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

O. D e Clerci©'*, H. R. Engledow3, J. J. Boltonb, R. J. Anderson0 and E. Coppejans" a Research Group Phycology, Biology Department, Ghent University, Krijgslaan 281 / S8,9000 Ghent, Belgium b Botany Department, University of Cape Town, Private Bag, 7700 Rondebosch, South Africa c 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 of 6 taxa of Phaeophyta and 14 Rhodophyta. Most species have a pantropical or Indo-Pacific distribution and are generally known from several localities in the Indian Ocean (e.g .Asteronema breviarticulatum, Ceramium cingulatum, 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, some species 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.

Introduction coast marine provinces has been well studied and characterised by a clearly defined overlap zone The South African coastline displays a unique floral (Bolton and Stegenga 1990, Jackelmanet al. 1991), in diversity, with biota ranging from kelp beds on the contrast to the eastern border of the Agulhas (south west coast to coral reefs near the Mozambican bor coast) marine province which is less clear. Seawater der. The correlation between surface seawater tem temperature, the main defining ecological parameter peratures and shore community compositions has in the distribution of seaweeds, gradually rises from been investigated by several researchers, who have around East London to northern Kwazulu-Natal firmly established that the main defining influences (Bolton 1986). At present, it is believed that a distinct are the warm Agulhas current flowing down the east sub-tropical marine province in Kwazulu-Natal is un coast from Mozambique to the region where it meets likely, as the flora seems to comprise an eastwardly the cold Benguela current (e.g. Stephenson 1948, declining number of Agulhas-province species, these Bolton 1986, Bolton and Anderson 1990). Stephenbeing replaced largely by Indo-West pacific species son (1948) divided the South African coastline into with the increase in water temperature (Bolton and three marine provinces: the cold-temperate west Anderson 1997). It should be noted, however, that coast, warm-temperate south coast, and sub-tropical our knowledge of the marine flora of Kwazulu-Natal east coast. More recently, seaweed biogeographers, is limited. Apart from a limited number of taxa, stud although also recognising three floral provinces, have ied in detail by students of G.F. Papenfuss in Berke described them differently. The seaweed flora of the ley (California) and a field guide on the seaweeds of west coast is regarded as warm-temperate by Bolton Maputaland by Seagrief (1980), no detailed studies (1986) and Timing (1990) or cool-temperate by were available relating to Natal algae prior to 1985. Emanuel et al. (1992) and Bolton and Anderson In the eighties and early nineties R.E. Norris and co (1997), rather than cold-temperate. The presence of a authors published a series of papers on the algae of distinct sub-tropical flora on the east coast (Kwazu- Kwazulu-Natal that gave some evidence that there is lu-Natal) is not accepted by either Timing (1990) or indeed a distinct flora in the region. In total they (see Bolton and Anderson (1997). These authors consider Norris 1992 and Silvaet al. 1996 for a complete refer the seaweeds of the east coast of South Africa to be ence list) reported 63 species as new records for the westernmost extension of the large Indo-West South Africa and described 23 new species, many of Pacific tropical region, and cast doubt on the exis which are currently only known from Kwazulu- tence of a distinct sub-tropical flora in Kwazulu-Na Natal. It has also been noted that elements of the tal. Kwazulu-Natal marine algal flora have a link with The transition zone between the west and south southwestern Australia (Norris and Aken 1985, 414 O. De Clerck et al.

Hommersand 1986). Kwazulu-Natal may therefore species which are newly recorded for South Africa. be more than a transition zone between the Agulhas Records on Chlorophyta have recently been pub and Indo-West Pacific floras, in that it actually consti lished separately by Leliaert et al. (2001). tutes a distinct flora with warm-temperate south western Australian affinities and a large number of endemics. Material and Methods In a joint research project conducted by the Botany Departments of the Ghent University (Bel Approximately 2400 specimens were collected over a gium) and the University of Cape Town (South period of two years (July 1998; August 1999; Decem Africa), the seaweeds of Kwazulu-Natal are being ber 1999; July-August 2000; February 2001), as a re studied from a biogeographical perspective. Several sult of a Bilateral Scientific and Technological Coop sites along the coast from Port Edward in the south eration Project between the Flemish community to Kosi Bay near the Mozambican border were sam (Belgium) and South Africa. Specimens were collect pled mainly between July 1998 and February 2001. ed at various sites distributed along the entire Kwazu The collections are now being identified and will lu-Natal coast (see Fig. 1) and were immediately eventually serve to provide a better understanding of processed as herbarium specimens, as well as being the distribution and floristic affinities of the algae partly preserved in 4 % formaldehyde/seawater. The along the Kwazulu-Natal coast. This paper reports 20 specimens, bearing serial numbers prefixed by ‘KZN’

Mozambique O Maputo

Swaziland — Rabbil Rock Kosi Bay — N23 — Sexton Reef Bhanga Nek — Linkia Reef South Africa Black Rock — Tiger Reef Island Rock Mablbi i 1/4 Mileh R eef 1— 2 .7 .9 Mile R eef Sodwana Bay f— AdlamAdlai s Reef * Deep D i i f i r Sponge Cape Vidal St. Lucia Mission Rocks

Mapelane

Zinkwazi

Shaka's Rock

Durban The Bluff Treasure Beach Umkomaas Aliwal Shoal

Scottsburah — Protea Banks — Orange Rocks Shelly Beach — — Boboyi Reef — Uvongo Reef Broker Reef .Trafalgar Port Edward Palm Beach 100 km

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 parts of the thallus and recurved branchlets; speci rial was stained in a mixture of 1 g aniline blue pow mens from exposed habitats generally form low, der, 50 mL Karo®, 45 mL distilled water, 5 mL acetic dense mats and lack slender straps. The South acid. Collections other than our own, those of R.E. African specimens represent the typical growth form Norris and co-workers (bearing serial numbers pre from exposed habitats. A major difference to previ fixed by ‘NAT’) deposited in UN and various other ous observations is the occurrence of iridescence. collectors in BOL were also examined. Herbarium Despite the examination of over 200 specimens by abbreviations follow Holmgrenet al. (1990). Digital De Clerck (1999), mainly from the East African coast images were taken, using an Olympus DP50 digital (Tanzania), iridescence was never observed. The tax camera (Melville, USA) mounted on a Leitz Diaplan onomic value and intraspecific variability of irides or Wild M10 microscope (Wetzlar, Germany). cence, however, remains poorly understood (Gaillard 1972).

Results Dictyota ciliolata Sonder ex Ktitzing, 1859:12, pi. 27, fig.l. Figs 5,6 Phaeophyta Type locality: La Guaira, Venezuela. Dictyotaceae Description: Thalli are erect, to 8 cm tali, attached Dictyota cervicornis Ktitzing, 1859:11,pi. 24, fig. 2. by means of a single stupose holdfast (Fig. 6). Figs 2-4 Stolonoidal fibres are absent. Straps, 2-3 mm wide, are slender and dichotomously branched (Fig. 5).The Type locality: Key West, Florida. margins are dentate, rarely smooth, while the surface Description: Thalli form low, dense mats (Fig. 2). is always smooth. The apices are rounded. The The individual specimens are to 5 cm tali and com medulla and cortex are uniformly one-layered. Spo posed of coarse,2-3 mm wide, subdichotomous rangia are single, scattered on both surfaces, but ab straps (Fig. 4). The margins are smooth, but the sur sent in the apical dichotomies. Sporangia, 95-110 pm face is beset with acute proliferations (Fig. 3). The in diameter, are borne on a single stalk cell and are apices are obtuse. Sporangia are scattered on both not surrounded by a conspicuous involucrum. Ga surfaces, but absent in the apical dichotomies. Spo metangia were not observed. rangia, 120-135 pm in diameter, are placed mostly Ecology: Specimens were collected in intertidal solitary and are borne on a single stalk cell, surround pools of the sublittoral fringe down to -12 m. ed by a conspicuous involucrum. Gametangia were Specimens: KZN 396 and KZN 401: Mabibi not observed. In situ the colour is brown with a blue- (9.viii.l999);KZN 2182: Mabibi (13.ii.2001). greyish iridescence. Discussion: Dictyota ciliolata is characterised by its Ecology: Specimens occur in the sublittoral fringe, stupose holdfast, dentate margins and the absence of generally on wave-exposed shores. stolonoidal fibres, although the margins of some Specimens: Simons 1262 (BOL): Kosi Bay specimens can be nearly smooth (Hornig et al. (30.vi.1965); Simons 1254 (BOL): Sodwana Bay 1992a,b). Several other dentate species of Dictyota (3.VÜ.1965); KZN 312: Adlam’s Reef Sodwana Bay occur in the Indo-Pacific and the Caribbean regions, (9.viii.l999);KZN 579:Rabbit Rock (13.viii.1999); as detailed by De Clerck and Coppejans (1999). KZN 736: N23 (15.viii.1999);KZN 1013: Cape Vidal Along the South African coast D. ciliolata is most (9.VÜ.1998);KZN 1014: Mission Rocks(8.vii.l998). similar to D. liturata J. Agardh and D. suhrii G. Mur Discussion: Dictyota cervicornis is perhaps the ray. The latter species, however, has a multilayered most commonDictyota species in the Indian Ocean, medulla near the margins, whereas the medulla of D. characterised by surface proliferations and sporangia ciliolata is uniformly one-layered. Dictyota ciliolata that are surrounded by an involucrum. It appears to differs from D. liturata in the placement of the spo be closely related to D. crispata Lamouroux and D. rangia, which are arranged singly or distributed in magneana De Clerck et Coppejans (De Clerck and longitudinal lines, respectively. Coppejans 1997,Coppejans et al. 2001), but differs mainly in the placement of sporangia, these being ab Dictyota hamifera Setchell, 1926:92, pi. 14, sent or present in the apical dichotomies respectively. figs. 1-6. Fig. 7 Other characters which distinguish D. crispata from D. cervicornis include the morphology of the apices Type locality: between Papenu and Huau, Tahiti. and the abundance and placement of surface prolif Description: The thallus is repent, up to 3 cm, at erations. Detailed comparisons between the three tached to hosts at various points by patches of rhi- species are made by Coppejans et al. (2001). Dictyota zoids. The straps, 1-2 mm wide, are dichotomously cervicornis exhibits a broad range of morphological branched. The margins are smooth except for the fal growth forms, which can often be related to its habi cate branchlets (Fig. 7), which may resemble teeth. tat. Specimens growing in the shallow subtidal are of The surface is smooth. The apices are rounded to tri- ten characterised by long slender straps in the distal dentate. The medulla and cortex are uniformly one- 416 O. De Clerck et al. Twenty marine benthic algae new to South Africa 417 layered. Sporangia or gametangia were not ob G. Murray and the Australian D. fastigiata Sonder. served. The latter two species, however, are characterised by Ecology: Specimens were collected at a depth of 18 a multilayered medulla, at least in the basal parts of to 25 m epiphytic on Plocamium sp. the thallus. Specimens: KZN 1617: Sodwana Bay, 2 Mile Reef ( 12.viii.2000). Padina gymnospora (Ktitzing) Sonder, Discussion: Dictyota hamifera was for a long time 1871:47. Figs 11,12 only known from the Pacific Ocean. Hornig et al. Type locality: St. Thomas, Virgin Islands. (1992a) reported the species for the first time from the Description: Thalli, attached by a small stupose rhi- Atlantic Ocean, while De Clerck ( 1999) and Coppejans zoidal base, are to 6.5 cm high, flabellate and com eta!. (2000) recorded the species from the East African posed of several lobes with inrolled margins (Fig. 11). coast (Kenya and Tanzania ). Dictyota hamifera is easily The superior surface (the side toward which the mar recognised by its falcate, swollen branchlets. gin is inrolled) is slightly to moderately calcified. Thalli are olive-brown in colour. In transverse sec Dictyota rigida De Clerck et Coppejans, 1999: tion the thallus is composed of 4-6 layers of cells in 189-191, figs. 10-17. Figs 8-10 the mid-regions and up to 8-9 layers near the base. Type locality: Kunduchi, Dar es Salaam,Tanzania. Hair rows are present on both sides of the thallus, but Description: Thalli are up to 6 cm high, attached by more conspicuous on the inferior side of the thallus. a small stupose holdfast. The dichotomously Tetrasporangia are arranged in concentric lines, branched straps are narrow (0.8-1.2 mm) and of uni above each hair row, mainly (but not exclusively) de form width throughout the thallus (Figs 8, 10). The veloped on the superior surface. An evanescent indu- margins and surface are smooth. The medulla and sium is present but is usually only clearly visible in cortex are uniformly one-layered. Sporangia form a very young sori (Fig. 12). Tetrasporangia are ovoid, well-defined line in the median part of the straps, but up to 100 pm long and 70 pm wide. Gametophytes are not present along the margins (Fig. 9). Sporangia, were not observed. 80-95 pm in diameter, are not surrounded by an in Ecology: Specimens are collected in shallow pools volucrum and are borne on a single stalk cell. Ga in the sublittoral fringe. metangia were not observed. Specimens: KZN 94: Durban, Treasure Beach Ecology: Specimens were collected in pools of the (3.viii.l999); KZN 294: Sodwana Bay, Adlam’s Reef sublittoral fringe. (9.viii.1999); KZN 361: Mabibi (9.viii.l999); KZN Specimens: KZN 311: Sodwana Bay, Adlam’s Reef 708: Bhanga Nek, N23 (15.viii.1999); KZN 745: Kosi (9.viii.l999); KZN 385: Mabibi (9.viii.l999). Bay (16.viii.1999). Discussion: Dictyota rigida was recently described Discussion: Padina species are distinguished pri from Tanzania (De Clerck and Coppejans 1999) and marily on the number of cell layers, the distribution subsequently reported from Kenya (Coppejans et al. of hair bands, position of the sporangial sori (on the 2000). Characters defining D. rigida include the erect superior versus inferior surface) and the type of al growth form, the stiff texture and the single stupose ternations of hair and sporangial bands. Padina gym holdfast (De Clerck and Coppejans 1999).Dictyota nospora is characterised by the thick thallus (up to 8 rigida is most similar to the South African D. suhrii cell layers), the distribution of sporangia primarily on

Figs 2 -4 . Dictyota cervicornis. Fig. 2. H abit (scale = 1 cm). Fig. 3. D etail of the surface with multiple proliferations (scale = 2 mm). Fig. 4. D etail of the api cal parts of the thallus (scale = 2 mm).

Figs 5 ,6 . Dictyota ciliolata. Fig. 5. H abit (scale = 1 cm). Fig. 6. D etail of the stupose base (scale = 2 mm). Fig. 7. Dictyota hamifera. H abit showing falcate branchlets (arrows) (scale = 2 mm). Figs 8-10. Dictyota rigida. Fig. 8. H abit (scale = 1 cm). Fig. 9. Tetrasporangia aggregated in the median region of the straps (scale = 1 mm). Fig. 10. D e 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 tetrasporangial sorus (scale = 100 pm). Figs 13-15. Asteronema breviarticulatum. Fig. 13. Habit of herbarium specimen illustrating the frayed, rope-like appearance (scale = 1 cm). Fig. 14. Detail of a hooked branch (scale = 50 pm). Fig. 15. D etail of an ovoid plurilocular sporangium (scale = 50 pm). 418 O. De Clerck et al. the superior surface of the thallus and the evanescent Type locality: Mage Island, Japan. indusium (Allender and Kraft 1983, Womersley Description: Thalli form woolly tufts composed of 1987). The species is most similar toP. crassa Yanta uniseriate filaments to 40 mm high (Fig. 16). Fila da, the latter supposedly having sporangia only on ments are opposite-distichously branched with inde the lower side of the thallus. Only P. boryana Thivy terminate branches being formed every (1-) 2 (-5) and P.plumbea (Areschoug) Levring have previously segments. The apical parts of the axes show a distinc been reported from South Africa (Levring 1940, tive sinusoidal curving (Fig. 17). Corticating rhizoids Gaillard 1975). Both species are characterised by a 2- arise from the lower cells of the laterals and form a layered thallus and hence are easily distinguished loose rag-like cortex (Fig. 19). The periaxial cells are from P. gymnospora. Due to confusion with P. 100-115 pm in diameter and bear spherical vesicular boergesenii Allender et Kraft in the past, the distribu cells, 8-15 pm in diameter on their abaxial side tion ofP. gymnospora is difficult to assess in the Indi (Fig. 18). Decussately divided tetrasporangia develop an Ocean (Silva et al. 1996). The latter, however, has on the adaxial side of the periaxial cells of the later been reported with certainty from Australia and als. Tetrasporangia are ovoid, 30-55 pm long and Oman (Womersley 1987, Wynne and Jupp 1998). 30-38 pm wide. Gametophytes were not observed. Ecology: Balliella crouanioides is a common com Incertae sedis ponent of the sublittoral zone (-20 to -35 m), where it forms distinctive orangy-iridescent tufts on a vari Asteronema breviarticulatum (J. Agardh) ety of substrates (coral debris, various algae, sponges, Ouriques et Bouzon, 2000:271. Figs 13-15 etc.). Type locality: San Agustín, Oaxaca, Mexico. Specimens: KZN 63 Aliwal Shoal (3.viii.l999); Description: Thalli, approximately 2.5 cm tali, form KZN 147: Aliwal Shoal (4.viii.l999);KZN 2093: Sod tufts of interwoven filaments, which have the appear wana Bay, Deep Sponge (10.ii.2001); KZN 250 and ance of a frayed rope at its tips (Fig. 13). Filaments KZN 257: Sodwana Bay, 2 Mile Reef (8.viii.l999); are uniseriate and irregularly branched with numer NAT 6232 (UN): Sodwana Bay, 2 Mile Reef ous short hooked branches arising at an angle of (14.X.1989); KZN 644: Bhanga Nek, Sexton Reef 80-90 ° (Fig. 14). Growth takes place by intercalary (14.viii.1999). cell divisions. Cells are 25-33 pm wide, 60-80 pm Discussion: Balliella species are recognisable by long, L/B: 1-1.3. Plurilocular sporangia, placed on a the presence of spherical vesicular cells (‘gland 1-2-celled stalk, are spherical to ovoid and measure cells’), which occur adaxially or abaxially on the peri 40 X 42 (-63) pm on average (Fig. 15). axial cells. Species are separated on the basis of their Ecology: growing in the intertidal, often attached habit, branching pattern, position and size of vesicu to barnacles in wave-exposed localities. lar cells and tetrasporangia. Wollaston (1984) report Specimens: KZN 451: Mabibi (ll.viii.1999); KZN ed B. subcorticata (Itono) Itonoet Tanaka from 654: Black Rock (14.viii.1999); KZN 728: N23 Mozambique, but Huisman (1988) expressed the (15.viii.1999). opinion that the specimen depicted was more similar Discussion: Asteronema breviarticulatum is a wide to B. crouanioides, based on the overall branching spread warm-temperate to tropical species charac pattern, arrangement of the tetrasporangia and size terised by the shape of the plurilocular sporangia and of the vesicular cells. Comparison of our material the presence of short hooked branches. Ouriques and with the specimen studied by Wollaston (UC Bouzon (2000) transferred this species, which was 1470004) reveals that they are identical and supports commonly known as Hincksia breviarticulata (J. Huisman’s proposition. Athanasiadis (1996) provi Agardh) Silva, to Asteronema on the basis of the stel sionally assigned a collection of specimens from late chloroplasts. Recent molecular studies by Drais- Kenya to B. crouanioides. Re-examination of the ma et al. (2001) and Rousseau et al. (2001) reveal, type collection proved to be necessary as some con however, that Asteronema is polyphyletic and not re troversies exist on the branching pattern and place lated to the Ectocarpales sensu lato. Unfortunately ment of the gland cells in this species. The South Asteronema breviarticulatum itself was not included African specimens match previous accounts of B. in the analysis. Pending further studies, A. breviartic crouanioides very well (Itono and Tanaka 1973, Itono ulatum is listed as incertae sedis rather than under the 1977, Huisman and Kraft 1984), although variation Ectocarpales. was observed in both the size (8-15 pm in diameter) and placement of the gland cells (abaxial but occa sionally adaxial). As these characters are regarded to Rhodophyta be of prime importance in delineating Balliella Ceramiales species, further research is needed on intraspecific variation. Ceramiaceae Balliella crouanioides (Itono) Itonoet Tanaka, 1973: Ceramium cingulatum Weber-van Bosse, 1923: 250. Figs 16-19 332-333, figs. 123,124. Figs 20-22 Twenty marine benthic algae new to South Africa 419

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 pm). Fig. 18. Cruciately divided adaxial tetrasporangia and abaxial gland cells (arrowheads) on the periax ial cells (scale = 25 pm). Fig. 19. Loose cortication at the nodes of the axes in the lower part of the thallus (scale = 100 pm).

Figs 20-22. Ceramium cingulatum. Fig. 20. H abit of the typically clavate axis (scale = 500 pm). Fig. 21. Whorled tetrasporangia (scale = 100 pm). Fig. 22. Detail of the nodal structure in a proximal axis showing two anterior (arrowheads) and two posterior (arrows) cortical filaments (scale = 100 pm).

Figs 23-26. Euptilota articulata. Fig. 23. Habit of herbarium specimen (scale = 1 cm). Fig. 24. Apical portion with alternate-distichous branching (scale = 100 pm). Fig. 25. Ultimate branchlet bearing lateral and term inal tetrasporangia (scale = 100 pm). Fig. 26. D etail of an ultimate branchlet with tetrasporangia (scale = 50 pm).

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 pm). Fig. 29. D etail of a tetrasporangial branchlet (scale = 100 pm). 420 O. De Clerck et al.

Type locality: Sape Strait, south of Sangeang Island, formed irregularly and repeat the branching pattern Indonesia. of the main axes. Tetrahedrally divided tetrasporan Description: Thalli to 2 mm high consisting of un gia (40-52 pm in diameter) are borne laterally or ter branched erect axes, that arise mainly from a single minally on the ultimate branchlets (Fig. 26). Gameto- point or from a creeping filament. Axes are some phytic plants were not observed. what clavate, measuring 90-100 pm wide near the Ecology: Euptilota articulata has only been collect base, 140-170 pm in the middle part of the thallus, ed once in situ, at a depth of 35 m in southern Natal. ending abruptly in an attenuate apex (Fig. 20). Axial The specimen in the Norris Herbarium (UN) was cells are lenticular, usually wider than long except collected from the drift. near the base. A total of 8 periaxial cells are present Specimens: KZN 1962: Protea Banks, Northern per segment, each producing 2 anterior and 2 posteri Pinnacle (5.Ü.2001); NAT 1310: Palm Beach or cortical initials of cortical filaments. Anterior ini (14.V.1983). tials cut off 1-2 acropetal filaments, which are up to 2 Discussion: The genus Euptilota contains at pres cells long (Fig. 22). Posterior initials either remain ent four species, all of which are characterised by an undivided or produce a single basipetal filament alternately-distichously branched thallus, in which (Fig. 22). Cortication remains incomplete, but adja axial cells each bear a single lateral (Millar 1990, cent cortical bands may be nearly confluent. Womersley 1998).Euptilota articulata differs from Tetrasporangia form in distinctive whorls, are slightly E. pappeana Ktitzing, the only species reported protruding and measure 28-35 pm in diameter from South Africa (Stegenga et al. 1997),in the pin (Fig. 21). Gametangia were not observed. nate branching of the determinate laterals as op Ecology: Epiphytic on Plocamium telfairiae at -18 posed to laterals, which are either unbranched or to 25 m. once to twice dichotomously branched in the latter Specimens: KZN 1619.2: Sodwana Bay, 2 Mile species. Euptilota articulata has previously been re Reef (12.viii.2000). ported for Australia (Millar 1990,Womersley 1998), Discussion: The South African specimens agree Japan (Itono 1977)and India (Umamaheswara Rao very well with the original description by Weber-van 1974). Bosse (1923) and a subsequent account of the species by Cormaci and Furnari (1991), who compared C. Euptilota fergusonii Cotton,1907:262-264. cingulatum with the Mediterranean C. giacconei Cor Figs 27-29 maci et Furnari. The only difference we observed was the mode of attachment. Typically, C. cingulatum has Type locality: ‘Pantura’, Sri Lanka. erect axes in the terminal parts or produces erect Description: Thalli are erect, to 15 cm high, pin branches. South African plants were usually strictly nately branched up to 5 orders and lack a percurrent erect, but the presence of rhizoids in the lower part of main axis (Fig. 27).The holdfast is densely rhizoidal. the thallus indicates the possibility of a creeping Axes are corticated from close to the apices with cor- growth form. Strictly erect versus ascending growth ticating filaments arising from the periaxial cells, ap forms are not considered to be major taxonomic difproximately 12-14 cells proximal to the apex. Proxi ferences in Ceramium (Dixon 1960, Womersley mal axial cells are 450-500 pm wide and 240-270 pm 1978).Ceramium cingulatum is unique among South long. Each axial cell bears a single lateral in a African representatives of the genus in its un distichous-alternate pattern (Fig. 28). Determinate branched erect axes. The species has been reported in branchlets are ecorticate, curved upwards and up to the Indian Ocean from Somalia, Tanzania and the 12 cells long. The basal 3 cells of a determinate lateral Seychelles (Jaasund 1970,Sartoni 1975, Wynne 1995). bear each a short adaxial side branch, the fourth to the ninth cell bear an arched abaxial branchlet and the remaining cells remain unbranched, except for Euptilota articulata (J. Agardh) Schmitz, 1896:7. the ultimate and penultimate cells, which bear 1-2 Figs 23-26 (-4) spine-like cells (Fig. 29). Indeterminate branch Type locality: Australia. lets are formed irregularly and repeat the branching Description:Thalli are erect, to 25 cm high and pin- pattern of the main axes. Tetrahedrally divided nately branched up to 5 orders (Fig. 23). A percurrent tetrasporangia (60 pm long, 40-50 pm wide) are main axis is lacking. The holdfast is densely rhizoidal. borne laterally on cells of the ultimate branchlets. Axes are corticated from close to the apices, with cor- Gametophytes were not observed. ticating filaments arising from the periaxial cells 5 to Ecology: Collected at two occasions only, at a 10 cells proximal to the apex. Near the base the axes depth of 15-25 m in northern Kwazulu-Natal (Sod attain a width of 1.5 mm. Each axial cell bears a single wana area). lateral in a distichous-alternate pattern (Fig. 24). De Specimens: KZN 342: Sodwana Bay, 2 Mile Reef terminate branchlets are ecorticate, straight, 12-17 (9.VÜL1999);KZN 419: Sodwana Bay, 7 Mile Reef cells long and pinnately branched (Fig. 25). Spine (9.VÜL1999). like cells are absent. Indeterminate branchlets are Discussion: Euptilota fergusonii is distinguished Twenty marine benthic algae new to South Africa 421 from otherEuptilota species in having spine-like cells Rhodomelaceae at the tips of the determinate laterals, the latter addi Digeneopsis subopaca Simons, 1970:10,11. Fig. 34 tionally being secundly, as opposed to alternately or subdichotomously, branched. Euptilota fergusonii Type locality: Santa Maria, Inhaca, Mozambique was originally described from Sri Lanka (Cotton Description:Thalli are erect, very stiff and blackish 1907) and has previously been reported from Tanza in colour, to 11 cm tali (Fig. 33). Indeterminate axes nia and Mozambique (Jaasund 1976, Wollaston are pinnately branched, compressed near the apices, 1984). but become rounded in the lower parts of the thallus. Determinate branchlets are stiff and incurved, Delesseriaceae 4-10 mm long and to 500 pm in diameter, bi- or tri- furcately branched in up to 2 or 3 orders. The thallus Hypoglossum minimum Yamada, 1936:138-140, is polysiphonous, consisting of an axial cell surround fig. 2. Figs 30-33 ed by 5 pericentral cells, which are obscured by a Type locality: Naha, Okinawa-jima, Ryukyu-retto, well-developed cortex from close to the apices. Japan. Tetrasporangia are formed in clustered stichidia near Description: Thalli consist of unbranched, lanceo the bases of determinate laterals. The stichidia are late blades (7-12 mm high and 0.8-2.3 mm broad), 100-190 pm thick, with each segment bearing 2 attached by means of a small discoid holdfast tetrasporangia. Tetrasporangia are up to 115 pm in (Fig. 30). Growth takes place from a single apical cell, diameter and tetrahedrally divided. Gametophytes with a typical type-one apical organisation (i.e. all were not observed. second order row cells produce third order rows). Ecology: Collected in intertidal rock pools and the The midrib remains uncorticated throughout. shallow sublittoral from Mapelane (just south of St. Tetrasporangial sori form discrete, paired ovate sori Lucia) to Mozambique. close to the midrib (Fig. 31), 900-1700 pm long and Specimens: KZN 368: Sodwana Bay, intertidal 400-600 pm wide. Tetrasporangial formation in (9.viii.1999); KZN 569: Bhanga Nek (13.viii.1999); volves the lateral pericentral cells but not the trans KZN 748: Kosi Bay (16.viii.1999); KZN 1641: Mabibi verse ones. Tetrasporangia, 62-80 pm in diameter, (13.viii.2000); KZN 1809: Cape Vidal (18.viii.2000); are cut off from second- and third order cell rows as KZN 1843: Mapelane, Crayfish Point (20.viii.2000); well as from cortical cells, resulting in sporangia that Pocock and Papenfuss 1024 (UN): St. Lucia lie in more than one plane of the blade. Spermatan- (21.vii.1938). gial sori form discrete, somewhat oblique patches Discussion: Digeneopsis subopaca, originally de scattered over the wings (Fig. 32). Female plants pro scribed from southern Mozambique (Simons 1970), duce a single cystocarp on the midrib in the distal re remains a poorly known representative of the gion of the blades (Fig. 33). Rhodomelaceae. In the absence of gametophytic ma Ecology: Collected on a single occasion, epiphytic terial the taxonomic affinities of this monotypic on Carpomitra longicarpa Simons at -43 m. genus are unclear. The morphology of the tetraspo Specimen: KZN 506: Tiger Reef, Bhanga Nek rangial stichidia and the compressed, bilateral nature (13.viii.1999). of the plants, however, seem to indicate a close rela Discussion: Hypoglossum minimum is a small tionship to the Amansieae. The South African record species, which can easily be confused with H. simu of Halopithys incurva (Hudson) Batters in Seagrief lans Wynne, Price et Ballantine and H. barbatum [1984, as H. pinastroides (S.G Gmelin) Ktitzing], Okamura, of which the latter has also been recorded seems to be based on a misidentification of a Pocock from Kwazulu-Natal (Wynne and Norris 1991). H y and Papenfuss collection (nr. 1024) which is referable poglossum minimum is distinguished from the two to D. subopaca. The relationship of Digeneopsis sub other species by its erect rather than creeping habit opaca to Halopithys and the other Amansieae is be (being attached by a single discoid holdfast), un ing studied by L. Phillips and GT. Kraft (pers. com.). branched blades, a type-one apical organisation and tetrasporangia that are cut off from lateral pericen Gigartinales tral cells. Although some of these features are also present in both H. simulans orH. barbatum, this com Dumontiaceae plete combination of characters is unique to H. mini Gibsmithia hawaiiensis Doty, 1963:458-465, figs. mum (Wynne et al. 1989, Wynne and De Clerck 1-17. Fig. 35 2000). The only other Indian Ocean record comes from the Maldives (Hackett 1977). Recently Stegen- Type locality: Honolulu, Oahu, Hawaiian Archipel ga et al. (2001) described H. imperfectum from the ago. South African south coast. The latter, however, dif Description: Thalli are to 4 cm high, pinkish-red in fers from H. minimum in its prostrate habit and by colour and composed of 4-5 gelatinous lobes, which fertile blades within which the proximal third-order are attached to a conspicuously cartilaginous, annu cell rows fail to reach the thallus margin. late stalk. The stalk is usually simple but may branch 422 O. De Clerck el al. twice in well-developed specimens. The lobes are un Specimens: KZN 614: Sexton Reef (14.viii.1999); divided and are broadly rounded, measuring approx KZN 1611: Sodwana Bay, 2 Mile Reef (12.viii.2000). imately 1 cm in diameter and 3 cm in length.The thal Discussion: Gibsmithia hawaiiensis was originally lus is multiaxial, composed of a filamentous medulla described from the Hawaiian Archipelago and ap and cortex embedded in a gelatinous matrix. The dis pears to be a common component of the deeper sub- tal ends of the cortical filaments produce the gelati tidal in the central, western and southern Pacific nous matrix. Specimens were completely sterile. Ocean (Kraft 1986, Abbott 1999). Its presence in the Ecology: Growing epilithically at a depth of 20 m Indian Ocean was until recently restricted to reports in northern Kwazulu-Natal. from Australia (Huisman 1992) and the Seychelles Twenty marine benthic algae new to South Africa 423

(Kalugina-Gutnik et al 1992). However, a report terised by its rectilinear-shaped cortical cells, the lack from the East African coast (Coppejans et al. 2000) of gland cells, the relatively sparse and large nutritive and the present collection indicates that G. hawaiien cells and the gonimoblasts, which arise apically from sis is the most widely distributed species of the genus. the auxiliary cells. This is the first record of this species for the Indian Ocean. Nemastomataceae Phacelocarpaceae Predaea weldii Kraft et Abbott, 1971:194, figs 1-13. Figs 36-39 Phacelocarpus tristichus J. Agardh, 1885:57-58. Figs 40,41 Type locality: Kaneohe Bay, Oahu Island, Hawaii. Description: The gelatinous thallus is about 3.5 cm Type locality: Mauritius. high, irregularly branched with numerous short,blunt- Description: Thalli are erect, 7-10 cm high, sparse tapering branchlets (Fig. 36).The colour is pale pink ly branched and attached by a small discoid holdfast ish in situ. The thallus is multiaxial, composed of a fila from which a short (up 1 cm) terete stipe arises mentous medulla and cortex embedded in a (Fig. 41). Axes are of uniform width, terete, 1.5 mm in gelatinous matrix. The cortical filaments are dichoto diameter (including teeth). Triangular teeth are mously branched, with rectilinear cells, 4-5 pm wide formed in 3 ranks along the axes (Fig. 40) except near by 8-13 pm long (Fig. 37). Gland cells are absent. Car- the base, where there are 2 ranks. Teeth are slightly pogonial branches are 3-celled (Fig. 39). Auxiliary upwardly directed, 500 pm long and 270 pm wide cells are placed intercalary in cortical filaments and near the base. Reproductive structures were not ob uteriform in shape with a prominent apical bulge. Nu served. tritive cells, grouped per 3-6 in chains 1-3 cells long, Ecology: Growing epilithically in large intertidal are present on the cortical cells immediately below pools (Island Rock) and in the sublittoral to a depth and on the 2 cells distal to the auxiliary cell (Fig. 38). of 43 m. The gonimoblast arises apically from the auxiliary cell Specimens: KZN 243: Sodwana Bay, 2 Mile Reef and not laterally in conjunction with the connecting (8.viii.1999); KZN 2180: Mabibi (13.ii.2001); KZN filament. Male gametophytes were not observed. 1680: Island Rock (14.viii.2000);KZN 516: Tiger Reef Ecology: Collected only once, attached to coral (13 .viii. 1999) rubble at a depth of 18 to 25 m in northern Kwazulu- Discussion: The genus Phacelocarpus contains 9 Natal. species primarily distributed in the Southern Hemi Specimen: KZN 1623: Sodwana Bay, 2 Mile Reef sphere (Indian Ocean, Australia and New Zealand), (12.viii.2000). with the exception of P. japonicus Okamura from Discussion: The genus Predaea is distinguished southern Japan (Womersley 1994). Phacelocarpus from all other red algae by the clusters of nutritive tristichus is predominantly distinguished from the cells (Kraft 1984). From the Indian Ocean only P. other species by the placement of teeth in 3 ranks feldmannii Bprgesen var. indica Balakrishnan et along the axes (although the number of ranks may Chawla (1984) and P. huismanii Kraft (1984) have vary from 2 to 4; Searles 1968), the majority of been hitherto reported. Predaea weldii is charac species having teeth in only 2 ranks. From South

Figs 30-33. Hypoglossum minimum. Fig. 30. H abit of a specimen epiphytic on Carpomitra longicarpa (scale = 1 cm). Fig. 31.Tetrasporic sorus near the apex of a blade (scale = 500 pm). Fig. 32. Male sori in irregular chevrons on both sides of the blade midline (scale = 500 pm). Fig. 33. Mature cystocarp near a blade apex (scale = 500 pm). 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. H abit of a liquid-preserved specimen (scale = 1 cm). Fig. 37. Cortical fascicles of rectilinear cells (scale = 100 pm). Fig. 38. Auxiliary cell with apical gonimoblast initial (arrow) and clusters of nutritive cells (arrow heads) on the contiguous cortical cells (scale = 10 pm). Fig. 39. Three-celled carpogonial branch (scale = 10 pm). Figs 40,41. Phacelocarpus tristichus. Fig. 40. Distal frond showing the tristichous placem ent of determinate branchlets (scale = 2 mm). Fig. 41. H abit of herbari um specimen (scale = 1 cm). Figs 42-44. Halymenia durvillei. Fig. 42. H abit of herbarium specimen (scale = 1 cm). Fig. 43. D etail of the blade apices (scale = 1 mm). Fig. 44. D etail of the thallus surface with minute spinose proliferations (scale = 2 mm). Fig. 45. Carpopeltis phyllophora. Habit of herbarium specimen (scale = 1 cm). 424 O. De Clerck et al.

Africa two species are known:P. oligocanthus Ktitz Carpopeltis phyllophora (J. Hookeret Harvey) ing and P. tortuosus Endlicher et Diesing, both occur Schmitz in Schmitz et Hauptfleisch, 1897:514. Fig. 45 ring along the South African south coast and extend Type locality: Port Arthur, Tasmania. ing into Kwazulu-Natal (Searles 1968, Seagrief 1988). Description: Thalli are erect, to 13 cm high, com Phacelocarpus oligocanthus is a robust species char plánate and composed of dichotomously to irregular acterised by teeth, which become submerged in the ly branched axes, 2 -7 mm in width. The base consists cortex of the main axes.Phacelocarpus tortuosus is of a small discoid holdfast, which extends into a con closer toP. tristichus in size and habit but differs in spicuous midrib in the lower parts of the thallus. The the bilateral placement of the teeth and flattened apices are broadly rounded to obtuse, but proliferate axes to 2.5 mm wide (Searles 1968).Phacelocarpus with mechanical damage. On transverse section the tristichus, originally described from Mauritius, ap thallus is 150-200 pm thick at mid-frond, composed pears to be widely spread throughout the western In of a pseudoparenchymatous cortex and a filamen dian Ocean (Silva et al. 1996). tous medulla. The cortex is 50-70 pm and 4-5 cells thick, composed of cells gradually decreasing in size Halymeniales towards the periphery, which are not arranged in an ticlinal rows. The medulla comprises about 1/3 third Halymeniaceae of the thallus thickness at maximum and is composed Halymenia durvillei Bory de Saint-Vincent, 1828: of compactly arranged filaments. Refractive gan 180-181,pi. 15. Figs 42-44 glionic cells are absent. Tetrasporangia are restricted to the apical parts of the branches, are decussately to Type locality: Port Praslin, New Ireland, Papua New cruciately divided and ovoid (20 pm long, 12 pm Guinea. wide). Gametophytes were not observed. Description: Thalli are erect, to 21 cm high and Ecology: Growing in lower intertidal pools and the composed of irregularly branched, lubricous straps sublittoral fringe. (Fig. 42). The base consists of a small discoid holdfast. Specimens: KZN 103: Durban, Treasure Beach Axes are branched up to 5 orders, taper from the (3.VÜL1999); KZN 1241: Cape Vidal (9.vii.l999); base to the apices and measure 4-20 mm in width. KZN 379:Mabibi (9.viii.l999). Apices are acute and the surface is covered with Discussion: The genus Carpopeltis, comprising acute proliferations (Figs 43,44), which may develop about 11 species, is mainly confined to the warmer into new axes. On transverse section the thallus is ap waters of the Indo-Pacific Ocean. Carpopeltis mail proximately 600 pm thick at mid-frond, composed of lardii (Montagne et Millardet) Chiang and C. beckeri a pseudoparenchymatous cortex and a filamentous Schmitz are known from South Africa. The latter rep medulla. The cortex is 60-80 pm thick, composed of resents an invalidly published manuscript name, the 6 -8 cells forming anticlinal rows. Medullary fila specimens on which it is based not belonging to Car ments are mainly anticlinally arranged, 8-12 pm in popeltis according to Papenfuss (see Silva et al. 1996). diameter, with relatively abundant inner-cortical re Carpopeltis phyllophora bears little resemblance to fractive ganglionic cells. The latter are irregularly C. maillardii; the former being a rather large and sup shaped, with branched arms. Decussately divided ple species, the latter a small, cartilaginous species tetrasporangia (14-20 pm long, 12-15 pm wide) are that branches profusely near the apices [Okamura scattered over the thallus and are cut off from sub 1909, as C. rigida (Harvey) Schmitz; Bprgesen 1943, surface cells. Gametophytes were not observed. as C. rigida', Chiang 1970], Comparison of the South Ecology: Growing in deep intertidal rock pools or African specimens with the type specimen (BM, s.n.), the shallow subtidal (-3 m). collected by Dr Jeannerett at Port Arthur, Tasmania, Specimens: KZN 429: Sodwana Bay, 1/4 Mile Reef reveals them to be morphologically very similar. It (10.viii.1999); KZN 2155: Sodwana Bay, 9 Mile Reef should be noted, however, that the South African (12.ii.2001); KZN 0782: Island Rock (17.viii.1999); specimens are rather variable with respect to thallus NAT 4685 (UN): Black Rock (21.xi.1986). width, with some being as narrow as 1-2 mm, where Discussion: Halymenia durvillei is a widespread as others are 5 -7 mm wide. and easily recognisable species. The branching habit is very distinctive and separates it from most other Plocamiales species of Halymenia, with foliose blades. From South Africa, only H. dilatata Zanardini (Norris and Plocamiaceae Aken 1985) has been previously reported. This Plocamium cf. mertensii (Greville) Harvey, 1849 species has leafy blades with mottled surfaces, which (1847-1849): 122. Figs 46-49 rule out possible confusion with H. durvillei, which has been reported under various synonyms (e.g. H. Type locality: Australia. ceylanica Harvey ex Ktitzing, H. formosa Harvey ex Description: Thalli are 9 (-17) cm tail, red in colour Ktitzing and H. venusta Bprgesen) within the tropical and alternately pinnately branched (Fig. 46). Individ Indian Ocean (De Smedt et al. 2001). ual axes, 2 -3 mm wide, have a plumose aspect, con Twenty marine benthic algae new to South Africa 425 secutive branchlets being of the same length. Later rangial stichidia occur as adventitious axes in the ax als have a tendency to produce narrow pinnules, giv ils of the ramuli or replace compound ramuli ing the fronds a distinctive fimbriate aspect (Fig. 47). (Fig. 49). Individual stichidia are oblong and curved, Ramuli form an alternate-distichous series of one up to 500 pm long and 80-20 pm wide, very occasion simple and one compound branch, cut off at an angle ally branched at their apices. No sexual thalli were of 45° (-60°) (Fig. 48). Simple ramuli are aceróse, found. mostly straight, 2.5-3 (-4) mm long and 0.5-0.7 mm Ecology: Occurring in the lower-intertidal pools wide. Margins of the ramuli are smooth. The central and the shallow subtidal of northern Kwazulu-Natal. axial filament is fairly visible after staining. Apices Specimens: KZN 327: Sodwana Bay, intertidal are slightly to strongly incurved. Clusters of tetraspo (9.viii.1999); KZN 374: Mabibi (9.viii.l999); 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. D etail of an apex (scale = 1 cm). Fig. 49. D etail of tetrasporangial stichidia (scale = 250 pm). Fig. 52. Figs 50-53. Plocamium telfairiae. Fig. 50. H abit of type-one thallus (scale = 1 cm). Fig. 51. H abit of type-two thallus (scale = 1 cm). Fig. 52. H abit of type-three thallus (scale = 1 cm). Fig. 53. Detail of dendroid tetrasporangial stichidia (scale = 250 pm). Fig. 54. Galaxaura rugosa. Habit of herbarium specimen (scale = 1 cm). 426 O. De Clerck et al.

589: Rabbit Rock (13.viii.1999); KZN 761: Kosi Bay(Fig. 52) has a similar outline and structure,but is gen (16.viii.1999). erally smaller (to 4.5 cm) and wider (1.5-2 mm).The Discussion: This species has hitherto only been ramuli of the latter are sub-pinnate, with a compound recorded from southern Australia. The distinctive ramulus subopposite a simple one. The angle of morphology of the tetrasporangial stichidia and the branching lies between that of type-one and type-two fimbriate aspect of the axes of the South African (40-50°). Similarly, the simple ramuli sizes lie be specimens agree closely with the description of this tween type-one and type-two, although they have a species as set out in Womersley (1994). On the other wider base (0.5-1 mm). The central axial filament is hand, no serrations on the abaxial edge of the simple not visible in type-one and type-two (or only near the ramuli were observed. This character, however, does apex), whilst it is in type-three. The tetrasporangial not appear to be fixed in the species. Also, the ten stichidia were found in type-one and type-three, dency of the proximal branchlets in an alternating se whereas only female material was collected for type- ries to become divided to a greater or lesser extent two. The former occur in the axils of ramuli or replac was not observed as typical of the Australian plants ing compound ramuli and having a stellate / tree-shape (G.T. Kraft, pers. comm.). This species is distinctly dif (Fig. 53). ferent from other South African species that have an The specimens examined agree with descriptions alternating-distichous series of two, having relatively and herbarium specimens of this species (including broad axes and relatively long, slender, clustered an isotype specimen in BM).This is the first record of stichidia. this species in South Africa, although Seagrief (1984) and Cormaci et al. (1991) credited this species to South Africa based on supposed records of Simons Plocamium telfairiae (W. Hookeret Harvey) Harvey (1964).The latter, however, reported the species only ex Ktitzing, 1849:885. Figs 50-53 from Mozambique and not from South Africa. This Type locality: Mauritius. species has a wide distribution within the Indian and Description: Thalli are more or less pyramidal in Pacific Oceans. There is also an Atlantic record from outline, rose-red in colour, to 9 cm high. The main Ghana by Lawson and John (1987). axes, 1 (-2) mm wide, are alternate-distichously branched at an angle of 25-70 °. Ramuli form an al Nemalionales ternate-distichous series of one simple and one com pound branch. The latter are strongly incurved near Galaxauraceae the apices. Stellate or dendroid tetrasporangial Galaxaura rugosa (Ellis et Solander) Lamouroux, stichidia are formed in the axils of the ramuli or re 1816:263. Fig. 54 place a compound ramulus. Ecology: Growing in intertidal rock pools and the Type locality: Jamaica. sublittoral fringe to a depth of 15 m in northern Description: Thalli are erect, to 8 cm high, hirsute Kwazulu-Natal. and calcified. The axes are terete, dichotomously Specimens: KZN 1069: Cape Vidal (9.vii.l998); branched, 2 mm in diameter and densely covered by KZN 249: Sodwana Bay, 2 Mile Reef (8.viii.l999); assimilatory filaments. The medulla consists of entan KZN 271: Sodwana Bay, Adlam’s Reef (9.viii.l999); gled filaments (12-16 pm in diameter). The cortex is KZN 329: Sodwana Bay, intertidal (9.viii.l999); KZN filamentous with both short and long assimilatory fil 585: Rabbit Rock (13.viii.1999); KZN 626: KZN 666: aments, which arise from inflated proximal cortex Black Rock (14.viii.1999); Sexton Reef (14.viii.1999);cells (33-42 pm in diameter). The cruciately divided KZN 762: Kosi Bay (16.viii.1999). tetrasporangia are borne apically on short assimilato Discussion: Three forms could be discerned forP. ry filaments. They are ovoid, 25-30 pm long and to telfairiae, distinguished on the branching angle and 20 pm wide. Gametophytes were not observed. differences in size of the axes. The density of branching Ecology: Epilithic in the lower intertidal zone. is similar in type-one (Fig. 50) and type-two (Fig. 51), Specimens: KZN 326: Sodwana Bay, intertidal but the angle of branching is more acute in type-one (9.VÜL1999); KZN 466: Mabibi (ll.viii.1999); KZN (25-) 30-45° (-55), as opposed to (30-) 40-55° 1685: Island Rock (14.viii.2000). (-70). This, combined with a narrower distance be Discussion: Despite taxonomic confusion in the tween consecutive branches, results in a greater de past (see Huisman and Borowitzka 1990), tetra- gree of overlapping ramuli in type-one thalli. Simple sporophytes of G. rugosa are easily recognisable by ramuli are subulate to triangular, mostly straight to their hirsute habit. Several species of Galaxaura and slightly incurved (rarely recurved). Simple ramuli in the morphologically similar genus Tricleocarpa are type-one are generally longer (1.5-2.5 mm) and known from South Africa.Galaxaura diesingiana Za broader (0.3-0.5 mm) than those of type-two (1-1.5 nardini, G. magna Kjellman and G. marginata (Ellis and 0.3 mm respectively). Compound ramuli are et Solander) Lamouroux are all complánate.Galax strongly incurved at the apex, with moderate overlap aura obtusata (Ellis et Solander) Lamouroux andTri ping in type-one and none in type-two. Type-three cleocarpa fragilis (Linnaeus) Huisman et Townsend Twenty marine benthic algae new to South Africa 427 are both terete, but are never hirsute and always lack species have been reported from Inhaca in southern the inflated proximal cells of assimilatory filaments. Mozambique. Their presence in the northern part of Galaxaura rugosa is widespread in the tropical Indi the Kwazulu-Natal coastline, therefore, only indi an Ocean. cates a minor range extension. In contrast, the distri bution range of Digeneopsis subopaca remains re stricted to northern Kwazulu-Natal and the Inhaca Peninsula in Mozambique. Together with Dasycladus Discussion ramosus Chamberlain (1958), these species are prob Distribution patterns of tropical seaweeds are incom ably endemic to the northern part of the overlap re plete for many taxa. Several regions have received gion between warm-temperate South African waters very little attention and remain seriously un(der)ex- and the tropical Indian Ocean. plored. There are not only limited data on small Balliella crouanioides, Gibsmithia hawaiiensis, islands and archipelagos but also for several conti Predaea weldii and Hypoglossum minimum are char nental coastal regions such as Oman, Iran and Mada acterised by disjunct tropical Indo-Pacific distribu gascar. This makes biogeographical assessments diffi tion patterns. All three were originally described cult, but a number of clear trends are indicated from the central or western Pacific Ocean and have (Table I). been reported from a few scattered localities within The majority of the Phaeophyta and Rhodophyta the Indian Ocean. Their small size and subtidal habi reported in this paper consist of widespread tropical tat probably contribute to the scarcity of reports.Bal species .Asteronema breviarticulatum, Dictyota cerviliella crouanioides and Gibsmithia hawaiiensis have cornis, D. ciliolata and Galaxaura rugosa are mainly recently been reported for the East African coast pantropical species, occurring in the Atlantic (Coppejans et al. 2000). Their presence along the (Caribbean Sea) and Indian Oceans as well as the Pa South African coast may indicate a wide distribution cific Ocean. Some are present along the Western within the Indo-Pacific region. Australian coast. Next to the pantropical taxa, sever Carpopeltis phyllophora and Plocamium mertensii al species have a defined Indo-Pacific or Indian are both Australian species. Norris and Aken (1985) Ocean distribution: Ceramium cingulatum, Euptilota and Hommersand (1986) regard this South African fergusonii, Halymenia durvillei and Phacelocarpus flora as having affinities with the western- and south tristichus. The presence of pantropical and Indo-Pa ern Australian algal floras, although there has been cific species along the northern part of the Kwazulu- some debate on what may have caused these floristic Natal coast is not surprising, as several of these similarities. According to Hommersand (1986), the

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

Mozambique Western Central Indonesia- Australia Pacific A tlantic Indian Indian Malaysia Ocean Ocean Ocean Ocean

Asteronema breviarticulatum _ + + + + + + Balliella crouanioides + + - + - + - Carpopeltis phyllophora - - - + + - - Ceramium cingulatum - + - + + - - Dictyota cenncomis + + + + + + + Dictyota ciliolata + + + + + + + Dictyota hamifera - + - + - + + Dictyota rigida - + - - - - - Digeneopsis subopaca + ------Euptilota articulata - - + - + + - Euptilota fergusonii + + + - - - - Galaxaura rugosa + + + + + + + Gibsmithia hawaiiensis - + - + + + - Halymenia durvillei - + + + + + - Hypoglossum m in ini um - - + - - + - Padina gymnospora 7 + ? ? + ? + Phacelocarpus tristichus + + + - - - - Plocamium mertensii - - - - + - - Plocamium telfairiae + - + - - + - Predaea weldii - - + + + +

General references: Womersley (1987; 1994; 1998); Millar and Kraft (1993); Silvaet al. (1996); Phillips (1997); Wynne (1998);Yoshida (1998); Abbott (1999); Coppejans and Millar (2000); Coppejanset al. (2000); Huisman (2000). 428 O. De Clerck et al. genus Carpopeltis and to a certain extentPlocamium macrocarpa (Areschoug) Schmidt, Ecklonia radiata represent examples of species clusters that originated (C. Agardh) J. Agardh and Pseudocodium de-vriesii along the west and south coasts of Australia and mi Weber-van Bosse (Barratt et al. 1984, Barratt et al. grated to South Africa during mid-to-late Miocene 1986, Wynne 1999). periods (16-6Mya) marked by a general decrease in temperature in the tropics. Euptilota articulata may also represent an example of the South African-Aus Acknowledgements tralian connection, although the species has also been reported from India and the North-West Pacific. We would like to offer our sincere gratitude to Jean The distribution of Padina gymnospora is difficult Harris, Nonhlanhla Nxumalo, Bridget Armstrong, to interpret due to taxonomic confusion in the past. Cloverly Lawrence and John Dives, of the Kwazulu- Studies on the genus Padina along the East African Natal Nature Conservation Services, for their full coast (Somalia to South Africa) have revealed that P. support in organising everything, everytime, every gymnospora is absent along the tropical coasts of where. We also extend our thanks to Peter Timm Kenya, Tanzania and the various tropical archipela from Triton Divers for his smoothly run services. gos (Muylle 2000). The species is, however, present in Funding for this project was provided by the Interna the northern part of the western Indian Ocean (So tional Scientific and Technological Cooperation malia, Yemen and Oman) as well as along the coast (BIL98/84) between the Ghent University and the of Kwazulu-Natal. The distribution pattern is inter University of Cape Town and FWO Research Project esting as both geographically isolated populations (3G002496). Further support was provided in South are in habitats characterised by slightly cooler water Africa by Marine and Coastal Management, the Na temperatures than occur in the tropical Indian tional Research Foundation and the Department of Ocean. Similar examples of disjunct distribution pat Environment and Tourism. terns between the northern Arabian Sea and the South African east coast are offered by: Dictyopteris A ccepted 19 May 2002.

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