Botanica Marina Vol. 45, 2002, pp. 413–431 © 2002 by Walter de Gruyter · Berlin · New York
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, Belgium
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
b Botany
* 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 IndoPacific region (Balliella crouanioides, Gibsmithia hawaiiensis, Predaea weldii, Hypoglossum minimum), possibly 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 represent a local endemic species.
Introduction
The South African coastline displays a unique floral
diversity, with biota ranging from kelp beds on the
west coast to coral reefs near the Mozambican border. The correlation between surface seawater temperatures and shore community compositions has
been investigated by several researchers, who have
firmly established that the main defining influences
are the warm Agulhas current flowing down the east
coast from Mozambique to the region where it meets
the cold Benguela current (e.g. Stephenson 1948,
Bolton 1986, Bolton and Anderson 1990). Stephenson (1948) divided the South African coastline into
three marine provinces: the cold-temperate west
coast, warm-temperate south coast, and sub-tropical
east coast. More recently, seaweed biogeographers,
although also recognising three floral provinces, have
described them differently. The seaweed flora of the
west coast is regarded as warm-temperate by Bolton
(1986) and Lüning (1990) or cool-temperate by
Emanuel et al. (1992) and Bolton and Anderson
(1997), rather than cold-temperate. The presence of a
distinct sub-tropical flora on the east coast (Kwazulu-Natal) is not accepted by either Lüning (1990) or
Bolton and Anderson (1997). These authors consider
the seaweeds of the east coast of South Africa to be
the westernmost extension of the large Indo-West
Pacific tropical region, and cast doubt on the existence of a distinct sub-tropical flora in Kwazulu-Natal.
The transition zone between the west and south
coast marine provinces has been well studied and
characterised by a clearly defined overlap zone
(Bolton and Stegenga 1990, Jackelman et al. 1991), in
contrast to the eastern border of the Agulhas (south
coast) marine province which is less clear. Seawater
temperature, the main defining ecological parameter
in the distribution of seaweeds, gradually rises from
around East London to northern Kwazulu-Natal
(Bolton 1986). At present, it is believed that a distinct
sub-tropical marine province in Kwazulu-Natal is unlikely, as the flora seems to comprise an eastwardly
declining number of Agulhas-province species, these
being replaced largely by Indo-West pacific species
with the increase in water temperature (Bolton and
Anderson 1997). It should be noted, however, that
our knowledge of the marine flora of Kwazulu-Natal
is limited. Apart from a limited number of taxa, studied in detail by students of G.F. Papenfuss in Berkeley (California) and a field guide on the seaweeds of
Maputaland by Seagrief (1980), no detailed studies
were available relating to Natal algae prior to 1985.
In the eighties and early nineties R.E. Norris and coauthors published a series of papers on the algae of
Kwazulu-Natal that gave some evidence that there is
indeed a distinct flora in the region. In total they (see
Norris 1992 and Silva et al. 1996 for a complete reference list) reported 63 species as new records for
South Africa and described 23 new species, many of
which are currently only known from KwazuluNatal. It has also been noted that elements of the
Kwazulu-Natal marine algal flora have a link with
southwestern Australia (Norris and Aken 1985,
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O. De Clerck et al.
Hommersand 1986). Kwazulu-Natal may therefore
be more than a transition zone between the Agulhas
and Indo-West Pacific floras, in that it actually constitutes a distinct flora with warm-temperate southwestern Australian affinities and a large number of
endemics.
In a joint research project conducted by the
Botany Departments of the Ghent University (Belgium) and the University of Cape Town (South
Africa), the seaweeds of Kwazulu-Natal are being
studied from a biogeographical perspective. Several
sites along the coast from Port Edward in the south
to Kosi Bay near the Mozambican border were sampled mainly between July 1998 and February 2001.
The collections are now being identified and will
eventually serve to provide a better understanding of
the distribution and floristic affinities of the algae
along the Kwazulu-Natal coast. This paper reports 20
species which are newly recorded for South Africa.
Records on Chlorophyta have recently been published separately by Leliaert et al. (2001).
Material and Methods
Approximately 2400 specimens were collected over a
period of two years (July 1998; August 1999; December 1999; July–August 2000; February 2001), as a result of a Bilateral Scientific and Technological Cooperation Project between the Flemish community
(Belgium) and South Africa. Specimens were collected at various sites distributed along the entire Kwazulu-Natal coast (see Fig. 1) and were immediately
processed as herbarium specimens, as well as being
partly preserved in 4% formaldehyde/seawater. The
specimens, 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 material was stained in a mixture of 1 g aniline blue powder, 50 mL Karo®, 45 mL distilled water, 5 mL acetic
acid. Collections other than our own, those of R.E.
Norris and co-workers (bearing serial numbers prefixed by ‘NAT’) deposited in UN and various other
collectors in BOL were also examined. Herbarium
abbreviations follow Holmgren et al. (1990). Digital
images were taken, using an Olympus DP50 digital
camera (Melville, USA) mounted on a Leitz Diaplan
or Wild M10 microscope (Wetzlar, Germany).
parts of the thallus and recurved branchlets; specimens from exposed habitats generally form low,
dense mats and lack slender straps. The South
African specimens represent the typical growth form
from exposed habitats. A major difference to previous observations is the occurrence of iridescence.
Despite the examination of over 200 specimens by
De Clerck (1999), mainly from the East African coast
(Tanzania), iridescence was never observed. The taxonomic value and intraspecific variability of iridescence, however, remains poorly understood (Gaillard
1972).
Results
Dictyota ciliolata Sonder ex Kützing, 1859: 12, pl. 27,
fig. 1.
Figs 5, 6
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 composed of coarse, 2–3 mm wide, subdichotomous
straps (Fig. 4). The margins are smooth, but the surface is beset with acute proliferations (Fig. 3). The
apices are obtuse. Sporangia are scattered on both
surfaces, but absent in the apical dichotomies. Sporangia, 120–135 µm in diameter, are placed mostly
solitary and are borne on a single stalk cell, surrounded by a conspicuous involucrum. Gametangia were
not observed. In situ the colour is brown with a bluegreyish 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 the
most common Dictyota species in the Indian Ocean,
characterised by surface proliferations and sporangia
that are surrounded by an involucrum. It appears to
be closely related to D. crispata Lamouroux and D.
magneana De Clerck et Coppejans (De Clerck and
Coppejans 1997, Coppejans et al. 2001), but differs
mainly in the placement of sporangia, these being absent or present in the apical dichotomies respectively.
Other characters which distinguish D. crispata from
D. cervicornis include the morphology of the apices
and the abundance and placement of surface proliferations. Detailed comparisons between the three
species are made by Coppejans et al. (2001). Dictyota
cervicornis exhibits a broad range of morphological
growth forms, which can often be related to its habitat. Specimens growing in the shallow subtidal are often characterised by long slender straps in the distal
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). The
margins are dentate, rarely smooth, while the surface
is always smooth. The apices are rounded. The
medulla and cortex are uniformly one-layered. Sporangia are single, scattered on both surfaces, but absent in the apical dichotomies. Sporangia, 95–110 µm
in diameter, are borne on a single stalk cell and are
not surrounded by a conspicuous involucrum. Gametangia were not observed.
Ecology: Specimens were collected in intertidal
pools 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 its
stupose holdfast, dentate margins and the absence of
stolonoidal fibres, although the margins of some
specimens can be nearly smooth (Hörnig et al.
1992a,b). Several other dentate species of Dictyota
occur in the Indo-Pacific and the Caribbean regions,
as detailed by De Clerck and Coppejans (1999).
Along the South African coast D. ciliolata is most
similar to D. liturata J. Agardh and D. suhrii G. Murray. The latter species, however, has a multilayered
medulla near the margins, whereas the medulla of D.
ciliolata is uniformly one-layered. Dictyota ciliolata
differs from D. liturata in the placement of the sporangia, which are arranged singly or distributed in
longitudinal 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, attached to hosts at various points by patches of rhizoids. The straps, 1–2 mm wide, are dichotomously
branched. The margins are smooth except for the falcate branchlets (Fig. 7), which may resemble teeth.
The surface is smooth. The apices are rounded to tridentate. The medulla and cortex are uniformly one-
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O. De Clerck et al.
Twenty marine benthic algae new to South Africa
layered. Sporangia or gametangia were not observed.
Ecology: Specimens were collected at a depth of 18
to 25 m epiphytic on Plocamium sp.
Specimens: KZN 1617: Sodwana Bay, 2 Mile Reef
(12.viii.2000).
Discussion: Dictyota hamifera was for a long time
only known from the Pacific Ocean. Hörnig et al.
(1992a) reported the species for the first time from the
Atlantic Ocean,while De Clerck (1999) and Coppejans
et al. (2000) recorded the species from the East African
coast (Kenya and Tanzania).Dictyota hamifera is easily
recognised 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 dichotomously
branched straps are narrow (0.8–1.2 mm) and of uniform width throughout the thallus (Figs 8, 10). The
margins and surface are smooth. The medulla and
cortex are uniformly one-layered. Sporangia form a
well-defined line in the median part of the straps, but
are not present along the margins (Fig. 9). Sporangia,
80–95 µm in diameter, are not surrounded by an involucrum and are borne on a single stalk cell. Gametangia were not observed.
Ecology: Specimens were collected in pools of the
sublittoral fringe.
Specimens: KZN 311: Sodwana Bay, Adlam’s Reef
(9.viii.1999); KZN 385: Mabibi (9.viii.1999).
Discussion: Dictyota rigida was recently described
from Tanzania (De Clerck and Coppejans 1999) and
subsequently reported from Kenya (Coppejans et al.
2000). Characters defining D. rigida include the erect
growth form, the stiff texture and the single stupose
holdfast (De Clerck and Coppejans 1999). Dictyota
rigida is most similar to the South African D. suhrii
417
G. Murray and the Australian D. fastigiata Sonder.
The latter two species, however, are characterised by
a multilayered medulla, at least in the basal parts of
the thallus.
Padina gymnospora (Kützing) Sonder,
1871: 47.
Figs 11, 12
Type locality: St. Thomas, Virgin Islands.
Description:Thalli, attached by a small stupose rhizoidal base, are to 6.5 cm high, flabellate and composed of several lobes with inrolled margins (Fig. 11).
The superior surface (the side toward which the margin is inrolled) is slightly to moderately calcified.
Thalli are olive-brown in colour. In transverse section the thallus is composed of 4–6 layers of cells in
the mid-regions and up to 8–9 layers near the base.
Hair rows are present on both sides of the thallus, but
more conspicuous on the inferior side of the thallus.
Tetrasporangia are arranged in concentric lines,
above each hair row, mainly (but not exclusively) developed on the superior surface. An evanescent indusium is present but is usually only clearly visible in
very young sori (Fig. 12). Tetrasporangia are ovoid,
up to 100 µm long and 70 µm wide. Gametophytes
were not observed.
Ecology: Specimens are collected in shallow pools
in 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); KZN
708: Bhanga Nek, N23 (15.viii.1999); KZN 745: Kosi
Bay (16.viii.1999).
Discussion: Padina species are distinguished primarily on the number of cell layers, the distribution
of hair bands, position of the sporangial sori (on the
superior versus inferior surface) and the type of alternations of hair and sporangial bands. Padina gymnospora is characterised by the thick thallus (up to 8
cell 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 apical 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. Detail 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 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 a
hooked branch (scale = 50 µm). Fig. 15. Detail of an ovoid plurilocular sporangium (scale = 50 µm).
418
O. De Clerck et al.
the superior surface of the thallus and the evanescent
indusium (Allender and Kraft 1983, Womersley
1987). The species is most similar to P. crassa Yamada, the latter supposedly having sporangia only on
the lower side of the thallus. Only P. boryana Thivy
and P. plumbea (Areschoug) Levring have previously
been reported from South Africa (Levring 1940,
Gaillard 1975). Both species are characterised by a 2layered thallus and hence are easily distinguished
from P. gymnospora. Due to confusion with P.
boergesenii Allender et Kraft in the past, the distribution of P. gymnospora is difficult to assess in the Indian Ocean (Silva et al. 1996). The latter, however, has
been reported with certainty from Australia and
Oman (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 appearance of a frayed rope at its tips (Fig. 13). Filaments
are uniseriate and irregularly branched with numerous short hooked branches arising at an angle of
80–90 ° (Fig. 14). Growth takes place by intercalary
cell divisions. Cells are 25–33 µm wide, 60–80 µm
long, L/B: 1–1.3. Plurilocular sporangia, placed on a
1–2-celled stalk, are spherical to ovoid and measure
40 x 42 (–63) µm on average (Fig. 15).
Ecology: growing in the intertidal, often attached
to barnacles in wave-exposed localities.
Specimens: KZN 451: Mabibi (11.viii.1999); KZN
654: Black Rock (14.viii.1999); KZN 728: N23
(15.viii.1999).
Discussion: Asteronema breviarticulatum is a widespread warm-temperate to tropical species characterised by the shape of the plurilocular sporangia and
the presence of short hooked branches. Ouriques and
Bouzon (2000) transferred this species, which was
commonly known as Hincksia breviarticulata (J.
Agardh) Silva, to Asteronema on the basis of the stellate chloroplasts. Recent molecular studies by Draisma et al. (2001) and Rousseau et al. (2001) reveal,
however, that Asteronema is polyphyletic and not related to the Ectocarpales sensu lato. Unfortunately
Asteronema breviarticulatum itself was not included
in the analysis. Pending further studies, A. breviarticulatum is listed as incertae sedis rather than under the
Ectocarpales.
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). Filaments are opposite-distichously branched with indeterminate branches being formed every (1–) 2 (–5)
segments. The apical parts of the axes show a distinctive sinusoidal curving (Fig. 17). Corticating rhizoids
arise from the lower cells of the laterals and form a
loose rag-like cortex (Fig. 19). The periaxial cells are
100–115 µm in diameter and bear spherical vesicular
cells, 8–15 µm in diameter on their abaxial side
(Fig. 18). Decussately divided tetrasporangia develop
on the adaxial side of the periaxial cells of the laterals. Tetrasporangia are ovoid, 30–55 µm long and
30–38 µm wide. Gametophytes were not observed.
Ecology: Balliella crouanioides is a common component of the sublittoral zone (–20 to –35 m), where
it forms distinctive orangy-iridescent tufts on a variety 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: Sodwana Bay, Deep Sponge (10.ii.2001); KZN 250 and
KZN 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 by
the presence of spherical vesicular cells (‘gland
cells’), which occur adaxially or abaxially on the periaxial cells. Species are separated on the basis of their
habit, branching pattern, position and size of vesicular cells and tetrasporangia. Wollaston (1984) reported B. subcorticata (Itono) Itono et Tanaka from
Mozambique, but Huisman (1988) expressed the
opinion that the specimen depicted was more similar
to B. crouanioides, based on the overall branching
pattern, arrangement of the tetrasporangia and size
of the vesicular cells. Comparison of our material
with the specimen studied by Wollaston (UC
1470004) reveals that they are identical and supports
Huisman’s proposition. Athanasiadis (1996) provisionally assigned a collection of specimens from
Kenya to B. crouanioides. Re-examination of the
type collection proved to be necessary as some controversies exist on the branching pattern and placement of the gland cells in this species. The South
African specimens match previous accounts of B.
crouanioides very well (Itono and Tanaka 1973, Itono
1977, Huisman and Kraft 1984), although variation
was observed in both the size (8–15 µm in diameter)
and placement of the gland cells (abaxial but occasionally adaxial). As these characters are regarded to
be of prime importance in delineating Balliella
species, further research is needed on intraspecific
variation.
Ceramium cingulatum Weber-van Bosse, 1923:
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 branching (scale = 100 µm). Fig. 18. Cruciately divided adaxial tetrasporangia and abaxial gland cells (arrowheads) on the periaxial 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. Detail
of 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 of
an 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).
420
O. De Clerck et al.
Type locality: Sape Strait, south of Sangeang Island,
Indonesia.
Description: Thalli to 2 mm high consisting of unbranched erect axes, that arise mainly from a single
point or from a creeping filament. Axes are somewhat clavate, measuring 90–100 µm wide near the
base, 140–170 µm in the middle part of the thallus,
ending abruptly in an attenuate apex (Fig. 20). Axial
cells are lenticular, usually wider than long except
near the base. A total of 8 periaxial cells are present
per segment, each producing 2 anterior and 2 posterior cortical initials of cortical filaments. Anterior initials cut off 1–2 acropetal filaments, which are up to 2
cells long (Fig. 22). Posterior initials either remain
undivided or produce a single basipetal filament
(Fig. 22). Cortication remains incomplete, but adjacent cortical bands may be nearly confluent.
Tetrasporangia form in distinctive whorls, are slightly
protruding and measure 28–35 µm in diameter
(Fig. 21). Gametangia were not observed.
Ecology: Epiphytic on Plocamium telfairiae at –18
to 25 m.
Specimens: KZN 1619.2: Sodwana Bay, 2 Mile
Reef (12.viii.2000).
Discussion: The South African specimens agree
very well with the original description by Weber-van
Bosse (1923) and a subsequent account of the species
by Cormaci and Furnari (1991), who compared C.
cingulatum with the Mediterranean C. giacconei Cormaci et Furnari. The only difference we observed was
the mode of attachment. Typically, C. cingulatum has
erect axes in the terminal parts or produces erect
branches. South African plants were usually strictly
erect, but the presence of rhizoids in the lower part of
the thallus indicates the possibility of a creeping
growth form. Strictly erect versus ascending growth
forms are not considered to be major taxonomic differences in Ceramium (Dixon 1960, Womersley
1978). Ceramium cingulatum is unique among South
African representatives of the genus in its unbranched erect axes.The species has been reported in
the Indian Ocean from Somalia, Tanzania and the
Seychelles (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 pinnately branched up to 5 orders (Fig. 23).A percurrent
main axis is lacking. The holdfast is densely rhizoidal.
Axes are corticated from close to the apices, with corticating filaments arising from the periaxial cells 5 to
10 cells proximal to the apex. Near the base the axes
attain a width of 1.5 mm. Each axial cell bears a single
lateral in a distichous-alternate pattern (Fig. 24). Determinate branchlets are ecorticate, straight, 12–17
cells long and pinnately branched (Fig. 25). Spinelike cells are absent. Indeterminate branchlets are
formed irregularly and repeat the branching pattern
of the main axes. Tetrahedrally divided tetrasporangia (40–52 µm in diameter) are borne laterally or terminally on the ultimate branchlets (Fig. 26). Gametophytic plants were not observed.
Ecology: Euptilota articulata has only been collected once in situ, at a depth of 35 m in southern Natal.
The specimen in the Norris Herbarium (UN) was
collected from the drift.
Specimens: KZN 1962: Protea Banks, Northern
Pinnacle (5.ii.2001); NAT 1310: Palm Beach
(14.v.1983).
Discussion: The genus Euptilota contains at present four species, all of which are characterised by an
alternately-distichously branched thallus, in which
axial cells each bear a single lateral (Millar 1990,
Womersley 1998). Euptilota articulata differs from
E. pappeana Kützing, the only species reported
from South Africa (Stegenga et al. 1997), in the pinnate branching of the determinate laterals as opposed to laterals, which are either unbranched or
once to twice dichotomously branched in the latter
species. Euptilota articulata has previously been reported for Australia (Millar 1990, Womersley 1998),
Japan (Itono 1977) and India (Umamaheswara Rao
1974).
Euptilota fergusonii Cotton, 1907: 262–264.
Figs 27–29
Type locality: ‘Pantura’, Sri Lanka.
Description: Thalli are erect, to 15 cm high, pinnately branched up to 5 orders and lack a percurrent
main axis (Fig. 27). The holdfast is densely rhizoidal.
Axes are corticated from close to the apices with corticating filaments arising from the periaxial cells, approximately 12–14 cells proximal to the apex. Proximal axial cells are 450–500 µm wide and 240–270 µm
long. Each axial cell bears a single lateral in a
distichous-alternate pattern (Fig. 28). Determinate
branchlets are ecorticate, curved upwards and up to
12 cells long. The basal 3 cells of a determinate lateral
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
the ultimate and penultimate cells, which bear 1–2
(–4) spine-like cells (Fig. 29). Indeterminate branchlets are formed irregularly and repeat the branching
pattern of the main axes. Tetrahedrally divided
tetrasporangia (60 µm long, 40–50 µm wide) are
borne laterally on cells of the ultimate branchlets.
Gametophytes were not observed.
Ecology: Collected at two occasions only, at a
depth of 15–25 m in northern Kwazulu-Natal (Sodwana 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
Twenty marine benthic algae new to South Africa
from other Euptilota species in having spine-like cells
at the tips of the determinate laterals, the latter additionally being secundly, as opposed to alternately or
subdichotomously, branched. Euptilota fergusonii
was originally described from Sri Lanka (Cotton
1907) and has previously been reported from Tanzania and Mozambique (Jaasund 1976, Wollaston
1984).
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, lanceolate 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. all
second order row cells produce third order rows).
The midrib remains uncorticated throughout.
Tetrasporangial sori form discrete, paired ovate sori
close to the midrib (Fig. 31), 900–1700 µm long and
400–600 µm wide. Tetrasporangial formation involves the lateral pericentral cells but not the transverse ones. Tetrasporangia, 62–80 µm in diameter,
are cut off from second- and third order cell rows as
well as from cortical cells, resulting in sporangia that
lie in more than one plane of the blade. Spermatangial sori form discrete, somewhat oblique patches
scattered over the wings (Fig. 32). Female plants produce a single cystocarp on the midrib in the distal region of the blades (Fig. 33).
Ecology: Collected on a single occasion, epiphytic
on Carpomitra longicarpa Simons at –43 m.
Specimen: KZN 506: Tiger Reef, Bhanga Nek
(13.viii.1999).
Discussion: Hypoglossum minimum is a small
species, which can easily be confused with H. simulans Wynne, Price et Ballantine and H. barbatum
Okamura, of which the latter has also been recorded
from Kwazulu-Natal (Wynne and Norris 1991). Hypoglossum minimum is distinguished from the two
other species by its erect rather than creeping habit
(being attached by a single discoid holdfast), unbranched blades, a type-one apical organisation and
tetrasporangia that are cut off from lateral pericentral cells. Although some of these features are also
present in both H. simulans or H. barbatum, this complete combination of characters is unique to H. minimum (Wynne et al. 1989, Wynne and De Clerck
2000). The only other Indian Ocean record comes
from the Maldives (Hackett 1977). Recently Stegenga et al. (2001) described H. imperfectum from the
South African south coast. The latter, however, differs from H. minimum in its prostrate habit and by
fertile blades within which the proximal third-order
cell rows fail to reach the thallus margin.
421
Rhodomelaceae
Digeneopsis subopaca Simons, 1970: 10, 11.
Fig. 34
Type locality: Santa Maria, Inhaca, Mozambique
Description: Thalli are erect, very stiff and blackish
in colour, to 11 cm tall (Fig. 33). Indeterminate axes
are 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 trifurcately branched in up to 2 or 3 orders. The thallus
is polysiphonous, consisting of an axial cell surrounded by 5 pericentral cells, which are obscured by a
well-developed cortex from close to the apices.
Tetrasporangia are formed in clustered stichidia near
the bases of determinate laterals. The stichidia are
100–190 µm thick, with each segment bearing 2
tetrasporangia. Tetrasporangia are up to 115 µm in
diameter and tetrahedrally divided. Gametophytes
were not observed.
Ecology: Collected in intertidal rock pools and the
shallow 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 described from southern Mozambique (Simons 1970),
remains a poorly known representative of the
Rhodomelaceae. In the absence of gametophytic material the taxonomic affinities of this monotypic
genus are unclear. The morphology of the tetrasporangial stichidia and the compressed, bilateral nature
of the plants, however, seem to indicate a close relationship to the Amansieae. The South African record
of Halopithys incurva (Hudson) Batters in Seagrief
[1984, as H. pinastroides (S.G. Gmelin) Kützing],
seems to be based on a misidentification of a Pocock
and Papenfuss collection (nr. 1024) which is referable
to D. subopaca. The relationship of Digeneopsis subopaca to Halopithys and the other Amansieae is being 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 Archipelago.
Description: Thalli are to 4 cm high, pinkish-red in
colour and composed of 4–5 gelatinous lobes, which
are attached to a conspicuously cartilaginous, annulate stalk. The stalk is usually simple but may branch
422
O. De Clerck et al.
twice in well-developed specimens. The lobes are undivided and are broadly rounded, measuring approximately 1 cm in diameter and 3 cm in length.The thallus is multiaxial, composed of a filamentous medulla
and cortex embedded in a gelatinous matrix. The distal ends of the cortical filaments produce the gelatinous matrix. Specimens were completely sterile.
Ecology: Growing epilithically at a depth of 20 m
in 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 originally
described from the Hawaiian Archipelago and appears to be a common component of the deeper subtidal in the central, western and southern Pacific
Ocean (Kraft 1986, Abbott 1999). Its presence in the
Indian Ocean was until recently restricted to reports
from Australia (Huisman 1992) and the Seychelles
Twenty marine benthic algae new to South Africa
(Kalugina-Gutnik et al. 1992). However, a report
from the East African coast (Coppejans et al. 2000)
and the present collection indicates that G. hawaiiensis is the most widely distributed species of the genus.
423
terised by its rectilinear-shaped cortical cells, the lack
of gland cells, the relatively sparse and large nutritive
cells and the gonimoblasts, which arise apically from
the auxiliary cells. This is the first record of this
species for the Indian Ocean.
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,blunttapering branchlets (Fig. 36). The colour is pale pinkish in situ.The thallus is multiaxial, composed of a filamentous medulla and cortex embedded in a
gelatinous matrix. The cortical filaments are dichotomously branched, with rectilinear cells, 4–5 µm wide
by 8–13 µm long (Fig. 37). Gland cells are absent. Carpogonial branches are 3-celled (Fig. 39). Auxiliary
cells are placed intercalary in cortical filaments and
uteriform in shape with a prominent apical bulge. Nutritive cells, grouped per 3–6 in chains 1–3 cells long,
are present on the cortical cells immediately below
and on the 2 cells distal to the auxiliary cell (Fig. 38).
The gonimoblast arises apically from the auxiliary cell
and not laterally in conjunction with the connecting
filament. Male gametophytes were not observed.
Ecology: Collected only once, attached to coral
rubble at a depth of 18 to 25 m in northern KwazuluNatal.
Specimen: KZN 1623: Sodwana Bay, 2 Mile Reef
(12.viii.2000).
Discussion: The genus Predaea is distinguished
from all other red algae by the clusters of nutritive
cells (Kraft 1984). From the Indian Ocean only P.
feldmannii Børgesen var. indica Balakrishnan et
Chawla (1984) and P. huismanii Kraft (1984) have
been hitherto reported. Predaea weldii is charac-
Phacelocarpaceae
Phacelocarpus tristichus J. Agardh, 1885: 57–58.
Figs 40, 41
Type locality: Mauritius.
Description: Thalli are erect, 7–10 cm high, sparsely branched and attached by a small discoid holdfast
from which a short (up 1 cm) terete stipe arises
(Fig. 41). Axes are of uniform width, terete, 1.5 mm in
diameter (including teeth). Triangular teeth are
formed in 3 ranks along the axes (Fig. 40) except near
the base, where there are 2 ranks. Teeth are slightly
upwardly directed, 500 µm long and 270 µm wide
near the base. Reproductive structures were not observed.
Ecology: Growing epilithically in large intertidal
pools (Island Rock) and in the sublittoral to a depth
of 43 m.
Specimens: KZN 243: Sodwana Bay, 2 Mile Reef
(8.viii.1999); KZN 2180: Mabibi (13.ii.2001); KZN
1680: Island Rock (14.viii.2000); KZN 516:Tiger Reef
(13.viii.1999)
Discussion: The genus Phacelocarpus contains 9
species primarily distributed in the Southern Hemisphere (Indian Ocean, Australia and New Zealand),
with the exception of P. japonicus Okamura from
southern Japan (Womersley 1994). Phacelocarpus
tristichus is predominantly distinguished from the
other species by the placement of teeth in 3 ranks
along the axes (although the number of ranks may
vary from 2 to 4; Searles 1968), the majority of
species 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 a
blade (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 contiguous
cortical 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 herbarium 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 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 Kützing and P. tortuosus Endlicher et Diesing, both occurring along the South African south coast and extending into Kwazulu-Natal (Searles 1968, Seagrief 1988).
Phacelocarpus oligocanthus is a robust species characterised by teeth, which become submerged in the
cortex of the main axes. Phacelocarpus tortuosus is
closer to P. tristichus in size and habit but differs in
the bilateral placement of the teeth and flattened
axes to 2.5 mm wide (Searles 1968). Phacelocarpus
tristichus, originally described from Mauritius, appears to be widely spread throughout the western Indian 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 New
Guinea.
Description: Thalli are erect, to 21 cm high and
composed of irregularly branched, lubricous straps
(Fig. 42).The base consists of a small discoid holdfast.
Axes are branched up to 5 orders, taper from the
base to the apices and measure 4–20 mm in width.
Apices are acute and the surface is covered with
acute proliferations (Figs 43, 44), which may develop
into new axes. On transverse section the thallus is approximately 600 µm thick at mid-frond, composed of
a pseudoparenchymatous cortex and a filamentous
medulla. The cortex is 60–80 µm thick, composed of
6–8 cells forming anticlinal rows. Medullary filaments are mainly anticlinally arranged, 8–12 µm in
diameter, with relatively abundant inner-cortical refractive ganglionic cells. The latter are irregularly
shaped, with branched arms. Decussately divided
tetrasporangia (14–20 µm long, 12–15 µm wide) are
scattered over the thallus and are cut off from subsurface cells. Gametophytes were not observed.
Ecology: Growing in deep intertidal rock pools or
the 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 widespread
and easily recognisable species. The branching habit
is very distinctive and separates it from most other
species of Halymenia, with foliose blades. From
South Africa, only H. dilatata Zanardini (Norris and
Aken 1985) has been previously reported. This
species has leafy blades with mottled surfaces, which
rule out possible confusion with H. durvillei, which
has been reported under various synonyms (e.g. H.
ceylanica Harvey ex Kützing, H. formosa Harvey ex
Kützing and H. venusta Børgesen) within the tropical
Indian 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, complanate and composed of dichotomously to irregularly branched axes, 2–7 mm in width. The base consists
of a small discoid holdfast, which extends into a conspicuous midrib in the lower parts of the thallus. The
apices are broadly rounded to obtuse, but proliferate
with mechanical damage. On transverse section the
thallus is 150–200 µm thick at mid-frond, composed
of a pseudoparenchymatous cortex and a filamentous medulla. The cortex is 50–70 µm and 4–5 cells
thick, composed of cells gradually decreasing in size
towards the periphery, which are not arranged in anticlinal rows. The medulla comprises about 1/3 third
of the thallus thickness at maximum and is composed
of compactly arranged filaments. Refractive ganglionic cells are absent. Tetrasporangia are restricted
to the apical parts of the branches, are decussately to
cruciately divided and ovoid (20 µm long, 12 µm
wide). Gametophytes were not observed.
Ecology: Growing in lower intertidal pools and the
sublittoral 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, comprising
about 11 species, is mainly confined to the warmer
waters of the Indo-Pacific Ocean. Carpopeltis maillardii (Montagne et Millardet) Chiang and C. beckeri
Schmitz are known from South Africa.The latter represents an invalidly published manuscript name, the
specimens on which it is based not belonging to Carpopeltis according to Papenfuss (see Silva et al. 1996).
Carpopeltis phyllophora bears little resemblance to
C. maillardii; the former being a rather large and supple species, the latter a small, cartilaginous species
that branches profusely near the apices [Okamura
1909, as C. rigida (Harvey) Schmitz; Børgesen 1943,
as C. rigida; Chiang 1970]. Comparison of the South
African specimens with the type specimen (BM, s.n.),
collected by Dr Jeannerett at Port Arthur, Tasmania,
reveals them to be morphologically very similar. It
should be noted, however, that the South African
specimens are rather variable with respect to thallus
width, with some being as narrow as 1–2 mm, whereas 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). Individual axes, 2–3 mm wide, have a plumose aspect, con-
Twenty marine benthic algae new to South Africa
secutive branchlets being of the same length. Laterals have a tendency to produce narrow pinnules, giving the fronds a distinctive fimbriate aspect (Fig. 47).
Ramuli form an alternate-distichous series of one
simple and one compound branch, cut off at an angle
of 45 ° (–60 °) (Fig. 48). Simple ramuli are acerose,
mostly straight, 2.5–3 (–4) mm long and 0.5–0.7 mm
wide. Margins of the ramuli are smooth. The central
axial filament is fairly visible after staining. Apices
are slightly to strongly incurved. Clusters of tetraspo-
425
rangial stichidia occur as adventitious axes in the axils 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 occasionally branched at their apices. No sexual thalli were
found.
Ecology: Occurring in the lower-intertidal pools
and 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-three
thallus (scale = 1 cm). Fig. 53. Detail of dendroid tetrasporangial stichidia (scale = 250 µm).
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
(16.viii.1999).
Discussion: This species has hitherto only been
recorded from southern Australia. The distinctive
morphology of the tetrasporangial stichidia and the
fimbriate aspect of the axes of the South African
specimens agree closely with the description of this
species as set out in Womersley (1994). On the other
hand, no serrations on the abaxial edge of the simple
ramuli were observed. This character, however, does
not appear to be fixed in the species. Also, the tendency of the proximal branchlets in an alternating series to become divided to a greater or lesser extent
was not observed as typical of the Australian plants
(G.T. Kraft, pers. comm.).This species is distinctly different from other South African species that have an
alternating-distichous series of two, having relatively
broad axes and relatively long, slender, clustered
stichidia.
Plocamium telfairiae (W. Hooker et Harvey) Harvey
ex 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 main
axes, 1 (–2) mm wide, are alternate-distichously
branched at an angle of 25–70 °. Ramuli form an alternate-distichous series of one simple and one compound branch. The latter are strongly incurved near
the apices. Stellate or dendroid tetrasporangial
stichidia are formed in the axils of the ramuli or replace a compound ramulus.
Ecology: Growing in intertidal rock pools and the
sublittoral fringe to a depth of 15 m in northern
Kwazulu-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); KZN
585: 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 and
differences in size of the axes.The density of branching
is 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 between consecutive branches, results in a greater degree of overlapping ramuli in type-one thalli. Simple
ramuli are subulate to triangular, mostly straight to
slightly incurved (rarely recurved). Simple ramuli in
type-one are generally longer (1.5–2.5 mm) and
broader (0.3–0.5 mm) than those of type-two (1–1.5
and 0.3 mm respectively). Compound ramuli are
strongly incurved at the apex, with moderate overlapping in type-one and none in type-two. Type-three
(Fig. 52) has a similar outline and structure, but is generally smaller (to 4.5 cm) and wider (1.5–2 mm). The
ramuli of the latter are sub-pinnate, with a compound
ramulus subopposite a simple one. The angle of
branching lies between that of type-one and type-two
(40–50 °). Similarly, the simple ramuli sizes lie between type-one and type-two, although they have a
wider base (0.5–1 mm). The central axial filament is
not visible in type-one and type-two (or only near the
apex), whilst it is in type-three. The tetrasporangial
stichidia were found in type-one and type-three,
whereas only female material was collected for typetwo. The former occur in the axils of ramuli or replacing compound ramuli and having a stellate / tree-shape
(Fig. 53).
The specimens examined agree with descriptions
and herbarium specimens of this species (including
an isotype specimen in BM). This is the first record of
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
(1964). The latter, however, reported the species only
from Mozambique and not from South Africa. This
species has a wide distribution within the Indian and
Pacific Oceans. There is also an Atlantic record from
Ghana 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, dichotomously
branched, 2 mm in diameter and densely covered by
assimilatory filaments. The medulla consists of entangled filaments (12–16 µm in diameter). The cortex is
filamentous with both short and long assimilatory filaments, which arise from inflated proximal cortex
cells (33–42 µm in diameter). The cruciately divided
tetrasporangia are borne apically on short assimilatory filaments. They are ovoid, 25–30 µm long and to
20 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); KZN
1685: Island Rock (14.viii.2000).
Discussion: Despite taxonomic confusion in the
past (see Huisman and Borowitzka 1990), tetrasporophytes of G. rugosa are easily recognisable by
their hirsute habit. Several species of Galaxaura and
the morphologically similar genus Tricleocarpa are
known from South Africa. Galaxaura diesingiana Zanardini, G. magna Kjellman and G. marginata (Ellis
et Solander) Lamouroux are all complanate. Galaxaura obtusata (Ellis et Solander) Lamouroux and Tricleocarpa fragilis (Linnaeus) Huisman et Townsend
Twenty marine benthic algae new to South Africa
are both terete, but are never hirsute and always lack
the inflated proximal cells of assimilatory filaments.
Galaxaura rugosa is widespread in the tropical Indian Ocean.
Discussion
Distribution patterns of tropical seaweeds are incomplete for many taxa. Several regions have received
very little attention and remain seriously un(der)explored. There are not only limited data on small
islands and archipelagos but also for several continental coastal regions such as Oman, Iran and Madagascar. This makes biogeographical assessments difficult, but a number of clear trends are indicated
(Table I).
The majority of the Phaeophyta and Rhodophyta
reported in this paper consist of widespread tropical
species. Asteronema breviarticulatum, Dictyota cervicornis, D. ciliolata and Galaxaura rugosa are mainly
pantropical species, occurring in the Atlantic
(Caribbean Sea) and Indian Oceans as well as the Pacific Ocean. Some are present along the Western
Australian coast. Next to the pantropical taxa, several species have a defined Indo-Pacific or Indian
Ocean distribution: Ceramium cingulatum, Euptilota
fergusonii, Halymenia durvillei and Phacelocarpus
tristichus. The presence of pantropical and Indo-Pacific species along the northern part of the KwazuluNatal coast is not surprising, as several of these
427
species have been reported from Inhaca in southern
Mozambique. Their presence in the northern part of
the Kwazulu-Natal coastline, therefore, only indicates a minor range extension. In contrast, the distribution range of Digeneopsis subopaca remains restricted to northern Kwazulu-Natal and the Inhaca
Peninsula in Mozambique. Together with Dasycladus
ramosus Chamberlain (1958), these species are probably endemic to the northern part of the overlap region between warm-temperate South African waters
and the tropical Indian Ocean.
Balliella crouanioides, Gibsmithia hawaiiensis,
Predaea weldii and Hypoglossum minimum are characterised by disjunct tropical Indo-Pacific distribution patterns. All three were originally described
from the central or western Pacific Ocean and have
been reported from a few scattered localities within
the Indian Ocean. Their small size and subtidal habitat probably contribute to the scarcity of reports. Balliella crouanioides and Gibsmithia hawaiiensis have
recently been reported for the East African coast
(Coppejans et al. 2000). Their presence along the
South African coast may indicate a wide distribution
within the Indo-Pacific region.
Carpopeltis phyllophora and Plocamium mertensii
are both Australian species. Norris and Aken (1985)
and Hommersand (1986) regard this South African
flora as having affinities with the western- and southern Australian algal floras, although there has been
some debate on what may have caused these floristic
similarities. According to Hommersand (1986), the
Table I. Known distributions of the species reported in this paper.
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
Mozambique
Western
Indian
Ocean
Central
Indian
Ocean
IndonesiaMalaysia
Australia
Pacific
Ocean
Atlantic
Ocean
–
+
–
–
+
+
–
–
+
–
+
+
–
–
–
?
+
–
+
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+
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+
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–
–
+
+
+
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+
?
+
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+
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+
+
+
+
+
+
+
–
–
–
–
+
+
+
–
?
–
–
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+
+
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+
+
+
+
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+
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+
+
+
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?
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+
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).
428
O. De Clerck et al.
genus Carpopeltis and to a certain extent Plocamium
represent examples of species clusters that originated
along the west and south coasts of Australia and migrated to South Africa during mid-to-late Miocene
periods (16–6Mya) marked by a general decrease in
temperature in the tropics. Euptilota articulata may
also represent an example of the South African-Australian connection, although the species has also been
reported from India and the North-West Pacific.
The distribution of Padina gymnospora is difficult
to interpret due to taxonomic confusion in the past.
Studies on the genus Padina along the East African
coast (Somalia to South Africa) have revealed that P.
gymnospora is absent along the tropical coasts of
Kenya, Tanzania and the various tropical archipelagos (Muylle 2000). The species is, however, present in
the northern part of the western Indian Ocean (Somalia, Yemen and Oman) as well as along the coast
of Kwazulu-Natal. The distribution pattern is interesting as both geographically isolated populations
are in habitats characterised by slightly cooler water
temperatures than occur in the tropical Indian
Ocean. Similar examples of disjunct distribution patterns between the northern Arabian Sea and the
South African east coast are offered by: Dictyopteris
macrocarpa (Areschoug) Schmidt, Ecklonia radiata
(C. Agardh) J. Agardh and Pseudocodium de-vriesii
Weber-van Bosse (Barratt et al. 1984, Barratt et al.
1986, Wynne 1999).
Acknowledgements
We would like to offer our sincere gratitude to Jean
Harris, Nonhlanhla Nxumalo, Bridget Armstrong,
Cloverly Lawrence and John Dives, of the KwazuluNatal Nature Conservation Services, for their full
support in organising everything, everytime, everywhere. We also extend our thanks to Peter Timm
from Triton Divers for his smoothly run services.
Funding for this project was provided by the International Scientific and Technological Cooperation
(BIL98/84) between the Ghent University and the
University of Cape Town and FWO Research Project
(3G002496). Further support was provided in South
Africa by Marine and Coastal Management, the National Research Foundation and the Department of
Environment and Tourism.
Accepted 19 May 2002.
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