Twenty Marine Benthic Algae New to South Africa, with Emphasis … · the seaweeds of the east coast of South Africa to be the westernmost extension of the large Indo-West Pacific

Botanica Marina Vol. 45, 2002, pp. 413–431 © 2002 by Walter de Gruyter · Berlin · New York

Introduction

The South African coastline displays a unique floraldiversity, with biota ranging from kelp beds on thewest coast to coral reefs near the Mozambican bor-der. The correlation between surface seawater tem-peratures and shore community compositions hasbeen investigated by several researchers, who havefirmly established that the main defining influencesare the warm Agulhas current flowing down the eastcoast from Mozambique to the region where it meetsthe cold Benguela current (e.g. Stephenson 1948,Bolton 1986, Bolton and Anderson 1990). Stephen-son (1948) divided the South African coastline intothree marine provinces: the cold-temperate westcoast, warm-temperate south coast, and sub-tropicaleast coast. More recently, seaweed biogeographers,although also recognising three floral provinces, havedescribed them differently. The seaweed flora of thewest coast is regarded as warm-temperate by Bolton(1986) and Lüning (1990) or cool-temperate byEmanuel et al. (1992) and Bolton and Anderson(1997), rather than cold-temperate.The presence of adistinct sub-tropical flora on the east coast (Kwazu-lu-Natal) is not accepted by either Lüning (1990) orBolton and Anderson (1997). These authors considerthe seaweeds of the east coast of South Africa to bethe westernmost extension of the large Indo-WestPacific tropical region, and cast doubt on the exis-tence of a distinct sub-tropical flora in Kwazulu-Na-tal.

The transition zone between the west and south

coast marine provinces has been well studied andcharacterised by a clearly defined overlap zone(Bolton and Stegenga 1990, Jackelman et al. 1991), incontrast to the eastern border of the Agulhas (southcoast) marine province which is less clear. Seawatertemperature, the main defining ecological parameterin the distribution of seaweeds, gradually rises fromaround East London to northern Kwazulu-Natal(Bolton 1986).At present, it is believed that a distinctsub-tropical marine province in Kwazulu-Natal is un-likely, as the flora seems to comprise an eastwardlydeclining number of Agulhas-province species, thesebeing replaced largely by Indo-West pacific specieswith the increase in water temperature (Bolton andAnderson 1997). It should be noted, however, thatour knowledge of the marine flora of Kwazulu-Natalis limited. Apart from a limited number of taxa, stud-ied in detail by students of G.F. Papenfuss in Berke-ley (California) and a field guide on the seaweeds ofMaputaland by Seagrief (1980), no detailed studieswere available relating to Natal algae prior to 1985.In the eighties and early nineties R.E. Norris and co-authors published a series of papers on the algae ofKwazulu-Natal that gave some evidence that there isindeed a distinct flora in the region. In total they (seeNorris 1992 and Silva et al. 1996 for a complete refer-ence list) reported 63 species as new records forSouth Africa and described 23 new species, many ofwhich are currently only known from Kwazulu-Natal. It has also been noted that elements of theKwazulu-Natal marine algal flora have a link withsouthwestern Australia (Norris and Aken 1985,

Twenty Marine Benthic Algae New to South Africa,with Emphasis on the Flora of Kwazulu-Natal

O. De Clercka*, H. R. Engledowa, J. J. Boltonb, R. J. Andersonc and E. Coppejansa

a Research Group Phycology, Biology Department, Ghent University, Krijgslaan 281 / S8, 9000 Ghent, Belgiumb Botany Department, University of Cape Town, Private Bag, 7700 Rondebosch, South Africac Marine and Coastal Management, Seaweed Unit, Private Bag X2, 8012 Roggebaai, Cape Town, South Africa

* Corresponding author: [email protected]

A total of 20 new records of benthic marine algae has been added to the flora of South Africa, consisting of6 taxa of Phaeophyta and 14 Rhodophyta. Most species have a pantropical or Indo-Pacific distribution andare generally known from several localities in the Indian Ocean (e.g. Asteronema breviarticulatum, Ceramiumcingulatum, Dictyota cervicornis, D. ciliolata, Euptilota fergusonii, Galaxaura rugosa, Halymenia durvillei,Phacelocarpus tristichus). Others are only known from a limited number of reports scattered within the Indo-Pacific region (Balliella crouanioides, Gibsmithia hawaiiensis, Predaea weldii, Hypoglossum minimum), possi-bly due to their subtidal habitat or small size. Apart from those algae with a large distribution range, somespecies show a distinctive southern Australian – South African distribution pattern (Carpopeltis phyllophora,Plocamium mertensii). Only Digeneopsis subopaca, originally described from Mozambique, appears to repre-sent a local endemic species.

Hommersand 1986). Kwazulu-Natal may thereforebe more than a transition zone between the Agulhasand Indo-West Pacific floras, in that it actually consti-tutes a distinct flora with warm-temperate south-western Australian affinities and a large number ofendemics.

In a joint research project conducted by theBotany Departments of the Ghent University (Bel-gium) and the University of Cape Town (SouthAfrica), the seaweeds of Kwazulu-Natal are beingstudied from a biogeographical perspective. Severalsites along the coast from Port Edward in the southto Kosi Bay near the Mozambican border were sam-pled mainly between July 1998 and February 2001.The collections are now being identified and willeventually serve to provide a better understanding ofthe distribution and floristic affinities of the algaealong the Kwazulu-Natal coast.This paper reports 20

414 O. De Clerck et al.

species which are newly recorded for South Africa.Records on Chlorophyta have recently been pub-lished separately by Leliaert et al. (2001).

Material and Methods

Approximately 2400 specimens were collected over aperiod of two years (July 1998; August 1999; Decem-ber 1999; July–August 2000; February 2001), as a re-sult of a Bilateral Scientific and Technological Coop-eration Project between the Flemish community(Belgium) and South Africa. Specimens were collect-ed at various sites distributed along the entire Kwazu-lu-Natal coast (see Fig. 1) and were immediatelyprocessed as herbarium specimens, as well as beingpartly preserved in 4% formaldehyde/seawater. Thespecimens, bearing serial numbers prefixed by ‘KZN’

Fig. 1. The coast of Kwazulu-Natal, South Africa showing the sample sites.

Twenty marine benthic algae new to South Africa 415

have been deposited in GENT and BOL. Slide mate-rial was stained in a mixture of 1 g aniline blue pow-der, 50 mL Karo®, 45 mL distilled water, 5 mL aceticacid. Collections other than our own, those of R.E.Norris and co-workers (bearing serial numbers pre-fixed by ‘NAT’) deposited in UN and various othercollectors in BOL were also examined. Herbariumabbreviations follow Holmgren et al. (1990). Digitalimages were taken, using an Olympus DP50 digitalcamera (Melville, USA) mounted on a Leitz Diaplanor Wild M10 microscope (Wetzlar, Germany).

Results

Phaeophyta

Dictyotaceae

Dictyota cervicornis Kützing, 1859: 11, pl. 24, fig. 2.Figs 2–4

Type locality: Key West, Florida.Description: Thalli form low, dense mats (Fig. 2).

The individual specimens are to 5 cm tall and com-posed of coarse, 2–3 mm wide, subdichotomousstraps (Fig. 4). The margins are smooth, but the sur-face is beset with acute proliferations (Fig. 3). Theapices are obtuse. Sporangia are scattered on bothsurfaces, but absent in the apical dichotomies. Spo-rangia, 120–135 µm in diameter, are placed mostlysolitary and are borne on a single stalk cell, surround-ed by a conspicuous involucrum. Gametangia werenot observed. In situ the colour is brown with a blue-greyish iridescence.

Ecology: Specimens occur in the sublittoral fringe,generally on wave-exposed shores.

Specimens: Simons 1262 (BOL): Kosi Bay(30.vi.1965); Simons 1254 (BOL): Sodwana Bay(3.vii.1965); KZN 312: Adlam’s Reef Sodwana Bay(9.viii.1999); KZN 579: Rabbit Rock (13.viii.1999);KZN 736: N23 (15.viii.1999); KZN 1013: Cape Vidal(9.vii.1998); KZN 1014: Mission Rocks (8.vii.1998).

Discussion: Dictyota cervicornis is perhaps themost common Dictyota species in the Indian Ocean,characterised by surface proliferations and sporangiathat are surrounded by an involucrum. It appears tobe closely related to D. crispata Lamouroux and D.magneana De Clerck et Coppejans (De Clerck andCoppejans 1997, Coppejans et al. 2001), but differsmainly in the placement of sporangia, these being ab-sent or present in the apical dichotomies respectively.Other characters which distinguish D. crispata fromD. cervicornis include the morphology of the apicesand the abundance and placement of surface prolif-erations. Detailed comparisons between the threespecies are made by Coppejans et al. (2001). Dictyotacervicornis exhibits a broad range of morphologicalgrowth forms, which can often be related to its habi-tat. Specimens growing in the shallow subtidal are of-ten characterised by long slender straps in the distal

parts of the thallus and recurved branchlets; speci-mens from exposed habitats generally form low,dense mats and lack slender straps. The SouthAfrican specimens represent the typical growth formfrom exposed habitats. A major difference to previ-ous observations is the occurrence of iridescence.Despite the examination of over 200 specimens byDe Clerck (1999), mainly from the East African coast(Tanzania), iridescence was never observed. The tax-onomic value and intraspecific variability of irides-cence, however, remains poorly understood (Gaillard1972).

Dictyota ciliolata Sonder ex Kützing, 1859: 12, pl. 27,fig. 1. Figs 5, 6

Type locality: La Guaira,Venezuela.Description: Thalli are erect, to 8 cm tall, attached

by means of a single stupose holdfast (Fig. 6).Stolonoidal fibres are absent. Straps, 2–3 mm wide,are slender and dichotomously branched (Fig. 5).Themargins are dentate, rarely smooth, while the surfaceis always smooth. The apices are rounded. Themedulla and cortex are uniformly one-layered. Spo-rangia are single, scattered on both surfaces, but ab-sent in the apical dichotomies. Sporangia, 95–110 µmin diameter, are borne on a single stalk cell and arenot surrounded by a conspicuous involucrum. Ga-metangia were not observed.

Ecology: Specimens were collected in intertidalpools of the sublittoral fringe down to –12 m.

Specimens: KZN 396 and KZN 401: Mabibi(9.viii.1999); KZN 2182: Mabibi (13.ii.2001).

Discussion: Dictyota ciliolata is characterised by itsstupose holdfast, dentate margins and the absence ofstolonoidal fibres, although the margins of somespecimens can be nearly smooth (Hörnig et al.1992a,b). Several other dentate species of Dictyotaoccur in the Indo-Pacific and the Caribbean regions,as detailed by De Clerck and Coppejans (1999).Along the South African coast D. ciliolata is mostsimilar to D. liturata J. Agardh and D. suhrii G. Mur-ray. The latter species, however, has a multilayeredmedulla near the margins, whereas the medulla of D.ciliolata is uniformly one-layered. Dictyota ciliolatadiffers from D. liturata in the placement of the spo-rangia, which are arranged singly or distributed inlongitudinal lines, respectively.

Dictyota hamifera Setchell, 1926: 92, pl. 14,figs. 1–6. Fig. 7

Type locality: between Papenu and Huau,Tahiti.Description: The thallus is repent, up to 3 cm, at-

tached to hosts at various points by patches of rhi-zoids. The straps, 1–2 mm wide, are dichotomouslybranched. The margins are smooth except for the fal-cate branchlets (Fig. 7), which may resemble teeth.The surface is smooth. The apices are rounded to tri-dentate. The medulla and cortex are uniformly one-



layered. Sporangia or gametangia were not ob-served.

Ecology: Specimens were collected at a depth of 18to 25 m epiphytic on Plocamium sp.

Specimens: KZN 1617: Sodwana Bay, 2 Mile Reef(12.viii.2000).

Discussion: Dictyota hamifera was for a long timeonly known from the Pacific Ocean. Hörnig et al.(1992a) reported the species for the first time from theAtlantic Ocean,while De Clerck (1999) and Coppejanset al. (2000) recorded the species from the East Africancoast (Kenya and Tanzania).Dictyota hamifera is easilyrecognised by its falcate,swollen branchlets.

Dictyota rigida De Clerck et Coppejans, 1999:189–191, figs. 10–17. Figs 8–10

Type locality: Kunduchi, Dar es Salaam,Tanzania.Description:Thalli are up to 6 cm high, attached by

a small stupose holdfast. The dichotomouslybranched straps are narrow (0.8–1.2 mm) and of uni-form width throughout the thallus (Figs 8, 10). Themargins and surface are smooth. The medulla andcortex are uniformly one-layered. Sporangia form awell-defined line in the median part of the straps, butare not present along the margins (Fig. 9). Sporangia,80–95 µm in diameter, are not surrounded by an in-volucrum and are borne on a single stalk cell. Ga-metangia were not observed.

Ecology: Specimens were collected in pools of thesublittoral fringe.

Specimens: KZN 311: Sodwana Bay, Adlam’s Reef(9.viii.1999); KZN 385: Mabibi (9.viii.1999).

Discussion: Dictyota rigida was recently describedfrom Tanzania (De Clerck and Coppejans 1999) andsubsequently reported from Kenya (Coppejans et al.2000). Characters defining D. rigida include the erectgrowth form, the stiff texture and the single stuposeholdfast (De Clerck and Coppejans 1999). Dictyotarigida is most similar to the South African D. suhrii

G. Murray and the Australian D. fastigiata Sonder.The latter two species, however, are characterised bya multilayered medulla, at least in the basal parts ofthe thallus.

Padina gymnospora (Kützing) Sonder,1871: 47. Figs 11, 12

Type locality: St.Thomas,Virgin Islands.Description:Thalli, attached by a small stupose rhi-

zoidal base, are to 6.5 cm high, flabellate and com-posed of several lobes with inrolled margins (Fig. 11).The superior surface (the side toward which the mar-gin is inrolled) is slightly to moderately calcified.Thalli are olive-brown in colour. In transverse sec-tion the thallus is composed of 4–6 layers of cells inthe mid-regions and up to 8–9 layers near the base.Hair rows are present on both sides of the thallus, butmore conspicuous on the inferior side of the thallus.Tetrasporangia are arranged in concentric lines,above each hair row, mainly (but not exclusively) de-veloped on the superior surface.An evanescent indu-sium is present but is usually only clearly visible invery young sori (Fig. 12). Tetrasporangia are ovoid,up to 100 µm long and 70 µm wide. Gametophyteswere not observed.

Ecology: Specimens are collected in shallow poolsin the sublittoral fringe.

Specimens: KZN 94: Durban, Treasure Beach(3.viii.1999); KZN 294: Sodwana Bay, Adlam’s Reef(9.viii.1999); KZN 361: Mabibi (9.viii.1999); KZN708: Bhanga Nek, N23 (15.viii.1999); KZN 745: KosiBay (16.viii.1999).

Discussion: Padina species are distinguished pri-marily on the number of cell layers, the distributionof hair bands, position of the sporangial sori (on thesuperior versus inferior surface) and the type of al-ternations of hair and sporangial bands. Padina gym-nospora is characterised by the thick thallus (up to 8cell layers), the distribution of sporangia primarily on

Figs 2–4. Dictyota cervicornis.Fig. 2. Habit (scale = 1 cm). Fig. 3. Detail of the surface with multiple proliferations (scale = 2 mm). Fig. 4. Detail of the api-cal parts of the thallus (scale = 2 mm).

Figs 5, 6. Dictyota ciliolata.Fig. 5. Habit (scale = 1 cm). Fig. 6. Detail of the stupose base (scale = 2 mm).

Fig. 7. Dictyota hamifera. Habit showing falcate branchlets (arrows) (scale = 2 mm).

Figs 8–10. Dictyota rigida.Fig. 8. Habit (scale = 1 cm). Fig. 9.Tetrasporangia aggregated in the median region of the straps (scale = 1 mm). Fig. 10. De-tail of the apical parts of the thallus (scale = 2 mm).

Figs 11, 12. Padina gymnospora.Fig. 11. Habit (scale = 1 cm). Fig. 12. Transverse section of a 6–7 layered thallus with a young, indusiate tetrasporangialsorus (scale = 100 µm).

Figs 13–15. Asteronema breviarticulatum.Fig. 13. Habit of herbarium specimen illustrating the frayed, rope-like appearance (scale = 1 cm). Fig. 14. Detail of ahooked branch (scale = 50 µm). Fig. 15. Detail of an ovoid plurilocular sporangium (scale = 50 µm).

the superior surface of the thallus and the evanescentindusium (Allender and Kraft 1983, Womersley1987). The species is most similar to P. crassa Yama-da, the latter supposedly having sporangia only onthe lower side of the thallus. Only P. boryana Thivyand P. plumbea (Areschoug) Levring have previouslybeen reported from South Africa (Levring 1940,Gaillard 1975). Both species are characterised by a 2-layered thallus and hence are easily distinguishedfrom P. gymnospora. Due to confusion with P.boergesenii Allender et Kraft in the past, the distribu-tion of P. gymnospora is difficult to assess in the Indi-an Ocean (Silva et al. 1996). The latter, however, hasbeen reported with certainty from Australia andOman (Womersley 1987,Wynne and Jupp 1998).

Incertae sedis

Asteronema breviarticulatum (J.Agardh)Ouriques et Bouzon, 2000: 271. Figs 13–15

Type locality: San Agustín, Oaxaca, Mexico.Description:Thalli, approximately 2.5 cm tall, form

tufts of interwoven filaments, which have the appear-ance of a frayed rope at its tips (Fig. 13). Filamentsare uniseriate and irregularly branched with numer-ous short hooked branches arising at an angle of80–90 ° (Fig. 14). Growth takes place by intercalarycell divisions. Cells are 25–33 µm wide, 60–80 µmlong, L/B: 1–1.3. Plurilocular sporangia, placed on a1–2-celled stalk, are spherical to ovoid and measure40 x 42 (–63) µm on average (Fig. 15).

Ecology: growing in the intertidal, often attachedto barnacles in wave-exposed localities.

Specimens: KZN 451: Mabibi (11.viii.1999); KZN654: Black Rock (14.viii.1999); KZN 728: N23(15.viii.1999).

Discussion: Asteronema breviarticulatum is a wide-spread warm-temperate to tropical species charac-terised by the shape of the plurilocular sporangia andthe presence of short hooked branches. Ouriques andBouzon (2000) transferred this species, which wascommonly known as Hincksia breviarticulata (J.Agardh) Silva, to Asteronema on the basis of the stel-late chloroplasts. Recent molecular studies by Drais-ma et al. (2001) and Rousseau et al. (2001) reveal,however, that Asteronema is polyphyletic and not re-lated to the Ectocarpales sensu lato. UnfortunatelyAsteronema breviarticulatum itself was not includedin the analysis. Pending further studies, A. breviartic-ulatum is listed as incertae sedis rather than under theEctocarpales.

Rhodophyta

Ceramiales

Ceramiaceae

Balliella crouanioides (Itono) Itono et Tanaka, 1973:250. Figs 16–19


Type locality: Mage Island, Japan.Description: Thalli form woolly tufts composed of

uniseriate filaments to 40 mm high (Fig. 16). Fila-ments are opposite-distichously branched with inde-terminate branches being formed every (1–) 2 (–5)segments. The apical parts of the axes show a distinc-tive sinusoidal curving (Fig. 17). Corticating rhizoidsarise from the lower cells of the laterals and form aloose rag-like cortex (Fig. 19). The periaxial cells are100–115 µm in diameter and bear spherical vesicularcells, 8–15 µm in diameter on their abaxial side(Fig. 18). Decussately divided tetrasporangia developon the adaxial side of the periaxial cells of the later-als. Tetrasporangia are ovoid, 30–55 µm long and30–38 µm wide. Gametophytes were not observed.

Ecology: Balliella crouanioides is a common com-ponent of the sublittoral zone (–20 to –35 m), whereit forms distinctive orangy-iridescent tufts on a vari-ety of substrates (coral debris, various algae, sponges,etc.).

Specimens: KZN 63 Aliwal Shoal (3.viii.1999);KZN 147: Aliwal Shoal (4.viii.1999); KZN 2093: Sod-wana Bay, Deep Sponge (10.ii.2001); KZN 250 andKZN 257: Sodwana Bay, 2 Mile Reef (8.viii.1999);NAT 6232 (UN): Sodwana Bay, 2 Mile Reef(14.x.1989); KZN 644: Bhanga Nek, Sexton Reef(14.viii.1999).

Discussion: Balliella species are recognisable bythe presence of spherical vesicular cells (‘glandcells’), which occur adaxially or abaxially on the peri-axial cells. Species are separated on the basis of theirhabit, branching pattern, position and size of vesicu-lar cells and tetrasporangia. Wollaston (1984) report-ed B. subcorticata (Itono) Itono et Tanaka fromMozambique, but Huisman (1988) expressed theopinion that the specimen depicted was more similarto B. crouanioides, based on the overall branchingpattern, arrangement of the tetrasporangia and sizeof the vesicular cells. Comparison of our materialwith the specimen studied by Wollaston (UC1470004) reveals that they are identical and supportsHuisman’s proposition. Athanasiadis (1996) provi-sionally assigned a collection of specimens fromKenya to B. crouanioides. Re-examination of thetype collection proved to be necessary as some con-troversies exist on the branching pattern and place-ment of the gland cells in this species. The SouthAfrican specimens match previous accounts of B.crouanioides very well (Itono and Tanaka 1973, Itono1977, Huisman and Kraft 1984), although variationwas observed in both the size (8–15 µm in diameter)and placement of the gland cells (abaxial but occa-sionally adaxial). As these characters are regarded tobe of prime importance in delineating Balliellaspecies, further research is needed on intraspecificvariation.

Ceramium cingulatum Weber-van Bosse, 1923:332–333, figs. 123, 124. Figs 20–22


Figs 16–19. Balliella crouanioides.Fig. 16. Habit of herbarium specimen (scale = 1 cm). Fig. 17. Detail of the sinusoidal apex and opposite-distichous branch-ing (scale = 100 µm). Fig. 18. Cruciately divided adaxial tetrasporangia and abaxial gland cells (arrowheads) on the periax-ial cells (scale = 25 µm). Fig. 19. Loose cortication at the nodes of the axes in the lower part of the thallus (scale = 100 µm).

Figs 20–22. Ceramium cingulatum.Fig. 20. Habit of the typically clavate axis (scale = 500 µm). Fig. 21.Whorled tetrasporangia (scale = 100 µm). Fig. 22. Detailof the nodal structure in a proximal axis showing two anterior (arrowheads) and two posterior (arrows) cortical filaments(scale = 100 µm).

Figs 23–26. Euptilota articulata.Fig. 23. Habit of herbarium specimen (scale = 1 cm). Fig. 24. Apical portion with alternate-distichous branching(scale = 100 µm). Fig. 25. Ultimate branchlet bearing lateral and terminal tetrasporangia (scale = 100 µm). Fig. 26. Detail ofan ultimate branchlet with tetrasporangia (scale = 50 µm).

Figs 27–29. Euptilota fergusonii.Fig. 27. Habit of herbarium specimen (scale = 1 cm). Fig. 28. Young lateral axes with curved, abaxial branchlets(scale = 500 µm). Fig. 29. Detail of a tetrasporangial branchlet (scale = 100 µm).

Type locality: Sape Strait, south of Sangeang Island,Indonesia.

Description: Thalli to 2 mm high consisting of un-branched erect axes, that arise mainly from a singlepoint or from a creeping filament. Axes are some-what clavate, measuring 90–100 µm wide near thebase, 140–170 µm in the middle part of the thallus,ending abruptly in an attenuate apex (Fig. 20). Axialcells are lenticular, usually wider than long exceptnear the base. A total of 8 periaxial cells are presentper segment, each producing 2 anterior and 2 posteri-or cortical initials of cortical filaments. Anterior ini-tials cut off 1–2 acropetal filaments, which are up to 2cells long (Fig. 22). Posterior initials either remainundivided or produce a single basipetal filament(Fig. 22). Cortication remains incomplete, but adja-cent cortical bands may be nearly confluent.Tetrasporangia form in distinctive whorls, are slightlyprotruding and measure 28–35 µm in diameter(Fig. 21). Gametangia were not observed.

Ecology: Epiphytic on Plocamium telfairiae at –18to 25 m.

Specimens: KZN 1619.2: Sodwana Bay, 2 MileReef (12.viii.2000).

Discussion: The South African specimens agreevery well with the original description by Weber-vanBosse (1923) and a subsequent account of the speciesby Cormaci and Furnari (1991), who compared C.cingulatum with the Mediterranean C. giacconei Cor-maci et Furnari.The only difference we observed wasthe mode of attachment. Typically, C. cingulatum haserect axes in the terminal parts or produces erectbranches. South African plants were usually strictlyerect, but the presence of rhizoids in the lower part ofthe thallus indicates the possibility of a creepinggrowth form. Strictly erect versus ascending growthforms are not considered to be major taxonomic dif-ferences in Ceramium (Dixon 1960, Womersley1978). Ceramium cingulatum is unique among SouthAfrican representatives of the genus in its un-branched erect axes.The species has been reported inthe Indian Ocean from Somalia, Tanzania and theSeychelles (Jaasund 1970, Sartoni 1975,Wynne 1995).

Euptilota articulata (J.Agardh) Schmitz, 1896: 7.Figs 23–26

Type locality:Australia.Description:Thalli are erect, to 25 cm high and pin-

nately branched up to 5 orders (Fig. 23).A percurrentmain axis is lacking.The holdfast is densely rhizoidal.Axes are corticated from close to the apices, with cor-ticating filaments arising from the periaxial cells 5 to10 cells proximal to the apex. Near the base the axesattain a width of 1.5 mm. Each axial cell bears a singlelateral in a distichous-alternate pattern (Fig. 24). De-terminate branchlets are ecorticate, straight, 12–17cells long and pinnately branched (Fig. 25). Spine-like cells are absent. Indeterminate branchlets are


formed irregularly and repeat the branching patternof the main axes. Tetrahedrally divided tetrasporan-gia (40–52 µm in diameter) are borne laterally or ter-minally on the ultimate branchlets (Fig. 26). Gameto-phytic plants were not observed.

Ecology: Euptilota articulata has only been collect-ed once in situ, at a depth of 35 m in southern Natal.The specimen in the Norris Herbarium (UN) wascollected from the drift.

Specimens: KZN 1962: Protea Banks, NorthernPinnacle (5.ii.2001); NAT 1310: Palm Beach(14.v.1983).

Discussion: The genus Euptilota contains at pres-ent four species, all of which are characterised by analternately-distichously branched thallus, in whichaxial cells each bear a single lateral (Millar 1990,Womersley 1998). Euptilota articulata differs fromE. pappeana Kützing, the only species reportedfrom South Africa (Stegenga et al. 1997), in the pin-nate branching of the determinate laterals as op-posed to laterals, which are either unbranched oronce to twice dichotomously branched in the latterspecies. Euptilota articulata has previously been re-ported for Australia (Millar 1990, Womersley 1998),Japan (Itono 1977) and India (Umamaheswara Rao1974).

Euptilota fergusonii Cotton, 1907: 262–264.Figs 27–29

Type locality: ‘Pantura’, Sri Lanka.Description: Thalli are erect, to 15 cm high, pin-

nately branched up to 5 orders and lack a percurrentmain axis (Fig. 27). The holdfast is densely rhizoidal.Axes are corticated from close to the apices with cor-ticating filaments arising from the periaxial cells, ap-proximately 12–14 cells proximal to the apex. Proxi-mal axial cells are 450–500 µm wide and 240–270 µmlong. Each axial cell bears a single lateral in adistichous-alternate pattern (Fig. 28). Determinatebranchlets are ecorticate, curved upwards and up to12 cells long.The basal 3 cells of a determinate lateralbear each a short adaxial side branch, the fourth tothe ninth cell bear an arched abaxial branchlet andthe remaining cells remain unbranched, except forthe ultimate and penultimate cells, which bear 1–2(–4) spine-like cells (Fig. 29). Indeterminate branch-lets are formed irregularly and repeat the branchingpattern of the main axes. Tetrahedrally dividedtetrasporangia (60 µm long, 40–50 µm wide) areborne laterally on cells of the ultimate branchlets.Gametophytes were not observed.

Ecology: Collected at two occasions only, at adepth of 15–25 m in northern Kwazulu-Natal (Sod-wana area).

Specimens: KZN 342: Sodwana Bay, 2 Mile Reef(9.viii.1999); KZN 419: Sodwana Bay, 7 Mile Reef(9.viii.1999).

Discussion: Euptilota fergusonii is distinguished


from other Euptilota species in having spine-like cellsat the tips of the determinate laterals, the latter addi-tionally being secundly, as opposed to alternately orsubdichotomously, branched. Euptilota fergusoniiwas originally described from Sri Lanka (Cotton1907) and has previously been reported from Tanza-nia and Mozambique (Jaasund 1976, Wollaston1984).

Delesseriaceae

Hypoglossum minimum Yamada, 1936: 138–140,fig. 2. Figs 30–33

Type locality: Naha, Okinawa-jima, Ryukyu-retto,Japan.

Description: Thalli consist of unbranched, lanceo-late blades (7–12 mm high and 0.8–2.3 mm broad),attached by means of a small discoid holdfast(Fig. 30). Growth takes place from a single apical cell,with a typical type-one apical organisation (i.e. allsecond order row cells produce third order rows).The midrib remains uncorticated throughout.Tetrasporangial sori form discrete, paired ovate soriclose to the midrib (Fig. 31), 900–1700 µm long and400–600 µm wide. Tetrasporangial formation in-volves the lateral pericentral cells but not the trans-verse ones. Tetrasporangia, 62–80 µm in diameter,are cut off from second- and third order cell rows aswell as from cortical cells, resulting in sporangia thatlie in more than one plane of the blade. Spermatan-gial sori form discrete, somewhat oblique patchesscattered over the wings (Fig. 32). Female plants pro-duce a single cystocarp on the midrib in the distal re-gion of the blades (Fig. 33).

Ecology: Collected on a single occasion, epiphyticon Carpomitra longicarpa Simons at –43 m.

Specimen: KZN 506: Tiger Reef, Bhanga Nek(13.viii.1999).

Discussion: Hypoglossum minimum is a smallspecies, which can easily be confused with H. simu-lans Wynne, Price et Ballantine and H. barbatumOkamura, of which the latter has also been recordedfrom Kwazulu-Natal (Wynne and Norris 1991). Hy-poglossum minimum is distinguished from the twoother species by its erect rather than creeping habit(being attached by a single discoid holdfast), un-branched blades, a type-one apical organisation andtetrasporangia that are cut off from lateral pericen-tral cells. Although some of these features are alsopresent in both H. simulans or H. barbatum, this com-plete combination of characters is unique to H. mini-mum (Wynne et al. 1989, Wynne and De Clerck2000). The only other Indian Ocean record comesfrom the Maldives (Hackett 1977). Recently Stegen-ga et al. (2001) described H. imperfectum from theSouth African south coast. The latter, however, dif-fers from H. minimum in its prostrate habit and byfertile blades within which the proximal third-ordercell rows fail to reach the thallus margin.

Rhodomelaceae

Digeneopsis subopaca Simons, 1970: 10, 11. Fig. 34

Type locality: Santa Maria, Inhaca, MozambiqueDescription:Thalli are erect, very stiff and blackish

in colour, to 11 cm tall (Fig. 33). Indeterminate axesare pinnately branched, compressed near the apices,but become rounded in the lower parts of the thallus.Determinate branchlets are stiff and incurved,4–10 mm long and to 500 µm in diameter, bi- or tri-furcately branched in up to 2 or 3 orders. The thallusis polysiphonous, consisting of an axial cell surround-ed by 5 pericentral cells, which are obscured by awell-developed cortex from close to the apices.Tetrasporangia are formed in clustered stichidia nearthe bases of determinate laterals. The stichidia are100–190 µm thick, with each segment bearing 2tetrasporangia. Tetrasporangia are up to 115 µm indiameter and tetrahedrally divided. Gametophyteswere not observed.

Ecology: Collected in intertidal rock pools and theshallow sublittoral from Mapelane (just south of St.Lucia) to Mozambique.

Specimens: KZN 368: Sodwana Bay, intertidal(9.viii.1999); KZN 569: Bhanga Nek (13.viii.1999);KZN 748: Kosi Bay (16.viii.1999); KZN 1641: Mabibi(13.viii.2000); KZN 1809: Cape Vidal (18.viii.2000);KZN 1843: Mapelane, Crayfish Point (20.viii.2000);Pocock and Papenfuss 1024 (UN): St. Lucia(21.vii.1938).

Discussion: Digeneopsis subopaca, originally de-scribed from southern Mozambique (Simons 1970),remains a poorly known representative of theRhodomelaceae. In the absence of gametophytic ma-terial the taxonomic affinities of this monotypicgenus are unclear. The morphology of the tetraspo-rangial stichidia and the compressed, bilateral natureof the plants, however, seem to indicate a close rela-tionship to the Amansieae. The South African recordof Halopithys incurva (Hudson) Batters in Seagrief[1984, as H. pinastroides (S.G. Gmelin) Kützing],seems to be based on a misidentification of a Pocockand Papenfuss collection (nr. 1024) which is referableto D. subopaca. The relationship of Digeneopsis sub-opaca to Halopithys and the other Amansieae is be-ing studied by L. Phillips and G.T. Kraft (pers. com.).

Gigartinales

Dumontiaceae

Gibsmithia hawaiiensis Doty, 1963: 458–465, figs.1–17. Fig. 35

Type locality: Honolulu, Oahu, Hawaiian Archipel-ago.

Description: Thalli are to 4 cm high, pinkish-red incolour and composed of 4–5 gelatinous lobes, whichare attached to a conspicuously cartilaginous, annu-late stalk. The stalk is usually simple but may branch

twice in well-developed specimens. The lobes are un-divided and are broadly rounded, measuring approx-imately 1 cm in diameter and 3 cm in length.The thal-lus is multiaxial, composed of a filamentous medullaand cortex embedded in a gelatinous matrix.The dis-tal ends of the cortical filaments produce the gelati-nous matrix. Specimens were completely sterile.

Ecology: Growing epilithically at a depth of 20 min northern Kwazulu-Natal.


Specimens: KZN 614: Sexton Reef (14.viii.1999);KZN 1611: Sodwana Bay, 2 Mile Reef (12.viii.2000).

Discussion: Gibsmithia hawaiiensis was originallydescribed from the Hawaiian Archipelago and ap-pears to be a common component of the deeper sub-tidal in the central, western and southern PacificOcean (Kraft 1986, Abbott 1999). Its presence in theIndian Ocean was until recently restricted to reportsfrom Australia (Huisman 1992) and the Seychelles


(Kalugina-Gutnik et al. 1992). However, a reportfrom the East African coast (Coppejans et al. 2000)and the present collection indicates that G. hawaiien-sis is the most widely distributed species of the genus.

Nemastomataceae

Predaea weldii Kraft et Abbott, 1971: 194, figs 1–13.Figs 36–39

Type locality: Kaneohe Bay, Oahu Island, Hawaii.Description: The gelatinous thallus is about 3.5 cm

high,irregularly branched with numerous short,blunt-tapering branchlets (Fig. 36).The colour is pale pink-ish in situ.The thallus is multiaxial, composed of a fila-mentous medulla and cortex embedded in agelatinous matrix. The cortical filaments are dichoto-mously branched, with rectilinear cells, 4–5 µm wideby 8–13 µm long (Fig. 37).Gland cells are absent.Car-pogonial branches are 3-celled (Fig. 39). Auxiliarycells are placed intercalary in cortical filaments anduteriform in shape with a prominent apical bulge. Nu-tritive cells, grouped per 3–6 in chains 1–3 cells long,are present on the cortical cells immediately belowand on the 2 cells distal to the auxiliary cell (Fig. 38).The gonimoblast arises apically from the auxiliary celland not laterally in conjunction with the connectingfilament.Male gametophytes were not observed.

Ecology: Collected only once, attached to coralrubble at a depth of 18 to 25 m in northern Kwazulu-Natal.

Specimen: KZN 1623: Sodwana Bay, 2 Mile Reef(12.viii.2000).

Discussion: The genus Predaea is distinguishedfrom all other red algae by the clusters of nutritivecells (Kraft 1984). From the Indian Ocean only P.feldmannii Børgesen var. indica Balakrishnan etChawla (1984) and P. huismanii Kraft (1984) havebeen hitherto reported. Predaea weldii is charac-

terised by its rectilinear-shaped cortical cells, the lackof gland cells, the relatively sparse and large nutritivecells and the gonimoblasts, which arise apically fromthe auxiliary cells. This is the first record of thisspecies for the Indian Ocean.

Phacelocarpaceae

Phacelocarpus tristichus J.Agardh, 1885: 57–58.Figs 40, 41

Type locality: Mauritius.Description: Thalli are erect, 7–10 cm high, sparse-

ly branched and attached by a small discoid holdfastfrom which a short (up 1 cm) terete stipe arises(Fig. 41).Axes are of uniform width, terete, 1.5 mm indiameter (including teeth). Triangular teeth areformed in 3 ranks along the axes (Fig. 40) except nearthe base, where there are 2 ranks. Teeth are slightlyupwardly directed, 500 µm long and 270 µm widenear the base. Reproductive structures were not ob-served.

Ecology: Growing epilithically in large intertidalpools (Island Rock) and in the sublittoral to a depthof 43 m.

Specimens: KZN 243: Sodwana Bay, 2 Mile Reef(8.viii.1999); KZN 2180: Mabibi (13.ii.2001); KZN1680: Island Rock (14.viii.2000); KZN 516:Tiger Reef(13.viii.1999)

Discussion: The genus Phacelocarpus contains 9species primarily distributed in the Southern Hemi-sphere (Indian Ocean, Australia and New Zealand),with the exception of P. japonicus Okamura fromsouthern Japan (Womersley 1994). Phacelocarpustristichus is predominantly distinguished from theother species by the placement of teeth in 3 ranksalong the axes (although the number of ranks mayvary from 2 to 4; Searles 1968), the majority ofspecies having teeth in only 2 ranks. From South

Figs 30–33. Hypoglossum minimum.Fig. 30. Habit of a specimen epiphytic on Carpomitra longicarpa (scale = 1 cm). Fig. 31.Tetrasporic sorus near the apex of ablade (scale = 500 µm). Fig. 32. Male sori in irregular chevrons on both sides of the blade midline (scale = 500 µm). Fig. 33.Mature cystocarp near a blade apex (scale = 500 µm).

Fig. 34. Digeneopsis subopaca. Habit of herbarium specimen (scale = 2 cm).

Fig. 35. Gibsmithia hawaiiensis. Habit of herbarium specimen (scale = 1 cm).

Figs 36–39. Predaea weldii.Fig. 36. Habit of a liquid-preserved specimen (scale = 1 cm). Fig. 37. Cortical fascicles of rectilinear cells (scale = 100 µm).Fig. 38.Auxiliary cell with apical gonimoblast initial (arrow) and clusters of nutritive cells (arrow heads) on the contiguouscortical cells (scale = 10 µm). Fig. 39.Three-celled carpogonial branch (scale = 10 µm).

Figs 40, 41. Phacelocarpus tristichus.Fig. 40. Distal frond showing the tristichous placement of determinate branchlets (scale = 2 mm). Fig. 41. Habit of herbari-um specimen (scale = 1 cm).

Figs 42–44. Halymenia durvillei.Fig. 42. Habit of herbarium specimen (scale = 1 cm). Fig. 43. Detail of the blade apices (scale = 1 mm). Fig. 44. Detail of thethallus surface with minute spinose proliferations (scale = 2 mm).

Fig. 45. Carpopeltis phyllophora. Habit of herbarium specimen (scale = 1 cm).

Africa two species are known: P. oligocanthus Kütz-ing and P. tortuosus Endlicher et Diesing, both occur-ring along the South African south coast and extend-ing into Kwazulu-Natal (Searles 1968, Seagrief 1988).Phacelocarpus oligocanthus is a robust species char-acterised by teeth, which become submerged in thecortex of the main axes. Phacelocarpus tortuosus iscloser to P. tristichus in size and habit but differs inthe bilateral placement of the teeth and flattenedaxes to 2.5 mm wide (Searles 1968). Phacelocarpustristichus, originally described from Mauritius, ap-pears to be widely spread throughout the western In-dian Ocean (Silva et al. 1996).

Halymeniales

Halymeniaceae

Halymenia durvillei Bory de Saint-Vincent, 1828:180–181, pl. 15. Figs 42–44

Type locality: Port Praslin, New Ireland, Papua NewGuinea.

Description: Thalli are erect, to 21 cm high andcomposed of irregularly branched, lubricous straps(Fig. 42).The base consists of a small discoid holdfast.Axes are branched up to 5 orders, taper from thebase to the apices and measure 4–20 mm in width.Apices are acute and the surface is covered withacute proliferations (Figs 43, 44), which may developinto new axes. On transverse section the thallus is ap-proximately 600 µm thick at mid-frond, composed ofa pseudoparenchymatous cortex and a filamentousmedulla. The cortex is 60–80 µm thick, composed of6–8 cells forming anticlinal rows. Medullary fila-ments are mainly anticlinally arranged, 8–12 µm indiameter, with relatively abundant inner-cortical re-fractive ganglionic cells. The latter are irregularlyshaped, with branched arms. Decussately dividedtetrasporangia (14–20 µm long, 12–15 µm wide) arescattered over the thallus and are cut off from sub-surface cells. Gametophytes were not observed.

Ecology: Growing in deep intertidal rock pools orthe shallow subtidal (–3 m).

Specimens: KZN 429: Sodwana Bay, 1/4 Mile Reef(10.viii.1999); KZN 2155: Sodwana Bay, 9 Mile Reef(12.ii.2001); KZN 0782: Island Rock (17.viii.1999);NAT 4685 (UN): Black Rock (21.xi.1986).

Discussion: Halymenia durvillei is a widespreadand easily recognisable species. The branching habitis very distinctive and separates it from most otherspecies of Halymenia, with foliose blades. FromSouth Africa, only H. dilatata Zanardini (Norris andAken 1985) has been previously reported. Thisspecies has leafy blades with mottled surfaces, whichrule out possible confusion with H. durvillei, whichhas been reported under various synonyms (e.g. H.ceylanica Harvey ex Kützing, H. formosa Harvey exKützing and H. venusta Børgesen) within the tropicalIndian Ocean (De Smedt et al. 2001).


Carpopeltis phyllophora (J. Hooker et Harvey)Schmitz in Schmitz et Hauptfleisch, 1897: 514. Fig. 45

Type locality: Port Arthur,Tasmania.Description: Thalli are erect, to 13 cm high, com-

planate and composed of dichotomously to irregular-ly branched axes, 2–7 mm in width. The base consistsof a small discoid holdfast, which extends into a con-spicuous midrib in the lower parts of the thallus. Theapices are broadly rounded to obtuse, but proliferatewith mechanical damage. On transverse section thethallus is 150–200 µm thick at mid-frond, composedof a pseudoparenchymatous cortex and a filamen-tous medulla. The cortex is 50–70 µm and 4–5 cellsthick, composed of cells gradually decreasing in sizetowards the periphery, which are not arranged in an-ticlinal rows. The medulla comprises about 1/3 thirdof the thallus thickness at maximum and is composedof compactly arranged filaments. Refractive gan-glionic cells are absent. Tetrasporangia are restrictedto the apical parts of the branches, are decussately tocruciately divided and ovoid (20 µm long, 12 µmwide). Gametophytes were not observed.

Ecology: Growing in lower intertidal pools and thesublittoral fringe.

Specimens: KZN 103: Durban, Treasure Beach(3.viii.1999); KZN 1241: Cape Vidal (9.vii.1999);KZN 379: Mabibi (9.viii.1999).

Discussion: The genus Carpopeltis, comprisingabout 11 species, is mainly confined to the warmerwaters of the Indo-Pacific Ocean. Carpopeltis mail-lardii (Montagne et Millardet) Chiang and C. beckeriSchmitz are known from South Africa.The latter rep-resents an invalidly published manuscript name, thespecimens on which it is based not belonging to Car-popeltis according to Papenfuss (see Silva et al. 1996).Carpopeltis phyllophora bears little resemblance toC. maillardii; the former being a rather large and sup-ple species, the latter a small, cartilaginous speciesthat branches profusely near the apices [Okamura1909, as C. rigida (Harvey) Schmitz; Børgesen 1943,as C. rigida; Chiang 1970]. Comparison of the SouthAfrican specimens with the type specimen (BM, s.n.),collected by Dr Jeannerett at Port Arthur, Tasmania,reveals them to be morphologically very similar. Itshould be noted, however, that the South Africanspecimens are rather variable with respect to thalluswidth, with some being as narrow as 1–2 mm, where-as others are 5–7 mm wide.

Plocamiales

Plocamiaceae

Plocamium cf. mertensii (Greville) Harvey, 1849(1847–1849): 122. Figs 46–49

Type locality:Australia.Description:Thalli are 9 (–17) cm tall, red in colour

and alternately pinnately branched (Fig. 46). Individ-ual axes, 2–3 mm wide, have a plumose aspect, con-


secutive branchlets being of the same length. Later-als have a tendency to produce narrow pinnules, giv-ing the fronds a distinctive fimbriate aspect (Fig. 47).Ramuli form an alternate-distichous series of onesimple and one compound branch, cut off at an angleof 45 ° (–60 °) (Fig. 48). Simple ramuli are acerose,mostly straight, 2.5–3 (–4) mm long and 0.5–0.7 mmwide. Margins of the ramuli are smooth. The centralaxial filament is fairly visible after staining. Apicesare slightly to strongly incurved. Clusters of tetraspo-

rangial stichidia occur as adventitious axes in the ax-ils of the ramuli or replace compound ramuli(Fig. 49). Individual stichidia are oblong and curved,up to 500 µm long and 80–20 µm wide, very occasion-ally branched at their apices. No sexual thalli werefound.

Ecology: Occurring in the lower-intertidal poolsand the shallow subtidal of northern Kwazulu-Natal.

Specimens: KZN 327: Sodwana Bay, intertidal(9.viii.1999); KZN 374: Mabibi (9.viii.1999); KZN

Figs 46–49. Plocamium mertensii.Fig. 46. Habit of herbarium specimen (scale = 2 cm). Fig. 47. Detail of the apical portion of a distinctively fimbriate axis(scale = 1 cm). Fig. 48. Detail of an apex (scale = 1 cm). Fig. 49. Detail of tetrasporangial stichidia (scale = 250 µm). Fig. 52.

Figs 50–53. Plocamium telfairiae.Fig. 50. Habit of type-one thallus (scale = 1 cm). Fig. 51. Habit of type-two thallus (scale = 1 cm). Fig. 52. Habit of type-threethallus (scale = 1 cm). Fig. 53. Detail of dendroid tetrasporangial stichidia (scale = 250 µm).

Fig. 54. Galaxaura rugosa. Habit of herbarium specimen (scale = 1 cm).

589: Rabbit Rock (13.viii.1999); KZN 761: Kosi Bay(16.viii.1999).

Discussion: This species has hitherto only beenrecorded from southern Australia. The distinctivemorphology of the tetrasporangial stichidia and thefimbriate aspect of the axes of the South Africanspecimens agree closely with the description of thisspecies as set out in Womersley (1994). On the otherhand, no serrations on the abaxial edge of the simpleramuli were observed. This character, however, doesnot appear to be fixed in the species. Also, the ten-dency of the proximal branchlets in an alternating se-ries to become divided to a greater or lesser extentwas not observed as typical of the Australian plants(G.T. Kraft, pers. comm.).This species is distinctly dif-ferent from other South African species that have analternating-distichous series of two, having relativelybroad axes and relatively long, slender, clusteredstichidia.

Plocamium telfairiae (W. Hooker et Harvey) Harveyex Kützing, 1849: 885. Figs 50–53

Type locality: Mauritius.Description: Thalli are more or less pyramidal in

outline, rose-red in colour, to 9 cm high. The mainaxes, 1 (–2) mm wide, are alternate-distichouslybranched at an angle of 25–70 °. Ramuli form an al-ternate-distichous series of one simple and one com-pound branch. The latter are strongly incurved nearthe apices. Stellate or dendroid tetrasporangialstichidia are formed in the axils of the ramuli or re-place a compound ramulus.

Ecology: Growing in intertidal rock pools and thesublittoral fringe to a depth of 15 m in northernKwazulu-Natal.

Specimens: KZN 1069: Cape Vidal (9.vii.1998);KZN 249: Sodwana Bay, 2 Mile Reef (8.viii.1999);KZN 271: Sodwana Bay, Adlam’s Reef (9.viii.1999);KZN 329: Sodwana Bay, intertidal (9.viii.1999); KZN585: Rabbit Rock (13.viii.1999); KZN 626: KZN 666:Black Rock (14.viii.1999); Sexton Reef (14.viii.1999);KZN 762: Kosi Bay (16.viii.1999).

Discussion: Three forms could be discerned for P.telfairiae, distinguished on the branching angle anddifferences in size of the axes.The density of branchingis similar in type-one (Fig. 50) and type-two (Fig. 51),but the angle of branching is more acute in type-one(25–) 30–45 ° (–55), as opposed to (30–) 40–55 °(–70). This, combined with a narrower distance be-tween consecutive branches, results in a greater de-gree of overlapping ramuli in type-one thalli. Simpleramuli are subulate to triangular, mostly straight toslightly incurved (rarely recurved). Simple ramuli intype-one are generally longer (1.5–2.5 mm) andbroader (0.3–0.5 mm) than those of type-two (1–1.5and 0.3 mm respectively). Compound ramuli arestrongly incurved at the apex, with moderate overlap-ping in type-one and none in type-two. Type-three


(Fig. 52) has a similar outline and structure,but is gen-erally smaller (to 4.5 cm) and wider (1.5–2 mm). Theramuli of the latter are sub-pinnate, with a compoundramulus subopposite a simple one. The angle ofbranching lies between that of type-one and type-two(40–50 °). Similarly, the simple ramuli sizes lie be-tween type-one and type-two, although they have awider base (0.5–1 mm). The central axial filament isnot visible in type-one and type-two (or only near theapex), whilst it is in type-three. The tetrasporangialstichidia were found in type-one and type-three,whereas only female material was collected for type-two.The former occur in the axils of ramuli or replac-ing compound ramuli and having a stellate / tree-shape(Fig. 53).

The specimens examined agree with descriptionsand herbarium specimens of this species (includingan isotype specimen in BM).This is the first record ofthis species in South Africa, although Seagrief (1984)and Cormaci et al. (1991) credited this species toSouth Africa based on supposed records of Simons(1964).The latter, however, reported the species onlyfrom Mozambique and not from South Africa. Thisspecies has a wide distribution within the Indian andPacific Oceans. There is also an Atlantic record fromGhana by Lawson and John (1987).

Nemalionales

Galaxauraceae

Galaxaura rugosa (Ellis et Solander) Lamouroux,1816: 263. Fig. 54

Type locality: Jamaica.Description: Thalli are erect, to 8 cm high, hirsute

and calcified. The axes are terete, dichotomouslybranched, 2 mm in diameter and densely covered byassimilatory filaments.The medulla consists of entan-gled filaments (12–16 µm in diameter). The cortex isfilamentous with both short and long assimilatory fil-aments, which arise from inflated proximal cortexcells (33–42 µm in diameter). The cruciately dividedtetrasporangia are borne apically on short assimilato-ry filaments. They are ovoid, 25–30 µm long and to20 µm wide. Gametophytes were not observed.

Ecology: Epilithic in the lower intertidal zone.Specimens: KZN 326: Sodwana Bay, intertidal

(9.viii.1999); KZN 466: Mabibi (11.viii.1999); KZN1685: Island Rock (14.viii.2000).

Discussion: Despite taxonomic confusion in thepast (see Huisman and Borowitzka 1990), tetra-sporophytes of G. rugosa are easily recognisable bytheir hirsute habit. Several species of Galaxaura andthe morphologically similar genus Tricleocarpa areknown from South Africa. Galaxaura diesingiana Za-nardini, G. magna Kjellman and G. marginata (Elliset Solander) Lamouroux are all complanate. Galax-aura obtusata (Ellis et Solander) Lamouroux and Tri-cleocarpa fragilis (Linnaeus) Huisman et Townsend


are both terete, but are never hirsute and always lackthe inflated proximal cells of assimilatory filaments.Galaxaura rugosa is widespread in the tropical Indi-an Ocean.

Discussion

Distribution patterns of tropical seaweeds are incom-plete for many taxa. Several regions have receivedvery little attention and remain seriously un(der)ex-plored. There are not only limited data on smallislands and archipelagos but also for several conti-nental coastal regions such as Oman, Iran and Mada-gascar.This makes biogeographical assessments diffi-cult, but a number of clear trends are indicated(Table I).

The majority of the Phaeophyta and Rhodophytareported in this paper consist of widespread tropicalspecies. Asteronema breviarticulatum, Dictyota cervi-cornis, D. ciliolata and Galaxaura rugosa are mainlypantropical species, occurring in the Atlantic(Caribbean Sea) and Indian Oceans as well as the Pa-cific Ocean. Some are present along the WesternAustralian coast. Next to the pantropical taxa, sever-al species have a defined Indo-Pacific or IndianOcean distribution: Ceramium cingulatum, Euptilotafergusonii, Halymenia durvillei and Phacelocarpustristichus. The presence of pantropical and Indo-Pa-cific species along the northern part of the Kwazulu-Natal coast is not surprising, as several of these

species have been reported from Inhaca in southernMozambique. Their presence in the northern part ofthe Kwazulu-Natal coastline, therefore, only indi-cates a minor range extension. In contrast, the distri-bution range of Digeneopsis subopaca remains re-stricted to northern Kwazulu-Natal and the InhacaPeninsula in Mozambique.Together with Dasycladusramosus Chamberlain (1958), these species are prob-ably endemic to the northern part of the overlap re-gion between warm-temperate South African watersand the tropical Indian Ocean.

Balliella crouanioides, Gibsmithia hawaiiensis,Predaea weldii and Hypoglossum minimum are char-acterised by disjunct tropical Indo-Pacific distribu-tion patterns. All three were originally describedfrom the central or western Pacific Ocean and havebeen reported from a few scattered localities withinthe Indian Ocean. Their small size and subtidal habi-tat probably contribute to the scarcity of reports. Bal-liella crouanioides and Gibsmithia hawaiiensis haverecently been reported for the East African coast(Coppejans et al. 2000). Their presence along theSouth African coast may indicate a wide distributionwithin the Indo-Pacific region.

Carpopeltis phyllophora and Plocamium mertensiiare both Australian species. Norris and Aken (1985)and Hommersand (1986) regard this South Africanflora as having affinities with the western- and south-ern Australian algal floras, although there has beensome debate on what may have caused these floristicsimilarities. According to Hommersand (1986), the

Table I. Known distributions of the species reported in this paper.

Mozambique Western Central Indonesia- Australia Pacific AtlanticIndian Indian Malaysia Ocean OceanOcean Ocean

Asteronema breviarticulatum – + + + + + +Balliella crouanioides + + – + – + –Carpopeltis phyllophora – – – + + – –Ceramium cingulatum – + – + + – –Dictyota cervicornis + + + + + + +Dictyota ciliolata + + + + + + +Dictyota hamifera – + – + – + +Dictyota rigida – + – – – – –Digeneopsis subopaca + – – – – – –Euptilota articulata – – + – + + –Euptilota fergusonii + + + – – – –Galaxaura rugosa + + + + + + +Gibsmithia hawaiiensis – + – + + + –Halymenia durvillei – + + + + + –Hypoglossum minimum – – + – – + –Padina gymnospora ? + ? ? + ? +Phacelocarpus tristichus + + + – – – –Plocamium mertensii – – – – + – –Plocamium telfairiae + – + – – + –Predaea weldii – – – + + + +

General references: Womersley (1987; 1994; 1998); Millar and Kraft (1993); Silva et al. (1996); Phillips (1997); Wynne(1998);Yoshida (1998);Abbott (1999); Coppejans and Millar (2000); Coppejans et al. (2000); Huisman (2000).

genus Carpopeltis and to a certain extent Plocamiumrepresent examples of species clusters that originatedalong the west and south coasts of Australia and mi-grated to South Africa during mid-to-late Mioceneperiods (16–6Mya) marked by a general decrease intemperature in the tropics. Euptilota articulata mayalso represent an example of the South African-Aus-tralian connection, although the species has also beenreported from India and the North-West Pacific.

The distribution of Padina gymnospora is difficultto interpret due to taxonomic confusion in the past.Studies on the genus Padina along the East Africancoast (Somalia to South Africa) have revealed that P.gymnospora is absent along the tropical coasts ofKenya, Tanzania and the various tropical archipela-gos (Muylle 2000).The species is, however, present inthe northern part of the western Indian Ocean (So-malia, Yemen and Oman) as well as along the coastof Kwazulu-Natal. The distribution pattern is inter-esting as both geographically isolated populationsare in habitats characterised by slightly cooler watertemperatures than occur in the tropical IndianOcean. Similar examples of disjunct distribution pat-terns between the northern Arabian Sea and theSouth African east coast are offered by: Dictyopteris


macrocarpa (Areschoug) Schmidt, Ecklonia radiata(C. Agardh) J. Agardh and Pseudocodium de-vriesiiWeber-van Bosse (Barratt et al. 1984, Barratt et al.1986,Wynne 1999).

Acknowledgements

We would like to offer our sincere gratitude to JeanHarris, Nonhlanhla Nxumalo, Bridget Armstrong,Cloverly Lawrence and John Dives, of the Kwazulu-Natal Nature Conservation Services, for their fullsupport in organising everything, everytime, every-where. We also extend our thanks to Peter Timmfrom Triton Divers for his smoothly run services.Funding for this project was provided by the Interna-tional Scientific and Technological Cooperation(BIL98/84) between the Ghent University and theUniversity of Cape Town and FWO Research Project(3G002496). Further support was provided in SouthAfrica by Marine and Coastal Management, the Na-tional Research Foundation and the Department ofEnvironment and Tourism.

Accepted 19 May 2002.

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