Phytogeography of the Fucales and Their Seasonal Growth MOHAMMED NIZAMUDDIN Institut Für Meereskunde, Kiel*} (Received: 18.12.1969}

Nizamuddin: Phytogcography of the Fucales and their Seasona] Growth 131

Botanica Marina Vol. XIII, p. 131—139, 1970

Phytogeography of the Fucales and their Seasonal Growth MOHAMMED NIZAMUDDIN Institut für Meereskunde, Kiel*} (Received: 18.12.1969}

L Introduction New Zealand inhabiting far North and fewer in the The order Fucales KYLIN emended PETROV comprises south of Cook Strait. to date 36 genera and six families. All the genera are Landsburgia comprises two species, L. wyricaeforwts }> confined to littoral and sublittoral regions. Cysioseira AGARDH and L. qmrcifotia (HOOKER et HARVEY) HARVEY C. AGARDH, Sargassum C. AGARDH are widely distrib- which are endemic to New Zealand including Chatham uted and Titrbinaria LAMOUROUX is confined to trop- Island. Marginaricl/a is also a small genus comprising two ical and sub-tropical regions. Other genera are re- species, M. boryana (A. RICHARD) TANDY and Äf. stricted in their geographical distribution. Most of the urmlliana (A. RICHARD) TANDY, They grow off rocky Fucalean genera and three families — Himanthaliaceae, coast and in sheltered pools in sub-littoral regions. Hormosiraceae and Seirococcaceae are limited to The former species is a deep water alga whereas the small regions. Such Isolation and restriction of the latter one is of shallow watet (LINDAUER et al, 1961). families and genera or few species to a particular place There are two genera, Horwosira (ENDLICHER) MENE- are of considerable biogeographical interest. GHINI and Xiphophora MONTAGNE, which are confined The World distribution of the Fucalean families and to Australia and New Zealand. Among the two species genera are shown in figs. l — 3. The known distribution of Xiphophora^ X. chondrophylia (R, BRÖ\VN ex TURNER) of the genera may be extended by collecting from other MONTAGNE ex HARVEY is confined to southern coast of regions* The summary of their distribution may give a Australia and New Zealand whereas X. gladiata (LA- correct picture of their biogeography. Even there are BILLARDIERE) MONTAGNE ex KJELLMAN is confined to regions that require a careful observation and recording south-eastern coast of Australia. This species varies before a complete picture of biogeography is obtained. with age and in different habitats. Both the species of The distribution given may mean that particular genus Xiphophora are sub-littoral* Hormos/ra is monotypic or species grow in suitable ecological habitats within and essentially intertidal (WOMERSLEY, 1959) and the regions. represented by H. banksü (TURNER) DECAISNE. BERG- Here an attempt has been made to catalogue the re- QUIST (1959) made an extensive study on the ecology of ported occurrences and distribution of all the families this species. and their genera to see whether a definite pattern of In Australien waters Cystophora J. AGARDH is the largest biogeographical control is evident. Further work may genus in the Cystoseiraceae and comprises 23 species distort or make this pattern more clear. The biogeog- and the biogeography of which has already been desc- raphy of the families and genera are grouped in three ribed by WOMERSLEY (1964), Almost all the species of major regions: — (i) distribution in southern hemi- this genus except C. torulosa (R. BROWN ex TURNER) sphere (ii) distribution in norhern hemisphere and J. AGARDH are sub-littoral; some occur under rough (in) distribution in both hemispheres. conditions while others in sheltered waters. Only a few species of this genus also grow in New Zealand C. platylobittw (MERTENS) J. AGARDH, C. torulosa, C. II. Distribution in Southern Hemisphere congesta WOMERSLEY and NIZAMUDDIN and C. retroflexa The genera Carpophyllum GREVILLE, Marginartella TANDY (LABILLARDIERE) J. AGARDH whereas C. scalaris J. and Landsburgia HARVEY are endemic to New Zealand AGARDH and C. distenta J. AGARDH are endemic to inhabiting sub-littoral region. Among five species of New Zealand. Carpöphyllum^ one species C. scalare SUHRING is con- The genus Acrocarpia ARESCHOUG comprises two spe- fined to South Africa and abundant at Port Alfred and cies, A. paniculata (TURNER) ARESCHOUG and A. Dwsea (IsAAC, 1953) and is sub-littoral. This is a robusta (J. AGARDH) WOMERSLEY. The genus is endemic unique species in this genus and to my opinion it to Australian waters. A. paniculata extends from Cape hardly resembles New Zealand species. A detailed Spencer, South Australia> around the south east coast study is needed to determine its systematic position. C. (including Tasmania) to Port Stephens, N, S. W, angustifolium J. AGARDH, C. maschalocarpum (TURNER) (WOMERSLEY, 1964; NIZAMUDDIN, 1960). A. robusta is GREVILLE, C. ßexuosum (£SPER) GREVILLE and C. known only from the southern part of Western (A, RICHARD) J. AGARDH are endemic to Austraüa. *) Department of Botany, University of Karachi, Karachi-32, The genus Caulocystis ARESCHOUG is endemic to Austra- Pakistan* lian waters and comprises two species, C. mifera (C.

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\ ATLANTIC OCEAN INDIAN OCEAN PACIFIC OCEAN ANTARCTIC \ PacificCout ^ \ sf America vi * M. \ 1 AiabianSea^ ^ 5 1 ΐ i 1 1 ™« «y 1U «3 ¹ *" «Elbt c Crt ÊË^ c — 5ί τ/ if c ™ Ë ·™ ^ i— ^ c -^ ^ ^i c 32 ^ ^ i^l^^wn^C'3g^^m^ 111 Adriati c Sc a GENERA \ Mcditrranca n 51!

Acrocarpia + 4- + 4- 4- + Acystis + Ascophyllum 4- Axillariella + Bifurcaria 4- + •h Bifurcariopsis + Carpoglossum + + Ψ

Carpophyllum ·*· 4· 4- Caulocystis 4- + 4- 4- + Coccopbora 4- + 4- + 4- 4- + Cystopbora + + 4· Cystophyllurn Cystoscira + 4- 4- 4- + 4 4- 4- 4· + -*· 4- Cystosphaera 4- Fucus + 4· 4- + + + 4· Halidrys -1-4 + •1- Hesperophycus 4- Himanthalia 4- 4- -h + 4 Kizikia 4-4- 4- Hormophysa 44+ 4-4 Hormos-ira 4- + 4- 4- 4- 4- Landsburgia 4- Marginarielto 4· 4- Myriodcsma -*- + 4- +· ?elvetia + 4- 4· Pelvetiopsis + Phyllospora + + 4- 4· Platythalia

Sargassum T-t" T-^* T4- ^T ^-U· TJ^ ~ér Scabena 4· + 4- + Scylolhalia 4- 4- Seirococcus + 4-4 Stokeyia + +· Stolonophora 4- W Turbinaria + 4- 4- 4- + 4- 4- 4- + + + + 4· H- 4- Xiphophora 4- -h + H- 4- 4-

Tab. l Distribution of the fucalean genera in the World

Fig. 2 Distribution of the fucalean families in the world, showing densities

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c. £·

Fig. 3 Distribution of the fucalean genera shown along the coast of the world (for detail see the list of the abbrevi- 6fl ation of the genera) a) Distribution of the genera alontj the Pacific coast b) Distribution of the genera along the north-east coast of Asia c) Distribution of the genera in Australian waters

A-l = Acrocarpia H-4 Hizikia A-2 =* Acystis H-5 Hormophysn A-3 = Ascophyüum H-6 Hormosira A-4 = Axillar iella L Landsburgia B-I - Bifurcaria M*l Marginariella B-2 = Bifurcariopsis /Vi-2 Mvriodesma C-l = Carpoglossam p. l C-2 — Carpophyllum P-2 Pelveiiopsis C-3 = Caalocystis P-3 Phyllospora C*4 = Coccophora PiatythQlia C-5 = Cystophora S- 1 Sargassum C-6 = Cystophyllum S-2 Scaberia C-7 = Cystoseira S-3 Scytöthalia C-8 = Cystosphaera S-4 Seirococcus F = FUCLTS S-5 Stokeyia H-1 - Haliärys S-6 Stolonophora H-2 = Hesperophycus T Turbinnria H-3 = Himanthalia X Xiphophora

32 -

Flg. 3a

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20 N

100 110 120 130 150 Fig. 3b

t i f 1

«'S,

- Ë .

11t) 120 130 ISO 160 170 190

Fig. 3c

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AGARDH) ARESCH. and C. ctphalornithos (LABILLARDIERE) folium SONDER and P. qtitrcifolium (TURNER) SONDER. ARESCH, These two species are separated mainiy on the Each inhabit sub-littoral region. shape of the vesicles. They inhabit generally calm to Scabtria GREVILLE is a unique fucalean member which medium rough coasts, from low tide level down but is monotypic and endemic to Australia extending from sometimes they may occur in pools. C. wifera extends Geraldton Bay, Western Australia, along the south east from Shark Bay, W, Australia around southern Austra- coast to Bondi, N.S.W., including Kangaroo Island and lia (including Tasmania) to Long Bay, N. S- W. C. northern coast of Tasmania and also in Lord Howe cephalornithos is distributed from Elision, S- Australia, Island. It is represented ^S.agardhii GREVILLE which around south eastern coast to Bondi, N. S- W., includ- grows in littoral to sub-littoral region (NIZAMUDDIN, ing Tasmania. 1962). Carpoglos$t4m KÜTZING i s monotypic and represented by Antarctica and sub-Antarctica are represented by a single one species, C confluens (R. BROWN ex TURNER) KÜTZING genus, Cystosphatra SKOTTSBERG which is endemic äs in Australian waters extending from Eliston, South well äs monotypic, represented by C. jacqtiinotn SKOTTS- Australia, around south eastern coast (including Tas- BERG. This is confined to West Antarctica growing along mania) to Western Port, Victoria. This species inhabits the steep rocks, It occurs in Bransfield Strait Harmony sub-littoral region (NIZAMUDDIN, I960, 1964). Cove on Nelson Island. Aiyrhdesma DECAISNE extraaustralian genus comprises South Africa is represented by two monotypic and to date eight species which are endemic to Australian endemic genera, Axillaridla (GRÜBER) SILVA and Bifur- watcrs extending from Western Australia around cariopsis PAPENFUSS, A. consfricta (GRÜBER) SILVA and southern Australia including Tasmania to Victoria B. capensis (ARESCHOUG) PAPENFUSS grow along the (NIZAMUDDIN and WOMERSLEY, 1966). M. harveyanum south western corner of the Cape Province (!SAAC, 1942. NFZAMUDDIN and WOMERSLEY is confined to South STEPHENSON, 1947), In addition to the above species Australia including Kangaroo Island, M. leptophyllum two more species, Bifurcaria brasstcaeforwis (KÜTZING) J. AGARDH and AL tuberosttm J. AGARDH are South BARTON and Carpophyllntn scalare SUHRING grow along Australian in origin extending from Eucla, Western the coast of South Africa. The ecology of these two Australia, to Port Elliot, South Australia* Eucla is just species have already been described with the other on the border of Western Australia and South Australia. species of the genus. It is more likely that the specimen found at Eucla may have drifted to this place. M. integrifolium HARVEY is distributed from Western Australia (Cottesloe) to » Distribution in Northern Hemisphere Victoria (Western Port) including north coast of Tas- Japan is represented by two endemic genera, Coccophora mania. AL quercifolium ( ) J. AGARDH is distributed GREVILLE and Hi^ikia OKAMURA which are monotypic from Geraldton, Western Australia» to Port Elliot. äs well, C. langsdorfii (TURNER) GREVILLE and //, fusi- South Australia (NIZAMUDDIN and WOMERSLEY, 1966), formis OKAMURA inhabit sub-littoral region. The distri- Af. serrtilatum (LAMOUROUX) DECAISNE is confined to bution of //- fusiformis extends upto Hong Kong. Both Western Australia (Dongara to Cape Riche). The other the species have also been reported from Korean coast west Australian species, M- peronii NIZAMUDDIN and (KANG, 1966). WOMERSLEY is distributed from DONGARA to MANDURAH. Along the coast of North America three endemic genera M. callophyllum J, AGARDH inhabit south eastern coast — Hesperopfrycus SETCHELL et GA R DNER, Pelvettopsis of Australia (Fishery Bay> S. A. to Port Philips, Vict). GARDNER and Stolonophora NIZAMUDDIN — gtow. He- Phyllospora C AGARDH, Scytothalia GREVILLE and sperophycus harveyanus (DECAISNE) SETCH. et GARDN. is Seirococcus GREVILLE are endemic to Australian waters the only species of the genus which grows in the upper and each genus is monotypic. They grow in sub-Üttoral littoral region and remains exposed to atmosphere du- region. P. comosa (LABILLARDIERE) C- AGARDH is ring low tide time, It is distributed from Central Cali- distributed from Encounter Bay, South Australia fornia (Montrey Bay) to Ensenada, Baja California (including Tasmania), around south east coast to Port (DAWSON 1945, 1946. SETCHELL et GARDNER, 1925). Macquarie, N. S. W. Scytothalia dorycarpa (TURNER) Pelvetiopsis is distributed from British Columbia to Cen- GREVILLE extends from Rottnest Island, Western tral California (Point Carmel). The genus comprises Australia, along south coast including Kangaroo Island two species, P. limitata GARDNER and P. arborescence to Port Elliot, South Australia. Seirococctts axillaris GARDNER and they grow in the upper littoral region. (R. BROWN ex TURNER) GREVILLE is distributed from Sfolonöphora is monotypic and represented by S. bran- Fishery Bay, South Australia, to Cape Paterson, Vic- dtgeti (SETCHELL et GARDNER) NIZAMUDDIN which is toria including Tasmania. Scy. dorycarpa is more south sublittoral and endemic to Guadalupe island, Mexico western in distribution whereas P. comosa and S. axil- (Pacific side). laris are south eastern in distribution. Species of Bifurcaria STACKHOUS, Fmm LINNAEUS, Ha- Platythalia SONDER is endemic to Western Australia lidrys LYNGBYE and Pelvetia DECAISNE et THURET are extending from Dongara to Cave Islet, Recherche Archi- widely separated and isolated from their point of origin, pelago. The genus comprises two species, P. angusti- A number of species and varieties were assigned to the

Botanica Marina / Vol / 1970 / Fase- 2 Brought to you by | Universidad Nacional Autonoma Authenticated Download Date | 2/12/19 7:03 PM 136 Nizamuddin: Phytogeography of the Fucales and their Seasonal Growth well known genus Fucus. POWELL (1963) delimited a Himanthalia LYNGBYE is monotypic and extends from number of species and varieties of this genus and only mid- to sub-littoral (BURROWS, 1964, DIXON, 1963. on the basis of ecological aspects he recognised onlysix JONES, 1960. MCALLISTER et ., 1967). The genus is species, According to BURROWS (1964) and DIXON (1963) represented by H. thngata (LiNN.) S. F. GRAY and FUCHS species extend from supra-üttoral (Shetland) to belongs to Western Europe extending from southern sub-littoral region. The distribution of the genus along part of Polar region to Spain including Iceland and the the European coast of Atlantic Ocean is from upper- Faeroes in the north, Its occurrence along the coast of to sub-littoral. F. dtsttchus LINN. with its two sub~ North America is doubtful äs it has not been reported species, is distributed along the Pacific coast of North since 1875. America with its southern limit, Santa Barbara, Cali- Acystis SCHIFFNER is monotypic and endemic to Aden fornia. F. virsoides J. AGARDH is endemic to the Adriatic (PAPENFUSS, 1969)- A. heinii SCHIFFNER grows in the Sea and grows in the littoral regions. F. ceranoides LINN., sub-littoral region. F. serratns LINN., F. spiralis LINN., and F. vesiculosus Sfokeyta THIVY and DOSHI is very much restricted in LINN, are common to the north temperate latitudes and distribution like Acystis. Stokeyia is monotypic and littoral in habit (Mc ALLISTER> 1967), F. vestculosus and endemic to the northern Arabian Sea. S. indica THIVY F. serratns grow in the Baltic Sea but F. vesiculosus ex- and Dosm grows from lower to sub-littoral region. tends upto Firdand. The southern limit of the genus is Plants growing m pools are strongly calcified whereas Morocco (GESSNERJ 1965). growing in other habitats lack calcification. This species Bifurcaria STACKHOUS comprises three species and each is common along the coast of KarachL grows in the lower littoral region, B. brassicaeforwis (KÜTZING) BARTON is endemic to south west of Cape . Distribution in Both Hemispheres Province from Cape Town to Cape Agulhas (I$AAC> The genus Turbinaria LAMOUROUX is an important alga 1953)* This species belongs to the Southern hemisphere. of reef floras of the tropics and sub-ttopics, It grows B. galapagensh (PiccoNE et GRUNOXV in PICCONE) Wo- in the intertidal äs well äs in the interatidal reefs (TAY- MERSLEY is endemic to Galapagos Islands, Pacific Ocean. LOR,, 1964), The well known species of the genus have The systematic position of this species is still uncertain. a wide ränge of distribution. T. turbinata (LiNN.) It needs further investigations. B. tuberculata STACKHOUS KUNTZE is distributed from Florida to Brasil and T. is found along the Atlantic shores of Europe. alata (TURNER) J, AGARDH from Bermuda to the West The genus Pehetia DECAISNE et THURET comprises three Indies but not in the eastern Atlantic coast. To date species, P. wrightii (HARVEY) OKAMURA, P. canaltculata Turbinaria has not been reported from the west coast (LiNN.) DECAISNE et THURET and P. fastigiata (J, of Americas. T. ornata (TURNER) J. AGARDH extends AGARDH) DE TONI, Each of these species is widely from Hawaii Islands, Pacific Ocean, to Somalia, on the separated geographically in distribution. P, fastigiata is east coast of Africa and T. decurrens BORY from New common along the Pacific coast of North America, from Guinea to Madagascar on the east of Africa, including Oregon to Ensenada. P, wrlghtü occurs along the coast Red Sea where it is very common (NA$R> 1939. SIMON- of Japan and Korea, P. canaluulata is common along the SEN> 1968). T. gracilis SONDER is endemic to the south European coast of Atlantic Ocean, These three species west coast of Australia and T, triquetra (J. AGARDH) of Pelvetla grow in the upper-littoral region, J. AGARDH is native of Red Sea and Somalia. T. lu^o- Halidrys LYNGBYE comprises two species, //. dioica mnsis TAYLOR belongs to the Philippines and T, kenyaen- GARDNER and H, siliquosa (LiNN.) LYNGBYE. H. dioica ensis TAYLOR to African State (east). T. elatensis TAYLOR is endemic to Californian coast i.e. from Monterey to is endemic to Elat, Gulf of Aqaba and Sinai, on the Ensenada and grows in the lower to sub-littoral regions. Gulf of Suez, T. murrayana BARTON has been reported //. siUquosa is littoral along the north Atlantic shores of from Kenya and Madagascar. This species also occurs Europe and extends eastward to the Baltic Sea upto in the East Indies, Indonesia, Celebes, New Guinea and Kategatt. Its northern limit is north of Norway (TAY- Singapore. There is a limited distribution of T. papen- LOR, 1954) and southward to Bay of Bikay (CHALON, fttssii TAYLOR from Egypt to Ethiopia. There is a wide 1909). Along the coasts of British Isles this species distribution of T. conotdes (J, AGARDH) KÜTZING around grows from mid- to sub-littoral (BURROWS, I960, 1964, Kenya, Tanganyka, South India, Andaman islands, DIXON, 1963. JÖNES, 1960 and MC ALLTSTBR et aL, Singapore, Indonesia, Philippines and the Pacific Islands 1967). (Phoenix, Canton Is., Tonga Is., Samoa Islands and Fiji Ascuphyllitm STACKHOUS is monotypic and represented Islands). T. crateriformis TAYLOR belongs to Kenya and by A. nodösum (LiNN.) LE JQLLIS which grows in the T. filamentosa YAMADA is endemic to Formosa. The last mid-littoral region and also in sheltered positions, The important species, T. condensata SONDER in KÜTZING is species grows along the northern part of the Atlantic reported from Kenya, Somalia, Ceylon, Malaya, Indo- coast including North America and Western Europe1), nesia and the Philippines. The genus Turbinaria never reaches beiow the Cape of Good Hope or Western *) LAK.OWITZ (1929) reported the occurrence of A. nödomm var, Australia in the south or above China Sea in the north. scorpiodes HAUCK. from Baltic Sea. SCHWENKE (1969) and my observation show that Ascoj.hyllum nodositm does not grow in The genera Cystoseira C. AGARDH and Sargassum C Bälde Sea. AGARDH are of great biogeographical importance.

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There is a great need of a monograph of these two Tab. l Numertcal and percentage distribution of the fucalean genera in gencra and therefore at this stage it is not adviseable to different geographical regions of the world discuss their biogeographical importance. A limited Number (^ccurrcnce account of their distribution is given below. of in Cystoseira grows in difFerent habitats from upper to sub- Geographical regions genera percentage littoral region and most of the species belongs to the Australasia 3 endemic 8.3 northern hemisphere and few species do occur in the Australia £1 endemic 25.0 New Zealand 2 endemic 8,3 southern hemisphere. On the Pacific coast of America Japan 2! endemic 5.5 thrce species — C. mgltcta SETCHELL and GARDNER, California, North America £! endemic Ã>,5 Ouadalupe Island, Pacific Ocean C. osmundacea (MENZIES) C. AGARDH and C. setchcllii (Mexico) l endemic 2.8 GARDNER — occur which are littoral to sub-littoral. Galapagos Islands (Mexico), Europe and South Africa endemic 2.8 On the coast of West Africa Cystoseiras are present in North Pacific and North Atlantic fairly large amounts forming an important zone espe- coasts endemic 2,8 Europe and North America endemic 2.8 cially on the coast of Senegal and Mauritiana (LAWSON, Europe and America (North Atlantic 1966). PAPENFUSS and JENSEN -(1967) merged Cysto- coasts) endemic 2.6 Europe and Pacific coast (California) endemic 2.8 phyllum and Cystoseira s the formet genus was based Europe endemic 2.8 on variable characters i. e. vesides. Such variable char- South Africa 2 endemic 5.5 acters are also present in Cystoseira. TAYLOR (1960) re- Antarctica and sub-Antarctica endemic 2.8 Aden endemic 2.8 ports C. myrica (GMELIN) J. AG. from the eastern coast Pakistan and India (North Arabian Sea endemic 2.8 of America (Florida and Bahamas). ROBERTS (1967) re- Northern and southern hemispheres 5l) 13,9 ports five British species of Cystoseiras, C. baccata (GME- ') Including Cystophyllum Total =· 36 genera LIN) SILVA, C. foeniculacea (L.) GREVILLE, C. myriophyl- ioides SAUVAGEAU, C. nodicau s (WiTH.) ROBERTS and interesting to note that the species of Bifurcaria, FUCHS\ C. tawariscifolia (HUDSON) PAPENFUSS. Majority of the Halidrys and Pelvetia are widely separated and isolated species occur along the coast of Mediterranean and from their other counterparts. It is an established fact Adriatic Sea. One species occur in Red Sea (C myrica) that such Isolation must have taken place in geological and along the east coast of Africa. The phytogeograph- era. These genera may be of great importance in the ical distribution of Cystoseira now is extended to the far evolution. north upto Japan due to the merger of Cystophyllum There are six families of the order Fucales. Cystoseira- (PAPENFUSS and JENSEN, 1968). ceae and Sargassaceae are world wide in distribution Sargassum the most important genus of the order is and their spread is continuous (fig. 2). The family widely distributed in the world. It occurs in tropics, Himanthaliaceae is endemic to North Atlantic whereas sub-tropics and temperate zones in both hemispheres. Hormosiraceae and the Seirococcaceae are restricted to The number of species is conspicuous in tropics and Australian waters, Southern Ocean. The family Fucaceae sub-tropics. Sargassum is conspicuous at and below the is discontinuously distributed (fig. 2). The phytogeo- top of the sub-littoral region of the warm temperate graphical distribution of the families with their densities coasts of southern hemisphere (WOMERSLEY, 1959). In are given in fig. 2. The distribution of the fucalean the sub-tropical regions of the world Sargassum species families in the world gives a definite pattern whereas extend from mid- to sub-littoral regions. the generic distribution is more continuous than the Australia is also the chief centre of certain groups of family. Sargassum. The sub-genus Pkyllotrichia is mostly confined to Australia and Tasmania although two species are VI. The seasonal growth and the reproduction known from outside Australia, one from Canary Islands and the other from Japan. New Zealand is represented It has well been established that adult plants of Asco- by only one species, S. verruculosum (LINDAUER, 1947); pkyllum nodosum, FUCHS serratus, F. vesicalosus, Halidrys Artkrophycus is largely confined to the southern part of siliquosa, Himanthalia elongata and Margmariella nrvilliana Australia and Tasmania; S. undulatum J. AG. and S. show seasonal periodicity of growth and reproduction. lacerifolium C. AG. are found in New Zealand whereas NAYLOR (1953) observed that the first two year's growth S. heterophyllum J. AG. and S. longifolium J. AG. inhabit in Margmariellaurvillianav* entirely vegetative resulting in South African coast. Eusargassum is widely distributed distichously branched axis of leafy-lateral. At the begin- in warm waters. S. natans LINN. and S. hysterix J. AGARDH ning of its third year receptacles and vesicles are pro- are the floating species of the genus which are found duced; the receptacles take about a year to mature, so in the Sargasso Sea and also along the coast of eastern the plant is three years old by the time it produces the gametes. In this species the fertility period lasts till early part of America. August i. e. from the middle of May (NAYLOR, 1953). V. Distribution of Genera in a Nutshell Moss and LACY (1963) observed that first year's growth It is evident that a majority of the fucalean genera are of Halidrys siliquosa is entirely vegetative producing restricted to the Australian waters i. e. about 41.3%, so pinnately branched main axis of leafy-laterals. The sec- are the majority of Sargassum species (tab. 1). It is also ond year's growth is differentiated into vesicles and at

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the end of the second year receptacles develop, Similar and Ascöphyllum nodosum by LAMB and ZIMMERMANN type of growth has also been observed by GIBB (1937) (1964). MCALLISTER */ aL (1967) observed the con- in Himanthalia elongata. Moss and LACY (1963) also ceptable maturation in F. spiralis in March, June and observed similar pattern and development of growth in September. The release of gametes in F. vesiculösm i s FUCUS serrattts and F. vesictthsus in which receptacles did from March till June whereas in F. serratus takes place not develop till the plants reached 15—20 cm in iength from October to December. In the case of Ascophyllum (KNIGHT and PARK, 1950). TAYLOR (1937) described nodosum shedding of gametes takes place from April to that AscQphylhiw nodosun? i s able to persist and regenerate June in Norway (PRINTZ, 1959). from the basal parts or from the denuded branches. After fruiting and shedding of gametes teceptacles dis- Stokeyia indua exhibit similar pattern of growth along integrate or cast off down to the point where they join the coast of Karachi (personal observations). a branch, The basal stalk in Halidrys siliquosa is left äs NIZAMUDDIN and WOMERSLEY (1966) described the tooth-like projections (Moss and LACY, 1963). A series development of annual fronds in Myriodesma where each of projections are found each year, so it is quite possible year the frond is lost and only the lower midrib remains, to count the age of the plant, At the start of new growth one to several fronds arise SKOTTSBERG (1907) described that Cystosphaera jacquinctti at the summit of the remaining midrib. This process become fertile from November to January. continues every year, So it is possible to count the age Himanthalia elongata becomes fertile from July till Jan- of the plant. Myrwdesma commonly attain an age of uary and the release of gametes occur during the late 3—6 vears^ , summer and autumn (GiBB, 1937) but MCALLISTER et Vesicles also show a seasonal rhythm of growth and aL (1967) observed the fertile conceptacles in June on are generally produced in the spring in Fucus species. the coast of Scotland, In Halidrys siliquosa the gametes It is also possible to determine the age of the plant by are released during the winter. In contrast to Moss and the number of series of vesicles present. In Halidrys LACY (1963) gametangia are present throughout the sUiqnosa vesicles begin to be formed from about Sep- year except May and June äs described by BLACKLER tember to November (Moss and LACY, 1963). (1956) in Halidrys siliquosa. The factor affecting the periodicity of development are not yet known. Moss The reproductive period from receptacles Initiation to and LACY (1963) postulate that temperature and Iength gamete discharge followed by the disintegration of of day must have their effect but there is an inherent receptacles in Halidrys siliquosa is about nine months factor which may be specific. (Moss and LACY, 1950) three months in F. venculosus (KNIGHT and PARK, 1950) but gamete liberation takes VII. Acknowledgement over a period of 2—3 months in the case of //. sili- qtma but according to Moss and LACY (1963) recep- The author is grateful to Alexander von Humboldt- tacles mature between May and August i. e, c. 4 Stiftung for awarding "Dozentenstipendium" to carry months. A similar pattern of development was also on research at the Institut für Meereskunde, Universität observed by KNIGHT and PARK (1950) in F. vesiculosus Kiel.

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