) ; (Pignatti 1962). Indian Ocean: Kenya (IsAAc 1967), Tanzania Bryopsis hypnoides Lamouroux Fig. 2, 3, 6 (JAAsuND 1976), (FARGHALY 1980), (B0RGESEN 1946), (AL-HAsAN Morphology: thallus erect, up to 10 cm & JONES 1989), (DE CLERCK & high; branching generally very dense and in all COPPEJANS 1994, 1996), (UMAMAHESWARA directions, resulting in a three-dimensional plu­ 1969), (DURAIRATNAM 1961), mule; main axis always recognizable; light to (NIZAMUDDlN & GESSNER 1970), Pakistan (SHA­ olive-green. Two extreme growth forms (with MEEL & TANAKA 1992). intermediates) : a) extremely supple woolly tufts, Pacific Ocean: (DURAIRATNAM 1961), no well defined conoidal outline; main axis Marshall Islands (DAWSON 1957), Galapagos without a bare base; b) very well individualized Islands (TAYLDR 1945), Panama (TAYLDR 1945). specimens with a marked, bare "stipe" (6-28 mm long) and a very dense, spongy, rather regular Specimens examined: Kenya: HEC 5645 : conoidal plumule (4-30 mm in diameter at its 10/07/1985 - Mombasa, Nyali ; HEC base). The onIy specimen from the Seychelles 6087 : 03/02/1986 - Mombasa, Shelly Beach; (SEY 477) shows an extremely branched bare HEC 6861 : 15/07/1987 - Diani; HEC 6903 : main axis and small plumules of 1-8 mm in dia­ 17/07/1987 - Gazi; HEC 7017: 28/07/1987­ meter. Mombasa, Nyali Beach; HEC 7044: 29/07/ Anatomy : main axis 230-1000 (exceptionally 1987 - Kanamai; HEC 7054: 30/07/1987 ­ up to 2000) tlm in diameter. Growth form a) Diani; HEC 7175 : 25/02/1988 - Gazi; HEC woolly habit : side axes 125-330 tlm in diameter, 8193: 31/07/1989 - Gazi; HEC 8299: 09/ not constricted at the base; primary side branches 08/1989 - Gazi; HEC 8149: 21/12/1988 ­ irregularly placed or more or less on 2 longi­ Gazi; HEC 8203: 01/08/1989 - Gazi; HEC tudinal rows (one of these generally with more 8496: 24/08/1989 - Diani; HEC 8634: 20/09/ densely placed branchlets), 60-90 (-130) tlm in dia­ 1990 - Mombasa, Iwatine Bay; HEC 8670: 05/ meter, strongly constricted at their base (30­ 09/1991 - Mombasa, Mwamba Beach; HEC 45 tlm); secondary branchlets, if present, 35­ 9474: 15/09/1992 - Gazi; HEC 9406: 11/09/ 45 tlm in diameter, with basal constriction to 15­ 1992 - Mombasa, Iwatine Bay. Zanzibar: HEC 23 tlm. Growth form b) with conoidal plumule : 9916: 05/08/1993 - Paje; HEC 10479: 20/08/ side axes (60-) 100 (-155) tlm in diameter, con­ 1994 - Uroa; HEC 10576: 23/08/1994 ­ stricted at the base (30-45 tlm) ; primary branch­ Nungwi; HEC 10690: 27/08/1994 - Chwaka. lets irregularly placed, 45-90 tlm in diameter, Tanzania: HEC 11314: 18/01/1996 - Mbudya constricted at the base. Plastids oval to rounded, (Kunduchi Beach); HEC 11362: 21/01/ 3,5-10 tlm long, each with a single pyrenoid. 1996 - Misali Island (E-coast of Pemba Island). Ecology: epilithic (on coral, rocks, shells), Seychelles: SEY 477: 29/12/1992 - Poivre epiphytic (on stipes of seagrasses and on larger . algae) or epipsammic, from the upper part of the This cosmopolitan species shows a pronoun­ intertidal down to the shallow infralittoral zone. ced variation in habit and ecology. The plumules Geographic distribution : first observation in of HEC 7054 are not conoidal, they are narrowing the Mediterranean Sea by LAMOUROUX (1809 : towards the base and fixing sand between their 3 4

FIG. 1-4. - General morphology. Fig. 1. Boodleopsis pusilla (Collins) Taylor, Joly & Bernatowicz : habit of HEC 8607 (scale bar = 3 cm). Figs 2, 3. Bryopsis hypnoides Lamouroux. Fig. 2. Habit of a woolly thallus (HEC 8299) (scale bar = 3 cm). Fig. 3. Habit of thalli with a naked 'stipe' and a conoidal plumule (scale bar = 0,5 cm). Fig. 4. Bryopsis indica Gepp & Gepp : habit of HEC 6825 (scale bar = 0,5 cm). 52 BELGIAN JOURNAL OF BOTANY 129

branchlets. HEC 6087 is a sparsely branched Pacific Ocean : Taiwan (LEWIS & NORRIS 1987), specimen: primary ramuli strongly constricted Philippines (VANNAJAN & TRONO 1977, SILVA et (15 11m) at their base, placed in small groups and al. 1987), Japan (cited in CRIBB 1954), Australia­ sorne of them forming a basal rhizoidal out­ Queensland (CRIBB 1954), growth. Morphologically B. hypnoides can be (WOMERSLEY & BAILEY 1970), Marshall Islands distinguished from the other species of Bryopsis (DAWSON 1957). found in the study area, by its three-dimensional Specimens examined: Kenya: HEC 5998 : plumule. According to JAASUND (1976: 17) the 24/01/1986 - Mombasa, Me. Kenzie Point; side axes have a diameter of 100-200 11m and the HEC 6825: 13/07/1987 - Tiwi; HEC 7008 : 27/ primary ramuli are 80-100 11m in diameter. DURAI­ 07/1987 Tiwi; HEC 7301: 10/03/1988 ­ RATNAM (1961 : 26) and BURROWS (1991 : 182) Shimoni, Wasini Island; HEC 8818: 22/09/ mention the presence of rhizoidal outgrowths at 1991 - Msambweni. Zanzibar: HEC 9877: 04/ the bases of lower branches. 08/1993 - Matemwe; HEC 9917: 05/08/ 1993 - Paje. Tanzania: HEC 11336: 18/01/ Bryopsis indica Gepp & Gepp Fig. 4, 7 1996 - Mbudya Island (Kunduchi Beach); HEC Morphology: small erect thalli (1-3,5 cm 11418 : 23/O1/ 1996 Pemba Island, Vitongoji. high) growing in dense or sparse tufts ; each plant N one of the previous descriptions ofthis spe­ with a marked bare "stipe" and a small terminal, cies (GEPP & GEPP 1909 : 379 ; B0RGESEN 1940 : oblongate to ovate plumule, 1-3 (exceptionally 44; CRIBB 1954: 18; WOMERSLEY & BAILEY 4) mm broad, 2-8 mm long; main axis sometimes 1970: 271 ; SARTONI 1976: 120; VANNAJAN & branched and then bearing lateral plumules next TRONO 1977 : 45) mentions the diameter of the to the terminal one ; light to olive-green. ramuli. The habit of our specimens agrees very Anatomy: main axis 215-520 11m in dia­ well with the illustrations of GEPP & GEPP (1909 : meter, bearing scars of lost branchlets along the fig. 10) and those of CRIBB (1954: pI. 1, fig. 2). major portion of the bare part; diameter of HEC 7301 has longer (2,5 mm) and thinner eventual side axes 185-230 11m, constricted to 125­ ramuli (60-) 80 (-95) 11m; according to CRIBB 170 11m at their base; desciduous ramuli relative­ (1954: 18) these appear to be older specimens. ly short (0,5-1,5 mm), mostly on 2 opposite, lon­ B. indica is very similar to B. australis Sonder gitudinal, straight rows ; sometimes each of these where the ramuli also are on 2 double rows rows doubled in 2 rows of alternating branch­ (although less regularly placed) ; WOMERSLEY & lets ; these ramuli 100-195 11m in diameter, strong­ BAILEY (1970 : 271) suggest that both names are ly constricted at their base: (60-) 80 (-95) 11m. synonyms. The distinction with B. pennata is not Plastids rounded, oval, irregular or spindle­ always evident either: the plants of this species shaped, 4-18 11m long, each with a single pyrenoid. are generally larger, the plumules longer and the Ecology : epilithic on vertical or overhanging ramuli coarser. walls of rock pools in the intertidal or the infra­ littoral fringe. Bryopsis pennata Lamouroux varo pennata Fig. 8, 9, 12,16, 20 Geographic distribution : originally described from the in the ludian Morphology : thalli gregarious, frequently in Ocean by GEPP & GEPP (1909 : 379). dense tufts, (2-) 3-10 (-12) cm high; main axis lndian Ocean: Kenya (ISAAC 1968), generally unbranched, length of the bare part (SARTONI 1976), Mauritius (B0RGESEN 1940), ("stipe") variable; plumule linear-lanceolate, disti­ Seychelles (WYNNE 1995), Chagos Archipelago chous, 1-9 mm broad, 1,5-3 cm long; dark green, (GEPP & GEPP 1909), India (Dixit 1968), sometimes iridescent. (Hackett 1977, Wynne 1993), Sri Lanka (cited in Anatomy: diameter of the main axis in­ CRIBB 1954), (Islam 1976), creasing towards the base, 200-690 11m; ramuli (Teo & Wee 1983). with a rather constant length within one specimen CODIALES FROM THE WESTERN INDIAN OCEAN 53

b

250 J.lm

a e

d

50 J.lm

FIG. 5-7. Camera lucida drawings. Fig. 5. Boodleopsis pusilla (Collins) Taylor, Joly & Bernatowicz (HEC 6741) : a. upper filaments with marked constrictions ; b. lower filaments ; c. gametangia. Fig. 6. Bryopsis hypnoides Lamouroux : a. apical part of a woolly thallus (REC 8193), main axis with ramuli; b. apica! part of a thallus (REC 9916) with a 'stipe' and a p1umule, main axis with ramuli. Fig. 7. Bryopsis indica Gepp & Gepp : a. apical part of REC 7008; b. basal part of REC 7008, main axis with scars and 1 side axis with ramuli; c. oval plastids (REC 7301) ; d. spind1e-shaped p1astids (REC 9917). 54 BELGIAN JOURNAL OF BOTANY 129

(1-4 mm), resulting in the linear aspect of the Pacific Ocean: Philippines (SILVA et al. 1987), plumule, and with a diameter of (90-) 155-185 , (CRANE 1981) Japan (DURAIRATNAM (-295) Ilm, constricted at their base (38-75 Ilm) 1961), Marshall Islands (DAWSON 1957), and with a truncated apex. Plumule distichous, (EGEROD 1952). but position of the ramuli either on 2 opposite, Specimens examined: Kenya: HEC 5596 : single, straight rows, or on a single and a double 05/07/1985 - Mombasa, Mc. Kenzie Point; row or on 2 opposite double rows. HEC 6724: 05/07/1987 - Mombasa, Nyali Three collections (HEC 6822, HEC 7417 and Beach; HEC 6728: 07/07/1987 - Mombasa, HEC 9510) are much more slender in all respects : Nyali Beach; HEC 6773 : 12/07/1987 - Mom­ diameter ofthe main axis 195-270 (-315) Ilm, with basa, Nyali Beach; HEC 6822: 13/07/1987 ­ thickened cell wall (1,3 Ilm) in the median and Tiwi; HEC 7345: 15/03/1988 - Mombasa, basal parts; the apparently naked basal part Nyali Beach; HEC 7417 : 20/03/1988 - Malindi, provided with scars; diameter of the eventually Casuarina Point; HEC 8608: 19/09/1990 ­ developed side axes 95-140 Ilm. Ramuli more or Mombasa, Iwatine Bay; HEC 9510: 19/09/ less on 2 single longitudinal, opposite, straight 1992 - Malindi, Casuarina Point. rows, 45-75 Ilm in diameter (down to 23 Ilm at the constricted base), 0,5-1,5 mm long. They In sorne tufts plumules of the pennata-type agree with the dimensions mentioned by Lawson are mixed to the leprieurii-type (with bare parts & John (1987 : 93). in the plumule) ; the identification of the specimen Plastids rounded, oval or irregular, 3,5­ was then based on the dominant form, but this 11,5 IlID long, each one with a single pyrenoid. fact indicates that the distinction of varieties is artificial. Except for the two "slender" collections Ecology : epilithic on coral substrate in reef described aboye, the Kenyan specimens are pools, sometimes under the overhanging cliff wall coarser than those from the West African coast under the supralittoral fringe (Bostrychietum), [LAWSON & JOHN 1987 : 93 : axes 240-360 Ilm, epiphytic on seagrass stipes or even epipsammic ramuli (47-) 60-150 Ilm]. HEC 6724 is a very large in the low midlittoral zone or in the infralittoral epiphytic specimen (up to 15 cm) with strongly fringe. ramified main axes and extremely small plumules Geographic distribution: first mentioned (2 x 5 mm). around the Antilles by LAMOUROUX (1809 :134). In his monograpical study of the genus Atlantic Ocean: North Carolina - ­ Bryopsis in the Mediterranean Sea PIGNATTI Bermuda (SCHNEIDER & SEARLES 1991), Florida (1962) follows B0RGESEN (1911) in considering (DAWES 1974), Bahamas (TAYLOR 1960), B. pennata as a variety of B. plumosa [B. plumosa (SCHNEIDER & SEARLES 1991), Cuba (TAYLOR (Hudson) C. Agardh varo pennata (Lamouroux) 1960), Jamaica (CHAPMAN 1961), Puerto Rico B0RGESEN]. We follow SILVA et al. (1996) in (TAYLOR 1960), Virgin lslands (TAYLOR 1960), keeping both species separate. Guadeloupe (TAYLOR 1960), (TAYLOR 1960), Colombia (TAYLOR 1960), France (FELD­ Bryopsis pennata Lamouroux varo leprieurii MANN 1937, COPPEJANS 1993); West Africa (Kützing) Collins & Hervey Fig. 10, 13 (Lawson & John 1987). Morphology: thalli in dense tufts, 2,5-16 cm lndian Ocean: Kenya (IsAAc 1967), Seychelles high ; main axis often branched ; plumule linear­ (KALUGINA-GUTNIK et al. 1992), (BARRATT lanceolate, distichous, 2-10 mm broad, 1-3 cm et al. 1984), (ORMOND & BANAIMOON long, with short bare parts ; dark green. 1994), India (DIXIT 1968), Maldives (SIGEE 1966), Anatomy and ecology similar to vaL pennata. Sri Lanka (B0RGESEN 1936, DURAIRATNAM 1961), Pakistan (SHAMEEL & TANAKA 1992), Geographic distribution : originally described Malaysia (PHANG 1986). from the Bermuda Islands by COLLINS & HERVEY CODIALES FROM THE WESTERN INDIAN OCEAN 55

FIG. 8-11. - General morphology. Figs 8, 9. Bryopsis pennata Lamouroux varo pennata. Fig. 8. Habit of HEC 6728. Fig. 9. Habit of HEC 6773 . Fig. 10. Bryopsis pennata Lamouroux varo leprieurii (Kützing) Collins & Hervey: habit of SEY 382A. Fig. 11. Bryopsis pennata Lamouroux varo secunda (Harvey) CoI1ins & Hervey: habit of HEC 7018 (aI1 scale bars 2 cm). 56 BELGIAN JOURNAL OF BOTANY 129

(1917 : 62). Cosmopolitan in temperate and (sub)­ from the sparsely set row incurved (fig. 14 b, c), tropical seas. still resulting in an asymmetric aspect. Atlantic Ocean: Bermuda lslands (COLLINS & Ecology similar to var. pennata. HERVEY 1917, TAYLOR 1960). lndian Ocean: Kenya (ISAAC 1971), Tanzania Geographic distribution: first observation (JAASUND 1976), Réunion (PAYRI 1985), Diego around the Bermuda lslands by COLLINS & Garcia Atoll (RHYNE 1971), Yemen (ORMOND & HERVEY (1917 : 62). BANAIMOON 1994), Singapore (TEO& WEE 1983). Atlantic Ocean: Bermuda lslands (COLLINS & HERVEY 1917, TAYLOR 1960), Florida (TAYLOR Specimens examined: Kenya: HEC 7197 : 1960), West lndies (CHAPMAN 1961), Jamaica 01/03/1988 - Mombasa, Mc Kenzie Point; (CHAPMAN 1961) ; West Africa (LAWSON & JOHN HEC 9417: 12/09/1992 - Mombasa, Mc Kenzie 1987). Point. Zanzibar: HEC 10527: 21/08/1994 ­ lndian Ocean : Tanzania (JAASUND 1976), Mau­ Bawe lsland; HEC 10657 : 25/08jl994 Ras ritius (JAGTAP 1993), Atoll (RHYNE Fumba. Tanzania: HEC 11198: 11/01/1996 ­ 1971), Yemen (ORMOND & BANAIMOON 1994), Mafia lsland. Seychelles: SEY 323: 23jl2/ India (DIXIT 1968), Singapore (TEO& WEE 1983). 1992 - lsland; SEY 382: 25jl2/ Pacific Ocean: Philippines (SILVA et al. 1987), 1992 - Ile Moyenne. Marshall lslands (DAwsoN 1957). Our specimens agree well with the dimen­ Specimens examined : Kenya: HEC 5834 : sions given by JAASUND (1976: 17): main axis 13/01jl986 - Mombasa, Mc Kenzie Point; 400-500 !-lm in diameter, ramuli 3 mm long and HEC 6717: 06/07jl987 - Mombasa; HEC 180-200 !-lm in diameter. Bryopsis pennata varo 7018: 28/07/1987 - Mombasa, Nyali Beach; leprieurii differs from var. pennata as well as from HEC 7085 : 05/08/ 1987 - Lamu, Ras Kitau; all other Bryopsis-species found in the study area, HEC 8671b: 05/09/1991- Mombasa, Mwamba by the presence of short bare parts in the plumule. Beach; HEC 9490: 16/09jl992 Msambweni. Zanzibar: HEC 10732: 30/08/1994 Bawe Bryopsis pennata Lamouroux var. secunda lsland. Seychelles: SEY 357 : 1O/12jl992 - Ile (Harvey) Collins & Hervey Fig. 11, 14 Sourie, Pointe du Se!. Syn.: Bryopsis plumosa (Hudson) C. B. pennata var. secunda is very similar to Agardh var. secunda Harvey the unilateral branching form of B. australis Sonder; the dimensions are very similar too, but Morphology : thalli in small dense tufts, 2­ this species is considered as endemic for Australia. 8 cm high; main axis generally markedly in­ B. harveyana J. Agardh also has a unilateral curved; plumule 1-5 mm broad and 0,7-2 cm branching pattern; this species which has been long; dark green, sometimes iridescent. collected in Mauritius (B0RGESEN 1946 : 34) dif­ Anatomy: diameter of the main axis in­ fers by the more slender ramuli (60-100 !-lm in creasing towards the base, 177-500 !-lm, bearing diameter) being irregularly arranged in a rather scars of lost branchlets ; diameter of eventual side broad zone. axes 185-375 !-lm, constricted at their base (77­ 115 !-lm) ; ramuli 0,5-4 mm long, 90-185 !-lm in Bryopsis plumosa (Hudson) C. Ag. Fig. 15, 19 diameter (35-75 !-lm at their base). Plumules Bas. : Ulva plumosa Hudson general1y unilateral on the ventral side of the axis Syn. : Bryopsis arbuscula Lamouroux (fig. 14 a), or partly distichous with one row on the ventral side and the other row on the dorsal Morphology: thalli erect, up to 3-11 cm side of the axis (fig. 14 b, c), one of these rows high, forming dense tufts; main axes mostly with markedly more densely set ramuli ; curvature straight and unbranched; plumules triangular, of the ramuli from the most densely set row in 1,5-2,5 cm long, 10-12 mm wide at the basis, with the same direction as the axis (fig. 14 b, c), those distichous ramuli in a single plane ; light green. CODIALES FROM THE WESTERN INDIAN OCEAN 57

50 ¡.¡m

a

b

a

FIG. 12-14. Camera lucida drawings. Fig. 12. Bryopsis pennata Lamouroux var. pennata : a, b. apical part of thallus (a. HEC 6822 ; b. HEC 5596), main axis with ramuli ; c. irregular plastids (HEC 5596). Fig. 13. Bryopsis pennata Lamouroux var. leprieurii (Kützing) Collins & Hervey: a, b. apical part of thallus (a. HEC 9417; b. HEC 7197), distichous plumule with short naked parts. Fig. 14. Bryopsis pennata Lamouroux var. secunda (Harvey) Collins & Hervey: a-c. apical parts of thallus ; a. unilateral plumule (HEC 9490) ; b, c. partly distichous plumule with ramuli incurved in the same direction, from the same thallus (REC 5834). 58 BELGIAN JOURNAL OF BOTANY 129

Anatomy: diameter of the main axis in­ rial from the Boulonnais (N. France). Sorne of creasing towards the base, 345-500 ~m; ramuli these specimens also have the ramuli placed on present from near the base, on 2 double, lon­ 2 double rows, with the branchlets of a row gitudinal, straight rows, becoming longer from alternating with those of the other row of the apex to base (0,5-6 mm), thus resulting in a couple. In specimens of B. pennata var. pennata triangular plume ; ramuli 100-160 ~m in diameter, from the E. African coast we observed that the constricted at the base (38-55 ~m), with blunt position of the ramuli can either be on 2 opposite, apices. Plastids oval or rounded, 3,5-6,5 ~m long, single, straight rows, or on a single and a doub1e each with a single pyrenoid. row or on 2 opposite double rows. We therefore do not distinguish a new form or variety in B. Ecology: a single specimen, epiphytic on plumosa based on the presence of these doub1e seagrass-stolons, in the upper midlittoral. rows as the genus Bryopsis has highly variable Geographic distribution: cosmopolitan in morphological characters ; we suggest to broaden (sub)tropical and temperate seas. the description of B. plumosa as to include the Atlantic Ocean : Bermuda Islands (TAYLOR 1960), here described growth formo As the type mate­ North & South Carolina (TAYLOR 1960), Florida rial of B. plumosa is lost it is impossible to check (TAYLOR 1960), Mexico (TAYLOR 1960), Cuba which growth form was originally collected. (TAYLOR 1960), Jamaica (CHAPMAN 1961), Virgin Moreover it is not a1ways easy to recognize the Islands (TAYLOR 1960), Guadeloupe (TAYLOR placement on 2 doub1e rows on dried material. 1960), Barbados (TAYLOR 1960). United Kingdom (HARVEY 1849, BURROWS 1991), France (DEBRAY Bryopsis sp. Fig. 17,21 1899, CHALON 1905, FELDMANN 1937, COPPE­ JANS 1995), (KOSTER 1941, Pignatti 1962) ; Morphology : thallus erect, up to 2 cm high, West Africa (LAWSON & JOHN 1987). forming a sparse tuft ; main axes slightly curved, Indian Ocean: Kenya (ISAAC 1967), South unbranched, arising from repent axes ; plumules Africa (SEAGRIEF 1988), (B0RGESEN 1934), marked1y unilateral, 10-13 mm long, 2 mm broad. India (Varma 1961), Laccadive Is1ands (JAGTAP Anatomy: diameter of the main axis in­ 1987), Malaysia (PHANG & WEE 1991), Singa­ creasing towards the base, 250-445 ~m; the pore (PURCHON & ENOCH 1954), Indonesia (VER­ apparently naked basal part provided with scars HEIJ & PRUD'HOMME VAN REINE 1993), Australia of shed branchlets ; ramu1i unilateral on a double (HmsMAN et al. 1990). row, becoming shorter from base to apex (2 to Pacific Ocean: China (TSENG 1984), Taiwan 0,1 mm), 35-55 (-85) ~m in diameter, constricted (LEWIS & NORRIS 1987), Philippines (VANNAJAN at the base (23-31 ~m). Plastids irregular, 4-9 ~m & TRONO 1977, SILVA et al. 1987), Australia long, each with a single pyrenoid. (WOMERSLEY 1984). Eco10gy : a single specimen, epilithic on reef Specimen examined : Kenya: HEC 8671a: p1atform in the infralittora1 zone. 05/09/ 1991 - Mombasa, Mwamba Beach. Geographic distribution: Indian Ocean: Sey­ chelles (this paper). Bryopsis plumosa differs from the other Specimen examined: SEY 679 : 02/01/1993 ­ distichous Bryopsis species by its characteristic He Desnreufs. triangular plumule. It is generally described and illustrated as being distichous (2 single, opposite, This specimen does not correspond to any longitudinal rows of branch1ets in a single plane) described species. It is closely related to B. har­ (KOSTER 1941 : 229 as f. typica, WOMERSLEY veyana J. Agardh also presenting a unilateral 1984 : 282, BURROWS 1991 : 183, SCHNEIDER & plumule, but in this species the ramu1i are irre­ SEARLES 1991 : 95, COPPEJANS 1995 : 100). After gularly inserted in a broad zone (B0RGESEN the observation of 2 double rows of ramu1i in 1946 : 35) and they have a diameter of 80-100 ~m the E. African specimen we checked living mate- (TSENG 1984: 280; B0RGESEN 1946: 35). The CODIALES FROM THE WESTERN INDIAN OCEAN 59

15

FIG. 15-18. - General morphology. Fig. 15. Bryopis plumosa (Rudson) C. Agardh (REe 8671a) (scale bar 3 cm). Fig. 16. Bryopis pennata Lamouroux var. pennata:, slender form (REC 9510) (scale bar I cm). Fig. 17. Bryopsis sp. (SEY 679A) (scale bar I cm). Fig. 18. Trichosolen sp. (REC 5642) (scale bar I cm). 60 BELGIAN JOURNAL OF BOTANY 129 ramuli of Bryopsis sp. are more slender than solen mucronata B0rgesen, described from India, those from the unilateral B. indica Gepp & Gepp is the only known species with such large plastids f. unilateralis Cribb (75-100 /lm) and B. pennata (7-12 /lm). B0RGESEN (1948: 27) described T. varo secunda (92-187 /lm). Because of the varia­ mauritiana B0rgesen from Mauritius, but the bility of many Bryopsis species and the absence plastids are only 4-7 /lm long. of more and fertile material we do not ascertain this entity to a known species. DISCUSSION TRICHOSOLEN Boodleopsis pusilla is a tropical cosmo­ Trichosolen sp. Fig. 18,22 politan species which could be expected to be found in the area explored by USo Its absence from Morphology : thallus erect, 2-6 cm high, at­ the collection from Tanzania and the Seychelles tached to the substrate by rhizoids; main axis therefore is remarkable. mostly branched, elegant ramuli (all of equal The 59 collections of Bryopsis studied here length, 1-2,5 mm) covering the axis completely ; belong to 7 different taxa (Table 1). Those from light green. Kenya (40 collections), collected during 9 different Anatomy : main axis gradually tapering from field trips, contain 7 taxa. B. hypnoides markedly the base to the apex: 520-90 /lm; branching is the most abundant species (17 collections) in dichotomous or irregular. Ramuli irregularly this area, followed by B. pennata var. secunda placed in all directions, (20-) 23 (-27) /lm in dia­ (6 collections). Our collection contains all 5 taxa meter, constricted at the base (to ± 15 /lm), with mentioned by LAWSON (1980) in his check-list for a blunt apex. Reference specimens sterile. Plastids Kenya. with irregular outline, 7-13 /lm long, each with Field work in Zanzibar (3 field trips) resulted a single pyrenoid. in 14 Bryopsis collections, ineluding 4 taxa. Next Ecology : epilithic on a vertical rock wall of to the 3 taxa described by JAASUND (1976) for a sandy rock pool in the low midlittoral zone. Tanzania, we also collected B. indica. During the Dutch Seychelles Expedition Specimen examined: Kenya: HEC 5642: 1992-1993 (1 month), only 5 Bryopsis collections 10/07/1985 - Mombasa, Nyali Beach. were gathered, containing 4 different taxa. In their The absence of reproductive structures list of benthic marine algae from the Seychelles (which are formed laterally on the ramuli and Islands, KALUGINA-GuTNIK et al. (1992) mention have a species-characteristic morphology) ex­ only 1 species: B. pennata. They probably did eludes any identification on species leve!. Tricho- not distinguish the 3 varieties of B. pennata.

TABLE 1 Survey 01 the taxa 01 Bryopsis and their distribution

Nr Species Kenya Tanzania Seychelles 1 B. hypnoides + + + 2 B. indica + + 3 B. pennata var. leprieurii + + + 4 B. pennata var. pennata + 5 B. pennata var. secunda + + + 6 B.plumosa + 7 B. sp. + number of species 6 4 4 CODIALES FROM THE WESTERN INDIAN OCEAN 61

o o o °0 0 0 b

500Jlm

FIG. 19-22. Camera lucida drawings. Fig. 19. Bryopis plumosa (Rudson) C. Agardh (REC 8671a) : a. apical part of the thal1us ; b. plastids, each with a single pyrenoid. Fig. 20. Bryopis pennata Lamouroux varo pennata : slender form : apical part of the thal1us (REC 9510). Fig. 21. Bryopsis sp. (SEY 679a) : a. apical part of the thal1us; b. basal part of the main axis with scars of lost branchlets ; c. irregular plastids. Fig. 22. Trichosolen sp. (REC 5642) : a. apical part of the thal1us, main axis with irregularly placed ramuli ; b. irregular plastids. 62 BELGIAN JOURNAL OF BOTANY 129

Data on Bryopsis from (sub-)tropical seas rium in Leiden, to P. Goetghebeur for taxonomic are scarce, and most of the descriptions are discussions and to coworker O. De Clerck for his incomplete (not mentioning dimensions). More­ valuable help with bibliographical search. This research over, as SETCHELL & GARDNER (1920 : 157) and was sponsored by the Fonds voor Wetenschappelijk BAARDSETH (1941 : 14) already stated, sorne of Onderzoek (Flanders, Belgium) (NWO research pra­ the traditionally used morphological and ana­ ject nrs. 2.0009.92 "Systematics, ecology and biogeo­ graphy of marine organisms in the lndian Ocean" and tomical characters are variable in different de­ G.0024.96 "Systematic study of AIgae, Fungi, Plantae grees and therefore of restricted diagnostic value. and lnvertebrates of the lndo-Melanesian region"). Intermediates between "typical" growth forms, i.e. agreeing with described entities from literature, can generally been found, and sometimes different REFERENCES "varieties" are present in a single tuft, especially in B. pennata. AL-HASAN R. & JONES W., 1989 - Marine algal flora Compared to the Mediterranean region and seagrasses of the coast of Kuwait. J. Univ. (PIGNATTI, 1962: 13 species if B. pennata is Kuwait (Sci.j 16 : 289-341. considered different from B. plumosa), the stu­ BAARDSETH E., 1941. - The Marine AIgae of Tristan died area apears to be poor in Bryopsis species da Cunha. Del Norsk. Videnskaps-Acad., . (4 species of which one with 3 "varieties"). Other 9:171p. tropical regions are similarly species poor: W. BARRATT L., ORMOND R., CAMPBELL A., HISCOCK S., HOGARTH P. & TAYLOR J., 1984. - Eco­ Africa (LAWSON & JOHN 1987): 3 species of logical study of rocky shores on the south coast which B. pennata with 2 varieties, Florida of Oman. Rep. lnt. Un. Cons. Nature & Nat. (TAYLOR 1967): 4 species of which B. pennata Res. UNEP, Geneva. 102 p. with 2 varieties, tropical E. America (TAYLOR B0RGESEN F., 1911. Sorne Chlorophyceae fram the 1960) : 5 species ; Philippines (SILVA et al. 1987) : Danish West lndies. BOl. Tidsskr. 31 : 127-152. 4 species of which B. pennata with 2 varieties. B0RGESEN F., 1934. Sorne marine algae from the Trichosolen had not previously been collected northern part of the Arabian Sea with remarks in this area; as the specimens are sterile, they on their geographical distribution. Del. Kg!. could not be identified at species leve!. Dansk. Vidensk. Selsk., Bio!. Medd. 11 : 1-72. Together with the 14 Codium species de­ B0RGESEN F., 1936. - Sorne marine algae from scribed in VAN DEN HEEDE & COPPEJANS (1996) Ceylon. Ceylon J. Sci. 12 : 57-96. B0RGESEN F., 1937a. - Contributions to a South the total number of taxa of Codiales for the study lndian marine algal flora I. Indian Bol. Soco 16 : area is 23. 1-57. B0RGESEN F., 1937b. - Contributions to a South lndian marine algal flora II. Indian Bol. Soco ACKNOWLEDGEMENTS 16 : 311-357. B0RGESEN F., 1938. - Contributions to a South We would like to thank the directors of the lndian marine algal flora lII. Indian BOl. Soco Kenya Marine and Fisheries Research lnstitute (Mom­ 17 : 206-242. basa, Kenya: MI. Allela and MI. Okemwa) and of B0RGESEN F., 1940. - Sorne marine algae from the lnstitute of Marine Sciences (Zanzibar, Tanzania: Mauritius, I. Chloraphyceae. Del. Kg!. Dansk. MI. Ngoile) for their help with the accomodation. Vidensk. Selsk., Bio!. Medd. 15: 1-84. Our gratitude also goes to E. Martens and Y. Ver­ B0RGESEN F., 1946. - Sorne marine algae fram Mau­ meulen for the local organisation of the field work ritius, an additional list of species to Part I. and laboratory research in Kenya and to M. Rich­ Chlorophyceae. Del. Kg!. Dansk. Vidensk. mond in Tanzania. The second author acknowledges Selsk., Bio!. Medd. 20: 1-64. the Netherlands Marine Research Foundation to join B0RGESEN F., 1948. - Sorne marine algae from Mau­ the Seychelles Expedition. Thanks also to W. F. ritius, additional lists to the Chlorophyceae and Prud'homme van Reine and E. Verheij for organizing Phaeophyceae. Del. KgI. Dansk. Vidensk. Selsk., and helping us during our stays at the Rijksherba- Biol. Medd. 10: 1-55. CODIALES FROM THE WESTERN INDIAN OCEAN 63

BORGESEN F, 1957. - Sorne marine algae from Mau­ Nord de la Franee. Bul!. Sci. France et Belgique ritius, final parto Det. Kg!. Dansk. Vidensk. 32 : 1-193. Selsk., Bio!. Medd. 23 : 1-35. DE CLERCK O. & COPPEJANS E., 1994. - Status of BURROWS E. M., 1991. - Seaweeds ofthe British Isles. the macroalgae and seagrass vegetation after the 2 Chlorophyta : 238 p. Natural History Museum, 1991 Gu1f War oi1 spilI. Courier Forsch.-Inst. London. Senckenberg 166 : 18-21. CHALON J., 1905. - Liste des AIgues Marines: 259 p. DE CLERCK O. & COPPEJANS E., 1996. - The marine Imprimerie J. E. Buschmann, Anvers. a1gae of the Wi1dlife Sanetuary for the Gulf CHAPMAN v., 1961. The marine algae of Jamaica. Region. In : KRUPP F, ABUZINADA A. & NADER Part 1. Myxophyceae and Ch1orophyceae. Bul!. 1. (eds.). A marine wildlife sanetuary for the Inst. Jamaica, Sci. Ser. 12: 1-159. Arabian Gulf. Environmental research and con­ servation following the 1991 Oil Spi1I. CHIHARA M. & TANAKA J., 1988. - Macroalgae in Courier Forsch.-Inst. Senckenberg Spec. vol.: Indonesian Mangrove Forests. Bull. Nat!. Sci. Mus. Tokyo Ser. B 14 : 93-106. 200-289. DIXIT S., 1968. Speeies 1ist of Indian Marine COLLlNS F, 1909. - The green algae of Bermuda. a1gae n. J. Univ. Bombay 36 : 9-24 Proc. Amer. Acad. Arts 37 : 229-270. DURAIRATNAM M., 1961. - Contribution to the Study COLLlNS & HERVEY A., 1917. - The algae of Ber­ F of the Marine AIgae of Ceylon. Fish. Res. St., muda. Contributions from the Bermuda Bio­ Ceylon Bull. 10: 1-181. logical Station for Research 69. Proc. Amer. EGEROD L., 1952. - An ana1ysis of the Siphonous 53: 1-195. Acad. Arts Ch10rophyeophyta with specia1 referenee to the COPPEJANS E., 1993. - Iconographie d'AIgues Médi­ Siphonocladales, Siphonales and Dasycladales terranéennes. Bibliotheca phycologica 63 : 317 pI. of Hawaü. University of California Press, Berke­ Cramer. ley and Los Angeles 25 : 325-454. COPPEJANS E., 1995. - Flore algologique des cotes EGEROD L., 1974. - Report of the Marine A1gae du Nord de la France et de la Belgique : 454 p. Colleeted on the Fifth Thai-Danish Expedition Jardin Botanique National de Belgique, Meise. of 1966, Ch10rophyeeae and Phaeophyeeae. Bot. COPPEJANS E. & GALLIN E., 1989. - Macroalgae Mar. 17: 130-157. associated with the mangrove vegetation of Gazi EGEROD L., 1975. - Marine AIgae of the Andaman Bay (Kenya). Bull. Soco Roy. Bot. Belg. 122: Sea Coast of Thai1and: Chlorophyceae. Bot. 47-60. Mar. 18: 41-66. COPPEJANS E., BEECKMAN H. & DE WIT M., 1992.­ FARGHALY M., 1980. - A1gues benthiques de la Mer The seagrass and associated macroalgal vegeta­ Rouge et du bassin occidental de l'Océan Indien tion of Gazi Bay (Kenya). Hydrobiologia 247 : (étude taxinomique et essai de répartition, no­ 59-75. tamment des Udotéacées) : 274 + 25 p. Thesis COPPEJANS E., KOOISTRA W. & AUDlFFRED P., Univ. Sci. & Techn. Langedoc, Montpellier. 1994. - Preliminary report on the research on FELDMANN J., 1937. - Les Cyanophycées, Chloro­ macroalgae. In: VAN DER LAND J. (ed.). Oceanic phycées et Phéophycées de la Cote des A1beres : reefs of the Seychelles, 2: 157-182. National 189 p. Imprimerie Wolf Rouen. Museum of Natural History, Leiden. FORTES M. & TRONO G., 1979. - Marine algal CRANE P., 1981. The marine Chlorophyceae and microphytes new to the Philippines. Kalikasan, Phaeophyceae of Penang Island. Malayan Nat. Philipp. J. Bio!. 8 : 51-68. J. 34 : 143-169. GEPP A. & GEPP E., 1909. - Marine algae (Chloro­ CRIBB A., 1954. Reeords of Marine A1gae from phyceae and Phaeophyceae) and marine phanero­ South-Eastern Queensland I: 15-37. Watson gams of the "Sealark" Expedition, collected by Ferguson & Company, Brisbane. J. Stan1ey Gardiner, M. A., F R. S., F L. S. ­ DAWES c., 1974. - Marine A1gae of the west coast Trans. Linn. Soco London 11 : 373-398. of Florida: 201 p. University of Miami Press GEPP A. & GEPP E. , 1911. - The Codiaceae of the Florida. Siboga Expedition including a Monograph of DAWSON Y, 1957. - An annotated list of marine algae Flabellarieae and Udoteae: 150 p. Drukkerij from Eniwetok Atoll, Marshall Islands. Pacific Brill, Leiden. Sci. 11 : 92-132. HACKETT H., 1977. - Marine a1gae known from the DEBRAY F, 1899. - F10ru1e des A1gues Marines du Maldive Islands. Atoll Res. Bull. 210 : 30 p. 64 BELGIAN JOURNAL OF BOTANY 129

HARVEY W., 1849. - A manual of the Britisch marine NIZAMUDDIN M. & GESSNER F, 1970. - The marine algae : 245 p. London, J. Van Voorst, Paternoster algae of the northern part of the Arabian Sea Row. and of the Persian Gulf. "Meteor" Forsch. Er­ HurSMAN J., KENDRICK G., WALKER D. & COUTÉ A, gebnisse 6 : 1-42. 1990. - The marine algae of Shark Bay, ORMOND R. & BANAIMOON S., 1994. Ecology of inter­ Western Australia. In: BERRY P, BRADSHAW tidal macroalgal assemblages on the Hadramout S. & WILSON B. (eds.). Research in Shark Bay. coast of southern Yemen, an area of seasonal Report on the France-Australe Bicentenary Ex­ . Mar. Ecol. Progr. Ser. 105: 105-120. pedition Cornmittee. W. Australian Museum: PAYRI c., 1985. Contribution to the knowledge of 89-100. the marine benthic flora of La Réunion Island ISAAC W., 1967. Marine botany of the Kenya Coast. (Mascareignes Archipelago, Indian ocean). In: 1. A first list of Kenya marine algae. J. E. Aji-ica Proc. 5th Int. Congr. Tahiti. Moo­ Nat. Hist. Soco Natl. Mus. 26 : 75-83. rea: Antenne Museum-E.P.H.E. 6 : 638-640. ISAAc W., 1968. - Marine botany ofthe Kenya Coast. PHANG S., 1986. - Malaysia's seaweed flora. In: 2. A second list of Kenya marine algae. J. E. WONG T, ONG J. & ONG K. (eds.). Towards Aji-ica Nat. Hist. Soco Natl. Mus. 27 : 1-28. greater productivity of our coastal waters. Proc. ISAAc W., 1971. - Marine botany of the Kenya Coast. 9th ann. sem. Mal. Soco Mar. Sd. : 17-45. 5. A third list of Kenya marine algae. J. E. Africa PHANG S. & WEE Y, 1991. - Benthic marine algae. Nat. Hist. Soco Natl. Mus. 28: 1-23. In: KIEW R. (ed.). The state of nature conser­ ISLAM A, 1976. - Contribution to the study of the vation in Malaysia: 51-61. Kuala Lumpur, marine algae of Bangladesh. Bibiotheca Phyco­ Malayan Nature Soco logica 19 : 253 p. PIGNATTI S., 1962. - Le specie Mediterranee del JAASUND E., 1976. Intertidal seaweeds in Tanzania, genere Bryopsis (Chlorophyceae-Siphonales). a field guide : 159 p. Univ. of Troms0. Atti 1st. Veneto Sd. Lett. Arti 120 : 31-58. JAGTAP T, 1987. - Distribution of algae, seagrasses PURCl-ION R. & ENOCH l., 1954. - Zonation of the and coral cornmunities from Is­ marine fauna and flora on a rocky shore near lands, eastern Arabian Sea. Ind. J. Mar. Sd. Singapore. Bull. Raffles Mus. 25: 47-65. 16 : 256-260. POCOCK M., 1958. - Preliminary list of marine algae JAGTAP T, 1993. - Studies on littoral and sublittoral. collected at Inhaca and on the neighbouring macrophytes around the Mauritius coast. Atoll . In: MACNAE W. & KALK M. (eds.). Res. Bull. 382 : 10 p. + 3 fig. A natural history of Inhaca Island, Mo<;ambi­ KALUGINA-GUTNIK A., PERESTENKO L. & TITLYANOVA que: 23-27. Johannesburg. Witwatersrand Univ. T, 1992. - Species composition, distribution Press. and abundance of algae and seagrasses of the RHYNE c., 1971. - Marine algae of Diego Garcia. Seychelles Islands. Atoll Res. Bull. 365: 1-23. In: STODDART D. & TAYLOR J. (eds.). Geo­ KOSTER J., 1941.- Quelques observations sur les Bryop­ graphy and ecology of Diego Garcia Atoll, sis du Golfe de Naples. Blumea 4: 225-258. Chagos Archipelago. Atoll Res. Bull. 149: 41-65. LAMOUROUX M., 1809. - Mémoire sur trois nouveaux SARTONI G., 1976. - Researches on the coast of genres de la famille des Algues marines. J. Bot. Somalia, the shore and dune of Sar Uanle, a 2: 129-136. study of the benthonic algal flora. Ital. J. Zool. LAWSON G., 1980. A check-list of East African 4: 115-143. seaweeds ( to Tanzania): 65 p. Dept. SCHNElDER C. & SEARLES R., 1991. - Seaweeds of Biol. Sci. Univ. Lagos, . the southeastern United States : Cape Hatteras LAWSON G. & JOHN D., 1987. - The marine algae to Cape Canaveral: 553 p. Duke Univ. Press, and coastal environment of tropical West Africa. USA Nova Hedwigia Beih. 93: 1-415. SEAGRIEF S., 1988. - Marine Algae. In : LUBKE R., LEWIS J. & NORRIS J., 1987. -A history and an­ GESS F & BRUTON M. (eds.). A field guide to notated acount of the benthic marine algae of the Eastern Cape coast: 35-72. Grahamstown. Taiwan: 38 p. Smithsonian Institution Press, Wildlife Soco RSA Washington D. C. SETCHELL W. & GARDNER N., 1920. - The Marine MOORJANI S. & SIMPSON B., 1988.- Seaweeds of the Algae of the Pacific Coast of . Kenyan coast: 285 p. Oxford Univ. Press, Part Il. Chlorophyceae. Univ. Caltl Publ. Bot. . 8: 139-374. CODIALES FROM THE WESTERN INDIAN OCEAN 65

SHAMEEL M. & TANAKA J., 1992. preliminary algae in the reference collections of the Central check-list of marine algae from the coast and Marine Fisheries Research Institute. Bull. Cena". inshore waters of Pakistan. In : NAKAIKE T. & Mar. Fish. Res. Inst. 9 : 37-48. MALIK S. (eds.). Cryptogamic Flora of Pakistan VANNAJAN S. & TRONO G., 1977. The marine 1: 1-64. : Nat. Sci. Mus.; Islamabad: benthic algae of Manila Bay, 1. lntroduction. Pakist. Mus. Nat. Hist. Cyanophyta and Chlorophyta. Philipp. J. Biol. SIGEE D., 1966. - Preliminary account of the land 6: 33-46. and marine vegetation of Addu Atoll. In : STOD­ VAN DEN HEEDE c., 1994. De Codiales (Ch10ro- DART D. Reef studies at Addu Atoll, phyta) van Kenya, Zanzibar en de Sychellen: Maldive lslands. Preliminary results of an ex­ 160 p. + 73 pI. MSc thesis University Gent, pedlltlcm to Addu Atoll in 1964. Atoll Res. Bull. VAN DEN HEEDE C. & COPPEJANS E., 1996. - The 116: 61-74. SIGEE 1966. genus Codiwn (Chlorophyta, Codiales) from SILVA P. c., BASSON P. W. & MOE R. L., 1996. ­ Kenya, Zanzibar and the Seychelles. Nova Hed­ Cata10g of the benthic marine algae of the ludian wigia 62 : 389-417. Ocean. Univ. Calif. Pub!. Bot. 79 : 1259 p. VARMA R., 1961. Flora of the pearl beds off Tuti- SILVA P., MEÑEZ E. & MOE R., 1987. Catalog of corin. J. Mar. Bio!. Ass. India 2 : 221-225. the Benthic Marine of the Philippines. VERHEIJ E. & PRUD'HOMME VAN REINE W., 1993. Smiths. Contr. Mar. Sci. 27 : 1-179. Seaweeds of the Spermonde Archipelago, SW TAYLOR W., 1945. Pacific Marine algae of the Allan Sulawesi, Indonesia. Blumea 37: 385-510. Hancock Expeditions to the Galapagos lslands : WOMERSLEY H., 1984. - The marine benthic flora of 528 p. University of South California Press, Los southern Australia, Part 1 : 329 p. South Austra­ Angeles. lian government Printing Division, Adela."ide. TAYLOR W., 1960. Marine algae of the eastern tro- WOMERSLEY H. & BAILEY A., 1970. - Marine Algae and subtropical coasts of the Americas. of the Solomon Islands. Phi/os. Trans. 259: University al Michigan Studies Scient(fic Series 257-352. 21 : 825 p. WYNNE M. 1993. - Benthic marine algae from the TAYLOR W., 1967. - The marine algae of Florida with Maldives, lndian Ocean, collected during the special reference to the Tortugas. Bibliotheca Te Vega Expedition. Contr. Univ. Michigan Phycologica 2 : 219 p. + 37 pI. Herb. 19 : 5-30. TEO L. & WEE Y, 1983. - Seaweeds of Singapore : WYNNE M. 1995. - Benthic marine algae from the 123 p. Singapore Univ. Press. Seychelles collected during the Te Vega TSENG c., 1984. - Common Seaweeds of China: lndian Ocean Expedition. Contr. Univ. Michigan 316 p. Inst. of Oceanology, Academia Sinica, Herb. 20 : 261-346. Qingdao, China. UMAMAHESWARA R., 1969. - Catalogue of marine Revised manuscript received 24 September 1996.