Marine algae and seagrasses of the Dahlak Archipelago, southern

Transcription

Nova Hedwigia
75
1—2
1—90
Stuttgart, Aug ust 2002
Marine algae and seagrasses of the Dahlak Archipelago,
southern Red Sea
by
Y. Lipkin
Department of Botany, Tel Aviv University, Tel Aviv, Israel
and
P.C. Silva *
Herbarium, University of California, Berkeley, California 94720-2465, U.S.A.
With 7 figures and 4 tables
Dedicated to the memory of G.F. Papenfuss (1903-1981), distinguished scholar,
teacher, and colleague, who laid the groundwork for this paper.
Lipkin, Y. & P.C. Silva (2002): Marine algae and seagrasses of the Dahlak Archipelago, southern
Red Sea. - Nova Hedwigia 75: 1-90.
Abstract: Two Israeli expeditions to the Eritrean (then Ethiopian) coast in 1962 and 1965 obtained
127 specific and infraspecific taxa of benthic marine algae representing 73 genera and 4 species of
seagrasses representing 3 genera. Collections were made mainly in the Dahlak Archipelago, but also
on the adjacent mainland coast. The spring (1962) expedition yielded 94 species of algae, whereas
the autumn (1965) expedition yielded 77, with 44 present in both seasons. Among these algae, 13
genera and 47 species had not been recorded previously from the Red Sea. Some of these 47 species
have been reported subsequently from the northernmost part of the basin. There were 58 Rhodophyta,
33 Chlorophyta, and 31 Phaeophyta. Blue-green prokaryotes (Cyanophyta) were well represented in
the collections, but only 5 species were identified.
Two new species of algae are described: Pterocladia microscopica Lipkin et Papenfuss and Hypnea
anastomosans Papenfuss, Lipkin et Silva. Two other new species have been published elsewhere:
Turbinaria papenfussii W.R. Taylor and Chlorodesmis papenfussii Ducker. A new name, Sargassum
steinitzii, is proposed for S. tenue J. Agardh, non S. tenue (Kützing) Endlicher. Two new combinations
are made: Gelidiopsis capitata for Acrocarpus capitatus Kützing and Chondrophycus glanduliferus
for Chondria glandulifera Kützing.
*
Address for correspondence: [email protected]
DOI: 10.1127/ 0029-5035/ 2002/0075-0001
0029-5035/ 02/0075-0001
$ 22.50
© 2002 J. Cramer in der Gebrüder Borntraeger
Verlagsbuchha ndlung, D-14129 Berlin · D-70176 Stuttgart
1
Introduction
The benthic marine flora of the southern Red Sea in general and the Dahlak Archipelago in particular is very poorly known. Of 493 species and subspecific taxa of
benthic marine algae recorded from the Red Sea prior to 1968 (Papenfuss 1968a),
only 218 are attributed to the area south of the Suakim-Al Qunfudhah line (ca.
19°N). Only 164 had been reported from the Ethiopian coast and only 26 from the
Dahlak Archipelago. All of these numbers are inflated as a result of taxonomic
recognition by Grunow (1915, 1916) of many ecological forms of Sargassum.
The Red Sea, which is situated between the latitudes 13°N and 30°N, with its mouth
to the southeast, received its algae from nearby African and Arabian coasts, which
are occupied chiefly by members of the widespread tropical Indo-Pacific marine
flora. In Papenfuss’s catalogue (Papenfuss 1968a), 101 algae (including 57 taxa of
Sargassum) were recorded only from the southern Red Sea. Because the northern,
temperate reaches of the Red Sea have been well investigated, the absence of these
101 algae in that area is evidence that the richness of the marine flora of the Red Sea
decreases northward.
The Dahlak Archipelago is located in the southern Red Sea off the Eritrean coast,
between the latitudes 15°N and 17°N and the parallels 39°E and 41°E (Fig. 1). It is
situated on a rather shallow shelf ca. 150 km wide and 200 km long. Maximal depths
between the islands range mostly between 10 and 60 m, with the exception of the
Mesewa Channel, which runs between the Eritrean mainland and Dahlak Kebir Island. This channel has a general depth of ca. 120 m, reaching almost 160 m in a
small area west of Dur Gaarn Island. Waters surrounding the islands of the archipelago,
because of their tropical location and shallowness, are very warm, even at the bottom,
being between 27°C and 30°C away from the islands and reaching nearly 40°C in
very shallow bays around the islands. At such high temperatures the concentration of
dissolved gases, particularly O2 and CO2, are low. Salinities are even higher than in
the open Red Sea, which, with salinities of ca. 40 ppt, is the most saline open sea on
earth. The shallowness exaggerates fluctuations of water conditions in response to
changes in weather, thus stressing the marine biota. To the northeast of the archipelago
lies the deep main channel of the Red Sea from which the swell comes in to wash the
outer islands, providing somewhat better conditions for marine life. Further
information on the environment is given by Lipkin (1987).
In the early 1960s two Israeli expeditions studied the marine life, and incidentally
certain aspects of terrestrial life, in the Dahlak Archipelago. The first Israel South
Red Sea Expedition (ISRSE) visited the archipelago in March-April 1962, camped
on Entedebir Island (where much of the work was done), and visited several other
islands, all with rocky and sandy beaches and surrounded by shallow shelves. Most
locations sampled were sheltered from severe wave action and yielded only a few
species. Exceptions were Romia Island and Harmil Island, located at the eastern
edge of the archipelago and exposed to the swell from the deep waters of the main
navigation channel of the Red Sea. Here a rather diverse biota was found, a short
visit yielding many species not encountered in other parts of the archipelago. The
mangrove habitat was visited only briefly, at Melita Bay in the Gulf of Zula on the
Eritrean mainland.
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Fig. 1. Map of Dahlak Archipelago showing shore collection sites of the Israel South Red Sea
Expeditions, 1962 and 1965. (See Table 1 for legend to numbered collection sites.) Map redrawn
from nautical chart 62008, Red Sea sheet IV, Masamirit I. to Zubair Is., published by the Hydrographic
Office, Washington D.C., Dec. 1912.
The second Israel South Red Sea Expedition, which visited the archipelago in October
1965, provided a glimpse of the autumn flora compared with the spring flora sampled
by the first expedition. It camped (and carried out much of its work) on Museri
Island, which differs markedly from Entedebir Island in having much of its coast
occupied by mangroves. Although collections were made at many habitats similar to
those studied in 1962 (in some cases the same sites), additional islands with different
habitats were visited briefly. General information was published by Oren (1962) for
ISRSE 1962 and by Lewinsohn & Fishelson (1968) for ISRSE 1965. Ecological
aspects of the marine flora of the archipelago were summarized by Lipkin (1987).
The present paper is a systematic treatment of the marine algae and seagrasses collected
by the two Israeli expeditions in the Dahlak Archipelago and on the adjacent Eritrean
coast.
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Material and methods
ISRSE 1962 camped on Entedebir Island (No. 6 on Fig. 1 and Table 1) and ISRSE 1965 camped on
Museri Island (No. 11 on Fig. 1 and Table 1). Collecting on the shores of the base camp island was
undertaken: between 9 March and 6 April during ISRSE 1962, and between 7 and 27 October during
ISRSE 1965. On these islands marine algae and seagrasses were collected from several sites,
representing various habitats (Oren 1962; Lewinsohn & Fishelson 1968; Lipkin 1987). Nine other
islands and three mainland sites (Fig. 1 and Table 1) were also visited, but briefly, with only a few
hours devoted to collecting. Cundabilu Island, situated close to Entedebir Island, was visited twice,
allowing almost three full days of collecting.
The marine algae and seagrasses of the first expedition were collected by G.F. Papenfuss and, to a
lesser extent, by I.E. Friedmann, who unfortunately had to quit the expedition after only a few days
because of illness. A few samples were taken from the intertidal zone by C. Lewinsohn and L.
Fishelson. Collections in 1965 were made by Y. Lipkin, who also carried out ecological studies
(Lipkin 1987). The 1962 samples came predominantly from shallow water, and in 1965 SCUBA
technique was added in order to collect algae from subtidal habitats to depths of ca. 10 m. Dredging
from a trawler was used to get samples from depths between 10 and 120 m.
Samples taken during ISRSE 1962 were numbered with the prefix E62/, whereas those of ISRSE
1965 bear the prefix ISRSE 65/. Many samples included both pressed and liquid-preservedspecimens,
while others were in either one form or the other. Complete sets of specimens from both expeditions
are deposited at the Phycological Collection of the National Natural History Museum of Israel,
temporarily housed at Tel Aviv University (TELA). The first set of duplicates is deposited at the
University of California (UC). Other herbaria cited in this paper are abbreviated in accordance with
Holmgren et al. (1990).
Representatives of a few genera among the 1962 collections have been studied and the results have
been published, as follows: Turbinaria (Taylor 1964); Caulerpa (Taylor 1967), Chlorodesmis
(Ducker 1967, 1969), Cystoseira (Papenfuss & Jensen 1967), Hormophysa (Papenfuss 1968b),
Leveillea (Scagel & Chihara, 1968) and Sarconema (Papenfuss & Edelstein 1974). Halimeda was
studied by L. Hillis-Colinvaux, but not reported. Y.M. Chiang identified most other algae from the
first expedition, but a few were identified by M. Chihara and R.H. Simons.
Concerning the 1965 collections, W.R. Taylor identified some specimens of Caulerpa and confirmed
our determinations of others, F. Drouet identified Calothrix pilosa, M.D. Hoyle confirmed identifications of some Gracilaria species, and H.W. Johansen confirmed identifications of the articulated
corallines.
Two of the samples of ISRSE 1962 were made the types of newly described species: Turbinaria
papenfussii W.R. Taylor (1964) and Chlorodesmis papenfussii Ducker (1969).
Taxa not recorded from the Red Sea prior to the collections of the two Israeli expeditions are marked
with an asterisk.
SYSTEMATIC LIST
CYANOPHYTA
CHROOCOCCALES
CHAMAESIPHONACEAE
*CHAMAECALYX Komárek et Anagnostidis
*Chamaecalyx clavatus (Setchell et Gardner) Komárek et Anagnostidis (Arch.
Hydrobiol., Suppl. 73: 199, 1986).
Chamaesiphon clavatus Setchell et Gardner (Proc. Calif. Acad. Sci., ser. 4, 19: 118, pl. 4, fig. 1,
1930).
4
Table 1: Shore collection sites in the Dahlak Archipelago (plotted on Fig. 1).
Type from Isla Guadalupe, Baja California, Mexico, leg. H. Mason 63, CAS 173619
in UC.
ISRSE SPECIMENS STUDIED : Museri Island: in front of Scopus Ridge, 8.x.1965, on
Sphacelaria brachygonia and S. tribuloides (ISRSE 65/2333ea).
DISTRIBUTION: Reported from Islas Revilla Gigedo off the Pacific coast of Mexico
(Setchell & Gardner 1930); Indonesia (Geitler 1932); Vietnam (Pham-hoàng 1969),
and now the Red Sea. Probably widely distributed in warm Indo-Pacific waters.
COMMENTS : Our plants grew in loose clusters on the hosts. Cells were clavate, 3040 µm tall and 8-10 µm in maximum diameter.
OSCILLATORIALES
OSCILLATORIACEAE (nom. cons.)
LYNGBYA C. Agardh
Lyngbya confervoides C. Agardh (Syst. Alg., p. 73, 1824).
Type from Cádiz, Atlantic coast of Spain, in the Agardh herbarium at LD (see Drouet
1968, p. 264).
Turbinaria triquetra (ISRSE 65/2333f), 9.x.1965, on Acanthophora nayadiformis
(ISRSE 65/2351k), 23.x.1965, epilithic (ISRSE 65/3091).
DISTRIBUTION: Widespread in warm and temperate waters.
COMMENTS : Our plants had filaments 13-17 µm in diameter, trichomes 11-13 µm in
diameter and hyaline sheaths. They formed a rather loose growth on other algae,
whereas on rock they formed mats 1-2 mm thick.
5
This alga has been reported from the northern Red Sea (see records in Papenfuss
1968a; Lipkin 1972a, 1972b; Aleem 1978a; Potts 1980), but not from the southern
part.
Lyngbya majuscula (Dillwyn) Harvey (in Hooker Brit. Fl. 2(1) [= J.E. Smith,
Engl. Fl. 5(1)], p. 370, 1833).
Conferva majuscula Dillwyn (Brit. Conferv., p. 40, suppl. pl. A, 1809).
Type from England, in LD (see Drouet 1968, p. 263).
ISRSE SPECIMENS STUDIED : Entedebir Island: Landing Bay, 21.x.1965 (ISRSE
65/3135, 3139). Museri Island: in front of Scopus Ridge, 8.x.1965 (ISRSE 65/
2331), 10.x.1965, on Halimeda discoidea (ISRSE 65/2315c), 16.x.1965 (ISRSE
65/3072).
DISTRIBUTION : Widespread in warm and temperate waters.
COMMENTS: Our plants formed large masses. Their filaments were thick, 40-75 µm
in diameter, sometimes tending to curl. The trichomes were 35-67.5 µm in diameter,
with hyaline sheaths.
NOSTOCALES
RIVULARIACEAE
CALOTHRIX C. Agardh
*Calothrix pilosa Harvey (Smithsonian Contr. Knowl. 10(2) [Nereis Bor. Amer.
3]: 106, pl. XVIII.C, 1858).
Type from Key West, Florida, leg. W.H. Harvey, in BM (ex K) (see Drouet 1973, p.
39).
ISRSE SPECIMENS STUDIED : Museri Island: Rhizophor a Lagoon, 12.x.1965 , on
pneumatophores of Rhizophora mucronata (ISRSE 65/2386), on mud (ISRSE 65/
2387); south of Rhizophora Lagoon, 15.x.1965, on Avicennia marina pneumatophores
(ISRSE 65/0329, 2283-228 6). Melita Bay in the Gulf of Zula, 2.iv.1962, on
pneumatophores of Avicennia marina, along with Caloglossa leprieurii, Bostrychia
simpliciuscula and Brachytrichia lloydii (E62/20232c). Desta Island: 26.x.1965, on
A. marina pneumatophores (ISRSE 65/3170).
DISTRIBUTION : Widespread in warm seas.
COMMENTS: Our plants had filaments 18-30 µmm in diameter, green trichomes 1020 µm in diameter and thick yellow to brownish sheaths. Pseudobranching was not
common. They grew in sheltered habitats, forming mats on pneumatophores of
Avicennia marina and rhizophores of Rhizophora mucronata as well as on mud.
Mats on A. marina reached 1 cm in thickness, on R. mucronata only 2-3 mm, and on
mud 1-2 mm. On the upper parts of pneumatophores of A. marina they often occupied
sectors 6-10 cm long, giving the pneumatophores a bottle-brush appearance (Lipkin
6
1987, fig. 5). A similar, but less common situation occurs in Sinai, in the northern
Red Sea (Dor 1984). Dor regarded the Sinai material as belonging to either Scytonema
cincinnatum Thuret or S. polycystum Bornet et Flahault.
Part of sample ISRSE 65/0329, which was sent to Francis Drouet for determination,
was cited by him (Drouet 1973, p. 73) under Scytonema hofmanii C. Agardh (Syn.
Alg. Scand., p. 117, 1817) as Lipkin 2305 in PH. Calothrix pilosa is one of numerous
species referred by Drouet to S. hofmanii.
STIGONEMATALES
MASTIGOCLADACEAE
*BRACHYTRICHIA Zanardini
*Brachytrichia lloydii (Crouan) P.C. Silva (in Silva, Basson, & Moe, Univ. Calif.
Publ. Bot. 79: 78, 1996).
Rivularia lloydii P. et H. Crouan (Fl. Finistère, p. 117, 1867).
Type from Belle-Ile, France, ix.1854, in Lloyd, Algues de l’Ouest de la France, no.
303, in herb. Bornet-Thuret, PC (see Drouet 1973, p. 232).
ISRSE SPECIMENS STUDIED: Melita Bay, Gulf of Zula, 2.iv.1962, on pneumatophores
of Avicennia marina, along with Caloglossa leprieurii, Bostrychia simpliciuscula
and Calothrix pilosa (E62/20232b).
DISTRIBUTION: Warm eastern Atlantic waters, Mediterranean Sea, Red Sea.
C OMMENTS : Our plants formed small dark patches, 1-3 mm in diameter and
hemispherical when wet, on pneumatophores of Avicennia marina, in one instance
immediately below a sector covered by Bostrychia simpliciuscula.
This species has been reported from intertidal rocks and A. marina pneumatophores
in the northern Red Sea and the Suez Canal as Brachytrichia balani (Safriel &
Lipkin 1964; Lipkin 1972a, b; Dor, 1984), but the collections of the ISRSE are the
first from the southern Red Sea.
RHODOPHYTA
ERYTHROPELTIDALES
ERYTHROTRICHIACEAE
ERYTHROTRICHIA J. Areschoug (nom. cons.)
Erythrotrichia carnea (Dillwyn) J. Agardh (Lunds Univ. Års-Skr., Afd. Math. och
Naturvitensk. 19(2): 15, 1883).
Conferva carnea Dillwyn (Brit. Conferv., fasc. 12, pl. 84, 1807).
Type from Loughor [Llwchwr], West Glamorgan, Wales, U.K., leg. W.W. Young,
in BM (ex herb. Forster).
7
Ceramium flaccidum, (ISRSE 65/3084n).
DISTRIBUTION : More or less cosmopolitan.
ACROCHAETIALES
ACROCHAETIACEAE (nom. cons.)
ACROCHAETIUM Nägeli
*Acrochaetium gracile Børgesen (Dansk Bot. Ark. 3: 26, figs. 19, 20, 1915).
Type from St. Thomas Harbor, St. Thomas, U.S. Virgin Islands, on Sargassum vulgare,
presumably in C.
ISRSE SPECIMENS STUDIED : Entedebir Island: Mersat Abiad, 12.iii.1962, on a snail’s
shell with other microscopic algae (E62/20031a); Landing Bay, 21.x.1965, on
Gelidiopsis acrocarpa (ISRSE 65/3141fa).
COMMENTS: Our plants were 1-1.25 mm tall with filaments ca. 5 µm in diameter, cell
length 13-19 µm, and sporangia ca. 7.5 × 10 µm.
We follow the classification of the Acrochaetiaceae proposed by Feldmann (1962)
(see discussion by Silva et al. 1996, p. 94).
NEMALIALES
GALAXAURACEAE
GALAXAURA Lamouroux
Galaxaura rugosa (Ellis et Solander) Lamouroux (Hist. Polyp. Corall. Flex., p.
263, 1816).
Corallina rugosa Ellis et Solander (Nat. Hist. Zooph., p. 115, pl. 22, fig.3, 1786).
Type from Jamaica (presumably gametophyte) (see Papenfuss et al. 1982, p. 421).
ISRSE SPECIMENS STUDIED : Entedebir Island: outside Landing Bay to the north,
13.iii.1962, 3 m depth (E62/20035), northern shore of Ras Papenfuss, 25.iii.1962
(E62/20149).
COMMENTS: Our plants formed clumps about 40 cm in diameter. No Galaxaura was
found during the second ISRSE. Although several species are very common in the
northern Red Sea (Lipkin, unpublished; see Papenfuss 1968a, for localities), the
genus is rather rare in the Dahlak Archipelago. Actinotrichia fragilis, another member
of the Galaxauraceae that is very common in the northern Red Sea, is apparently
absent from the Dahlak Archipelago.
8
Our plants agree with Galaxaura lapidescens var. annuligera Piccone et Grunow (in
Piccone, Nuovo Giorn. Bot. Ital., 16: 312, 1884; type from the Bay of Assab, Eritrea)
regarded as a taxonomic synonym of G. subverticillata by Papenfuss et al. (1982,
pp. 424, 425). Galaxaura subverticillata in turn was synonymized under G. rugosa
by Huisman & Borowitzka (1990, p. 153).
Many records of G. lapidescens (Ellis et Solander) Lamouroux from the Red Sea
(see Papenfuss 1968a, p. 72), including De Toni & Levi (1888, p. 53) and Lipkin
(in Mergner & Schuhmacher 1974, pp. 263, 291, 303, misspelled G. lepidescens),
were probably based on G. rugosa.
GELIDIALES
GELIDIACEAE
GELIDIUM Lamouroux (nom. cons.)
Gelidium corneum (Hudson) Lamouroux (Ann. Mus. Hist. Nat. [Paris] 20: 129,
1813).
Fucus corneus Hudson (Fl. Angl., p. 474, 1762).
Type from Devonshire, England, leg. D. Miller, in BM (ex herb. Buddle, Sloane
collection) (see Dixon 1967).
ISRSE SPECIMENS STUDIED: Entedebir Island: entrance to Landing Bay, 15.iii.1962,
1 m depth, on a snail (E62/20053); Goliath Bay, 16.iii.1962 (E62/20069), 23.iii.1962
(E62/20153), 6.iv.1962 (E62/20258).
DISTRIBUTION : Nearly cosmopolitan.
COMMENTS : Our plants were to 2-3 cm tall, and fairly well branched.
Gelidium crinale (Hare ex Turner) Gaillon (Dict. Sci. Nat. [Levrault] 53: 362,
1828).
Fucus crinalis Hare ex Turner (Fuci, vol. 4, p. 4, pl. 198, figs. a-c, e-g, 1815).
Lectotype from Ilfracombe, Devonshire, England, leg. R. Hare, in BM (ex K) (see
Dixon & Irvine 1977a: 140).
ISRSE SPECIMENS STUDIED: Entedebir Island: Goliath Bay, 23.iii.1962 (E62/20154),
6.iv.1962, on Digenea simplex (E62/20257).
COMMENTS: Gelidium crinale and G. pusillum are traditionally recognized as two
distinct species, the former characterized as being caespitose and having filiform
branches, the latter as forming a turf and having flat, oval or lanceolate branches
(Feldmann & Hamel 1936, p. 107). They have been merged by Dixon & Irvine
(1977a, b), however, on the basis of a study that included sequential observations of
marked plants. We have no reason to doubt the validity of this conclusion, especially
9
when considering the fact that both type localities are in Devonshire, England, within
Dixon & Irvine’s study area. In the Red Sea, however, as in many other parts of the
world, the name G. pusillum has been applied to any very small turf-forming
Gelidium. It is unlikely that any of these recipients of the name is conspecific with
G. pusillum, which in Britain reaches a height of 15 cm. Realizing that taxonomic
problems within Gelidium cannot be solved in a floristic account, we follow
traditional, albeit faulty, species concepts in recognizing both G. crinale and G.
pusillum among collections at hand. Our plants of G. crinale reached 4 cm in height,
comparable with nearly fully developed plants in the northern Red Sea and eastern
Mediterranean.
Gelidium pusillum (Stackhouse) Le Jolis (Mém. Soc. Imp. Sci. Nat. Cherbourg
10: 139, 1863).
Fucus pusillus Stackhouse (Nereis Brit., p. 16, pl. VI, 1795).
Lectotype from Sidmouth, Devonshire, England, in BM (see Dixon & Irvine 1977a:
139).
ISRSE SPECIMENS STUDIED : Museri Island: off Mangrove Bay, 14.x.1965, on rock,
ca. 1 m depth (ISRSE 65/2397-tetrasporic, 3020g-tetrasporic); Camping Bay, off
Ras Pandion, 16.x.1965, on dendroid crustose coralline (ISRSE 65/3047g).
DISTRIBUTION : Cosmopolitan.
COMMENTS: Our plants were to 3 cm tall, but mostly only 10-15 mm tall, all bearing
tetrasporangia.
This species has been reported from the Red Sea only three times, all from the
northernmost part (Nasr 1947; Rayss & Dor 1963; Natour et al. 1979b).
Piccone & Grunow (in Piccone 1884) described G. pusillum var. conchicola from Mesewa,
Eritrea. In this variety, the axes were said to be somewhat broader (to 1 mm), more
complanate, and more proliferous than in other forms of the species. In our plants, the
axes were 1.5-2 mm wide, somewhat flat, and not much branched, thus agreeing with
the description and illustration given by Stackhouse (1795). Several varieties and formae
have been described within this morphologically variable species, but their taxonomic
value remains to be demonstrated (see Dixon & Irvine 1977b, p. 129).
*PTEROCLADIELLA Santelices et Hommersand
This genus was segregated from Pterocladia by Santelices & Hommersand (1997)
on the basis of the following suite of characters: intercalary carpogonia directed
towards both surfaces of the thallus; nutritive filaments growing centripetally and
forming a virtually solid cylinder around the central axis; and cystocarps usually
attached to one side of the cystocarp floor with chains of carposporangia on the
remaining three sides. The type is P. capillacea (S.G. Gmelin) Santelices et
Hommersand (Fucus capillaceus S.G. Gmelin), a well-known species widely distributed in warm temperate and subtropical waters. but not reported from the Red Sea.
10
*Pterocladiella caloglossoides (Howe) Santelices (J. Appl. Phycol. 10: 244, 1998).
Gelidium caloglossoides Howe (Mem. Torrey Bot. Club 15: 96, pl. 34, fig. 7, pl. 35, figs. 1-12,
1914).
Type from Isla San Lorenzo, Peru, Coker 59, in NY (see Dawson 1953, p. 76;
Stewart & J. Norris 1981).
ISRSE SPECIMENS STUDIED : Museri Island: Camping Bay, off Ras Pandion, 16.x.1965,
on dendroid crustose coralline, with new layers of coralline enveloping bases of
Pterocladiella (ISRSE 65/3047e-cystocarpic).
DISTRIBUTION: Widespread, but uncommon in most places where it occurs.
COMMENTS : Our plants were up to 1 cm tall, with flat, subpinnately branched fronds.
Branches were lanceolate to linear, 1-1.5 mm wide in their wider parts and bearing
cystocarps on both main axes and lateral branches.
Red Sea plants closely resemble Pterocladia parva Dawson (1953, p. 77, pl. 6, fig.
2; type locality: San Felipe, Gulf of California, Mexico). Although they are more
regularly branched than the type, they agree with specimens in UC from near the
type locality (Dawson 410, 7.ii.1946). Stewart & J. Norris (1981) considered P.
parva and P. caloglossoides conspecific.
*Pterocladiella microscopica Lipkin et Papenfuss, sp. nov.
Planta usque ad 2.5(-3) mm alta; frondibus erectis 100-250 µm latis, plerumque arcuatis, simplicibus
complanatis, linearibus vel spathulatis, interdum lanceolatis, raro teretibus, binis ad quaternis aggregatis,
ca. 0.5 mm distantis, e stolonibus repentibus 50-65 µm ortis; apices frondium nunc acuti nunc
rotundati, interdum parum impressi. Hapteron cylindraceus brevis opposito quoque fasciculus frondes
descendentis. Cystocarpia singula intra trientum superiorem frondium erectarum; ostiolis distantibus.
Tetrasporangia intra quartam parte superioram, 15-18 µm diam.
Plants to 2.5(-3) mm tall, composed of a prostrate system of creeping stolons 5060 µm diam., at intervals of ca. 1/2 mm giving rise to a group of 2-4 erect branches
on the upper side, and a short, descending, cylindrical attachment protuberance (“holdfast”) on the lower (“ventral”) side; erect branches simple, tending to curve to the
left or right from the point of issue (Fig. 2A), usually flattened, linear or spatulate,
sometimes lanceolate, rarely terete, 100-250 µm wide, apices pointed or rounded,
apex sometimes a little sunken. Cystocarps, one per erect branch, situated in its
upper third; ostiole usually at the distal end of cystocarp (Fig. 2B). Tetrasporangia
in upper fourth of branch, 15-18 µm in diameter (Fig. 2C).
Holotype: ISRSE E62/20031 (cystocarpic and tetrasporic), on a snail’s shell, with
many microscopic epiphytes, Mersat Abiad, Entedebir Island, Dahlak Archipelago,
leg. G.F. Papenfuss, 12.iii.1962, UC 1462284 (isotype in TELA).
ISRSE
SPECIMENS STUDIED :
see Holotype.
DISTRIBUTION: Known only from the type locality.
Among previously described species of Pterocladiella, only P. minima (Guiry &
Womersley) Santelices & Hommersand (1997) matches P. microscopica in its
11
Fig. 2. Pterocladiella microscopica (holotype, E62/20031): A, habit; B, cystocarpic branch; B’, side
view of cystocarp; B”, oblique top view of cystocarp; C, tetrasporangial branch. Scale bars: A =
500 µm, B and C = 200 µm, B’ and B” = 50 µm.
extremely small size. However, in that alga, which was originally described from
southern Australia as Gelidialla minima Guiry & Womersley (1992), the tetrasporangia
are arranged in distinctly diagonal rows, whereas in P. microscopica they form an
irregular pattern with indistinct diagonal rows (Fig. 3). It should be noted, however,
that both of these species are based on single collections so that the extent of variability
in the arrangement of tetrasporangia in either species is unknown.
12
Fig. 3. Pterocladiella microscopica (holotype, E62/20031): tetrasporangial sorus, surface view.
Scale bar = 100 µm.
GELIDIELLACEAE
GELIDIELLA J. Feldmann et G. Hamel
Gelidiella acerosa (Forsskål) J. Feldmann et G. Hamel (Rev. Gén. Bot. 46: 533,
1934).
Fucus acerosus Forsskål (Fl. Aegypt.-Arab., p. 190, 1775).
Type from Al Mukha, Yemen, southern Red Sea, leg. P. Forsskål, in C (see Børgesen
1932, pp. 5-6, pl. I; fig. 3.).
ISRSE SPECIMENS STUDIED : Entedebir Island: Padina Bay, 11.iii.1962, just below
intertidal zone, on shells (E62/20030); Goliath Bay, 23.iii.19 62 (E62/2017413
tetrasporic). Nocra Island: south side opposite village, 16.iii.1962, 1-2 m deep (E62/
20067).
DISTRIBUTION : Widespread in most warm seas.
COMMENTS: Our plants were well developed, to 4 cm tall, rather red compared with
their usual yellow colour in the shallow, highly illuminated habitats in the northern
Red Sea (Safriel & Lipkin 1964, p. 188, as Gelidium sp.).
*Gelidiella lubrica (Kützing) J. Feldmann et G. Hamel (Rev. Gén. Bot. 46: 535,
1934).
Acrocarpus lubricus Kützing (Phycol. General., p. 405, pl. 60.II, 1843).
Type from Napoli, Italy, in L, no. 941, 46...46 (see Womersley & Bailey 1970, p.
305).
ISRSE SPECIMENS STUDIED : Entedebir Island: Mersat Abiad, 26.iii.1962, on snail
shells (E62/20172 – sparsely tetrasporic). Museri Island: off Mangrove Bay,
14.x.1965, on a vermetid shell (ISRSE 65/3004).
DISTRIBUTION : Widely distributed in warm Indo-west Pacific waters and the Red Sea.
COMMENTS: Our plants covered rather densely the shells of snails, both mobile and
sessile (vermetids), sometimes while the animals still lived. They did not grow directly
on the shell, but on encrusting coralline algae, forming mats to 5 mm thick. Creeping
stolons were ca. 150 µm in diameter. The vertical branches emerging from the creeping
stolons were 10-12 mm tall, about 100-230 µm wide proximally and about 145175 µm wide distally. They were simple when young, the older branches becoming
sparsely subpinnately branched.
This alga has not been reported previously from the Red Sea.
*Gelidiella myrioclada (Børgesen) J. Feldmann et G. Hamel (Rev. Gén. Bot. 46:
533, 1934).
Echinocaulon myriocladum Børgesen (Kew Bull. 1934: 5, figs. 4, 5, 1934).
Type from Malabar Hill, Bombay, India, on rocks in exposed places near high water
mark, leg. F. Børgesen, no. 5235, in C.
ISRSE SPECIMENS STUDIED : Entedebir Island: northern shore of Ras Papenfuss,
25.iii.1962, forming a felt on sand-covered rock (E62/20179).
DISTRIBUTION : Vietnam, India, Tanzania, and now Red Sea. Probably widespread in
the Indian Ocean.
COMMENTS: Our plants bore many tetrasporangia in transverse series, as in G. pannosa
(J. Feldmann) J. Feldmann & Hamel 1934: 531, fig. 2).
14
CORALLINALES
CORALLINACEAE
AMPHIROA Lamouroux
*Amphiroa kuetzingiana Trevisan (Nomencl. Alg., p. 34, 1845).
Amphiroa verrucosa Kützing (Polyp. Calcif., p. 18, 1841), not A. verrucosa Lamouroux 1816.
Amphiroa verruculosa Kützing (Sp. Alg., p. 700, 1849) (superfluous substitute name).
Type from Split (Spalato), Croatia.
ISRSE
3003).
SPECIMENS STUDIED:
Museri Island: off Mangrove Bay, 14.x.1965 (ISRSE 65/
DISTRIBUTION: Mediterranean and Red Sea.
COMMENTS : Only one plant was encountered. It was 1.5 cm tall and bore many tetrasporangiate conceptacles. The axes were 200-250 µm in diameter, regularly dichotomously branched at the geniculum or along the intergeniculum. The segments
were 1.8-3.2 mm long. The genicula consisted of two layers of cells, those in the
lower layer being 120-160 µm long and those in the upper 20-40 µm long.
The existence of Amphiroa kuetzingiana Trevisan 1845 as a substitute name for the
later homonym A. verrucosa Kützing 1841, with priority over A. verruculosa Kützing
1849, was overlooked by Weber-van Bosse (1904, p. 101) and subsequent authors.
JANlA Lamouroux
Jania adhaerens Lamouroux (Hist. Polyp. Corall. Flex., p. 270, 1816).
Type stated by Lamouroux to be from the Mediterranean, but with uncertainty.
ISRSE SPECIMENS STUDIED : Cundabilu Island: western end of islet, 20.iii.1962 (E62/
20100-male, female and tetrasporic). Enteraia Island: 24.iii.1962, on Cystoseira
myrica (E62/20159). Entedebir Island: Goliath Bay, 14.iii.1962, 1 m depth (E62/
20046); Landing Bay, 21.x.1965, on lower part of Halimeda discoidea in shallow
water (ISRSE 65/2314b). Desta Island: off the northern coast, 26.x.1965, rocks at
shallow water (ISRSE 65/3169d). Museri Island: in front of Scopus Ridge, 10.x.1965,
epilithic and on H. discoidea, Padina boergesenii and C. myrica (ISRSE 65/2315a,
2366i, 2368f, 2373a); 9.x.1965, in Caulerpa racemosa community, epilithic and on
Avrainvillea amadelpha and Caulerpa racemosa (ISRSE 65/2343d, 2344f, 2347b),
14.x.1965, on H. discoidea (ISRSE 65/2317e), 19.x.1965, epilithic (ISRSE 65/
3085i); off Mangrove Bay, 14.x.1965, epilithic and on Chondria (ISRSE 65/3004h,
3033f, 3038f, 3040); Camping Bay, off Ras Pandion, 16.x.1965, epilithic (ISRSE
65/3047a, 3054d, 3067b-cystocarpic). Romia Island: 29.iii.1962, epilithic and on
Digenea simplex (E62/20199a).
DISTRIBUTION: Widespread in most warm seas.
COMMENTS : The alga, a rather common epiphyte and lithophyte in the southern Red
Sea, was, however, much less abundant than in many habitats in the eastern
15
Fig. 4. Jania adherens (E62/20100): A, B, thalli with gall-like growths of uncertain origin; C,
deformations in close view. 1, normal conceptacle-bearing terminal intergeniculum; 2, normal vegetative terminal intergeniculum;3, deformed conceptacle-bearingterminal intergeniculum;4, proliferous
terminal intergenicula. Scale bars = 1 mm.
16
Mediterranean. Tetrasporophytes and gametophytes of both sexes were collected in
spring, but only female plants were obtained in autumn.
Some of our plants had deformations on various parts of their thalli (Fig. 4A). The
cause of these deformations (Fig. 4, B, C), which appeared on both sterile and fertile
plants, is unknown.
GIGARTINALES
GRACILARIACEAE
GRACILARIA Greville (nom. cons.)
Gracilaria arcuata Zanardini (Mem. Reale Ist. Veneto Sci. Lett. Arti, 7: 265, pl. V,
fig. 2, 1858).
Type from Al ’Aqabah, Jordan, leg. Portier, in herb. Zanardini, Museo Civico di
Storia Naturale, Venezia (see De Toni & Levi 1888).
ISRSE SPECIMENS STUDIED : Museri Island: Camping Bay, 19.x.1965, on rock flat ca.
30 cm below low water (ISRSE 65/2200, 2223); in front of Scopus Ridge, 9.x.1965,
on stones in Caulerpa racemosa community growing on calcareous sand, ca. 30 cm
below low water (ISRSE 65/2336), on sponge, ca. 15 cm below low water (ISRSE
65/2241, 2242); off Mangrove Bay, 14.x.1965 (ISRSE 65/3010).
DISTRIBUTION: Warm waters of Red Sea and western Indian Ocean.
COMMENTS : Our plants were to 4.5 cm tall and sterile. They grew on stones and other
solid substrates in shallow water, but were not common.
Gracilaria bursa-pastoris (S.G. Gmelin) P.C. Silva (Univ. Calif. Publ. Bot. 25:
265, 1952).
Fucus bursa-pastoris S.G. Gmelin (Hist. Fuc., p. 121, pl. VIII, fig. 3, 1768).
Type from the Mediterranean Sea.
ISRSE
SPECIMENS STUDIED :
Nocra Island: 16.iii.1962 (E62/20095).
DISTRIBUTION: Widespread in all warm waters.
COMMENTS : Our plants were to 12 cm tall and bore tetrasporangia. This alga has been
previously reported from the Red Sea under the name Gracilaria compressa (C.
Agardh) Greville by Newton (1953, pp. 396, 414).
Gracilaria canaliculata Sonder (Abh. Naturwiss. Naturwiss. Verein Hamburg, 5(2):
56, 1871) (treated as a new name for Sphaerococcus canaliculatus Kützing, Tab.
Phycol. 18, p. 29, pl. 82, figs. d, e, 1868; not S. canaliculatus C. Agardh, Sp. Alg.
1, p. 260, 1822).
Type from Wagap, New Caledonia, leg. Vieillard.
17
ISRSE SPECIMENS STUDIED: Cundabilu Island: western end of islet, 20.iii.1962 (E62/
20073a). Entedebir Island: outside Landing Bay to the north, 13.iii.1962, 3 m depth
(E62/20039); Landing Bay, 21.x.1965, in shaded cavelet in fossil reef limestone, ca.
50 cm depth, with cystocarps and with parasitic Gracilariophila sibogae (ISRSE 65/
0374a, 3114), with tetrasporangia (ISRSE 65/2219). Museri Island: in front of Scopus
Ridge, 10.x.1965 (ISRSE 65/2371), 19.x.1965 (ISRSE 65/2331a – young plants,
3087); Camping Bay, off Ras Pandion, 16.x.1965 (ISRSE 65/2227, 3050).
DISTRIBUTION : Warm Indo-west Pacific waters.
COMMENTS: Our plants were to 4.5 cm tall in both spring and autumn. In the spring
only tetrasporangial plants were collected, whereas in autumn cystocarpic as well as
tetrasporic plants were obtained. Gracilariophila sibogae was present on plants
collected at Entedebir Island in 1965. The plants usually grew on rocks in shallow
waters, down to 3 m depth.
Gracilaria debilis (Forsskål) Børgesen (Dansk Bot. Ark. 8(2): 7, 1932).
Fucus debilis Forsskål (Fl. Aegypt.-Arab., p. 191, 1775).
Type from Al Mukha, Yemen, southern Red Sea, leg. P. Forsskål, in C (see Børgesen,
1932, p. 7, pl. I, fig. 5; Bird et al., 1986, p. 2046, fig. 2).
ISRSE
SPECIMENS STUDIED :
Entedebir Island: Goliath Bay, 23.iii.1962 (E62/20168).
DISTRIBUTION : Indian Ocean and Red Sea (see note below).
COMMENTS: Our plants were sterile, to 4.5 cm tall, with missing apices – probably
the result of grazing by fish. Bird et al. (1986, p. 2047) examined these plants
(which they cited as UC 1461727 and UC 20168), compared them with the holotype,
and concluded that “in structure these are... quite like the Forsskål collection”, thus
confirming our provisional identification.
Gracilaria debilis was transferred to the genus Polycavernosa Chang et Xia (1963)
by Fredericq & J. Norris (1985) after they had studied material from Belize. Bird et
al. (1986), however, showed that the western Atlantic alga known for a long time as
G. debilis is in fact G. cornea J. Agardh.
*Gracilaria dura (C. Agardh) J. Agardh (Alg. Medit. Adriat., p. 151, 1842).
Sphaerococcus durus C. Agardh (Sp. Alg. 1(2), p. 310, 1822).
Type from Cádiz, Atlantic coast of Spain; in LD.
ISRSE
SPECIMENS STUDIED:
Romia Island: 29.iii.1962 (E62/20185).
DISTRIBUTION : Mediterranean Sea and Atlantic coast of Spain and Morocco; Red Sea;
western Indian Ocean.
COMMENTS : Many plants (all sterile) were collected, with a maximum height of
12 cm. Some bore Gracilariophila sibogae as a parasite.
18
*GRACILARIOPHILA Setchell et Wilson
*Gracilariophila sibogae Weber-van Bosse (Siboga-Exped. Monogr. 59d, p. 443,
1928).
Type from Dongkalang (Dongala), Teluk Tomini (Palos Bay), Sulawesi, Indonesia,
on Gracilaria arcuata.
ISRSE specimens studied: Entedebir Island: Landing Bay, 21.x.1965, on Gracilaria
canaliculata in shaded cavelet in fossil reef limestone, ca. 50 cm depth (ISRSE 65/
0374, 3114a). Romia Island: 29.iii.1962, on Gracilaria dura (E62/20185a).
DISTRIBUTION: Warm Indian Ocean waters and the Red Sea.
COMMENTS : Our plants were stalked, the largest being 6 mm in diameter and to 4 mm
high, including the stalk (see Weber-van Bosse, 1928, fig. 181). Most of the small
plants were smooth, whereas most of the larger ones had broad, rounded
protuberances. Our plants are clearly referable to Gracilariophila, but the absence
of reproductive structures makes it impossible to determine which of the six described
species they represent, if any. On the basis of the size and shape of the papillae, we
tentatively assign our plants to G. sibogae even though they are much larger than the
type, which was 1.5 mm in diameter and 1.5 mm high.
SOLIERIACEAE
SARCONEMA Zanardini
Sarconema filiforme (Sonder) Kylin (Lunds Univ. Årsskr., N.F., Avd. 2, 28(8):
22, 1932).
Dicranema filiforme Sonder (Bot. Zeitung 3: 56, 1845).
Type from western Australia, Preiss 2557, in MEL (see Papenfuss & Edelstein 1974).
ISRSE SPECIMENS STUDIED: Entedebir Island: Landing Bay, 11.iii.1962, ca. 1 m depth
(E62/20003-tetrasporic) (erroneously cited by Papenfuss & Edelstein, 1974, as E62/
2003); outside Padina Bay to the north, 13.iii.1962, 3 m depth (E62/20036-tetrasporic,
E62/20037); Goliath Bay, 23.iii.1962 (E62/20167). Museri Island: Camping Bay,
19.x.1965, on rock flat ca. 30 cm below low water (ISRSE 65/2201). Harmil Island:
28.iii.1962 (E62/20287-tetrasporic). (All except the Museri Island collection were
cited by Papenfuss & Edelstein, 1974.)
DISTRIBUTION: Widespread in warm Indo-west Pacific waters.
COMMENTS: Plants obtained in spring were large (to 16 cm high) and bore tetrasporangia, whereas those collected in autumn were small (to 8 cm high) and sterile.
These differences suggest that in the southern Red Sea fronds begin development in
summer and reach maturity the following spring or early summer.
This alga was reported previously from the Dahlak Archipelago by Papenfuss (1968a:
88) as Sarconema furcellatum Zanardini, a name later placed in the synonymy of S.
filiforme by Papenfuss & Edelstein (1974).
19
*Sarconema scinaioides Børgesen (Kew Bull. 1934: 12, 1934).
Type from the vicinity of Karachi, Pakistan, leg. Kotwal, in NY (see Papenfuss &
Edelstein 1974).
ISRSE SPECIMENS STUDIED : Entedebir Island: Landing Bay, 21.x.1965, entrance to
cavelet in fossil reef limestone, ca. 1 m depth (ISRSE 65/3136).
DISTRIBUTION : Western Indian Ocean and the Red Sea.
COMMENTS : Our plants were juvenile, only 4 cm tall, with terete axes 2-4 times
bifurcate, 1.5 mm in diameter at the base, tapering towards the apex. The peripheral
cortical cells were small, oblong, 12-15 µm by 5-6 µm; the inner cortical cells were
large, irregularly isodiametric, 60-105 µm in diameter. The medulla was composed
of distinctive thick-walled filaments, 8-12 µm in diameter in cross section.
WURDEMANNIACEAE
*WURDEMANNIA Harvey
*Wurdemannia miniata (Sprengel) J. Feldmann et G. Hamel (Rev. Gén. Bot. 46:
544, 1934).
Sphaerococcus miniatus Sprengel (Syst. Veg., ed. 16, 4(1): 340, 1827) (see Silva et al. 1987, p. 72
for a discussion on the correct basionym).
Type from the vicinity of Montpellier, French Mediterranean coast, leg. Draparnaud.
on rock flat ca. 30 cm below low water (ISRSE 65/2228).
COMMENTS: Our plants were subpinnately branched, to 18 mm tall, and sterile.
HYPNEACEAE
HYPNEA Lamouroux
*Hypnea anastomosans Papenfuss, Lipkin, et Silva, sp. nov.
Plantae caespitosae, ca. 2-3(-4) cm altae; axes prope bases plantae 200-500 µm diam. gradatim ad
180 µm prope peripheriae plantae decrescentes, saepe inter se ubi tangentes anastomosantes, repentes,
haptera adventitia discoidea brevissima substratis et algis ceteris ubi tangentes formantes; ramuli
plerumque breves, 200-500 µm longi, gradatim in apices acuti decrescentes, interdum in rami
crescentes. Cystocarpia subsphaericae, ca. 250-280 µm diam. plerumque partibus inferioribus
ramulorum insidentibus. Tetrasporangia 40-87 µm longae, 22-29 µm diam. in partibus superioribus
et interdum in partibus centralibus ramulorum; circumcirca peripheriam, interdum non nisi in partes
peripheriarum.
Red Sea plants growing usually on rocks, but sometimes as epiphytes on other algae;
forming small entangled cushions 2-3(-4) cm high, with the main axes irregularly
ramified, branches thin, 250-500 µm in diameter near the base of plant, gradually
20
tapering to about 180 µm in diameter near their periphery, often anastomosing when
in contact with one another (Fig. 5A), creeping and forming adventive, short, discoid
holdfasts when in contact with a rocky substrate or neighboring alga; branchlets
usually short, 0.2-0.5 mm long, gradually tapering to a pointed apex, sometimes
elongating to become a part of the main axial system. Cystocarps subspherical, ca.
250-280 µm in diameter, usually borne on the lower parts of branchlets (Fig. 5B).
Tetrasporangia borne at the distal part of branchlets (stichidia), sometimes at the
central part, usually all around the branchlet, sometimes not on the entire
circumference. Mature tetrasporangia 40-87 µm long, 22-29 µm in diameter
(Fig. 5C).
Holotype: ISRSE 65/2346 (tetrasporic and cystocarpic), in Caulerpa racemosa
community in front of Scopus Ridge, Museri Island, Dahlak Archipelago, leg. Y.
Lipkin, 9.x.1965, in TELA (isotype in UC).
ISRSE SPECIMENS STUDIED: Entedebir Island: Landing Bay, 21.x.1965 (ISRSE 65/
3122a, 3139a – both cystocarpic); Museri Island: in front of Scopus Ridge, 9.x.1965,
in Caulerpa racemosa community (ISRSE 65/2346 – tetrasporic and cystocarpic);
10.x.1965 (ISRSE 65/2380m-tetrasporic); off Mangrove Bay, 14.x.1965 (ISRSE
65/2398d-cystocarpic); Camping Bay, off Ras Pandion, 16.x.1965, (ISRSE 65/3047b).
DISTRIBUTION: Widespread in warm seas.
COMMENTS : The name Hypnea esperi Bory (1828 [1826-1829], p. 157) has been
widely applied to this small, broadly distributed, tropical to subtropical species (e.g.,
Børgesen 1924, 1950; Tanaka 1941; Dawson 1944, 1954, 1956, 1957; Rayss & Dor
1963; Pham-hoàng 1969; Chapman 1971; Egerod 1971; Isaac, 1971; Lipkin 1972a,
b). Papenfuss (1964, p. 38), however, pointed out that this name is an illegitimate
substitute for Fucus nootkanus Esper, a name applied to a species of uncertain identity.
Bory’s plant (from Concepción, Chile) was first given a valid name, at varietal
level, by J. Agardh (1851 [1851-1863], p. 442) as H. musciformis var. esperi. The
small tropical alga in question, however, differs markedly from Bory’s plant, which
was described as being 15 cm tall. It also differs from the small Pacific plant (probably
from Japan) illustrated by Kützing (1868, p. 9, pl. 26) under the name H. esperi.
Dawson (1961, p. 234) examined Kützing’s material and concluded that it was a
small variant of H. cervicornis J. Agardh. Thus, the small tropical alga in question
remains undescribed and unnamed.
This alga has been reported, as H. esperi, from the northern Red Sea (Rayss & Dor
1963; Aleem 1978b, 1978c; Lundberg & Lipkin 1979) and the Suez Canal (Lipkin
1972a, 1972b; Aleem 1984). It is reported here for the first time from the southern
part of this sea.
Hypnea cornuta (Kützing) J. Agardh (Sp. Alg. 2(2), p. 449, 1851).
Chondroclonium cornutum Kützing (Sp. Alg., p. 741, 1849).
Type locality stated by Kützing to be unknown; “ad oras Guineae” fide J. Agardh
(l.c.), in herb. Lenormand, in PC.
21
Fig. 5. Hypnea anastomosans. (holotype, ISRSE65/2346): A, habit showing anastomosing branches;
B, cystocarpic branch; C, tetrasporangial branch. Scale bars: A = 1 mm, B and C = 500 µm.
ISRSE SPECIMENS STUDIED : Entedebir Island: Padina Bay, 12.iii.1962, (E62/20022cystocarpic); Mersat Abiad, 26.iii.1962 (E62/20165); Goliath Bay, 3.iv.1962 (E62/
20213), 6.iv.1962, intertidal (E62/20255). Museri Island: in front of Scopus Ridge,
8.x.1965 (ISRSE 65/2326h, 2330f, 2331o), 10.x.1965, (ISRSE 65/2315e, 2368i,
2380d), 9.x.1965, on rock flat ca. 20 cm below low water, in Chondrophycus
papillosus community (ISRSE 65/0385, 2337, 2341g), on rock flat ca. 1 m below
low water, in Caulerpa racemosa community (ISRSE 65/2353, 2345h-with parasitic
Hypneocolax); off Mangrove Bay, 14.x.196 5 (ISRSE 65/3020-with parasitic
Hypneocolax, 3024, 3035a); Camping Bay, 19.x.1965 (ISRSE 65/2221, 2222tetrasporic, 3080d). Romia Island: 29.iii.1962 (E62/20201-tetrasporic, 20283). Harmil
Island: 28.iii.1962 (E62/20248, 20282).
DISTRIBUTION : Widespread in all warm seas.
COMMENTS: This species was one of the most common algae in the Dahlak Archipelago.
It often grew epiphytically on a variety of other algae, in a wide range of usually
well-illuminated habitats in shallow water. Well-developed plants, up to 15 cm tall,
were collected both in spring and in autumn. Plants bearing cystocarps were found
only in spring, whereas tetrasporophytes were found in both spring and autumn.
Reproductive as well as sterile plants from the two seasons bore great numbers of the
stellate propagules typical of the species. Many propagules in initial stages of
development were found attached to other algae.
22
Hypnea valentiae (Turner) Montagne (in Barker-Webb & Berthelot, Hist. Nat. Iles
Canaries, p. 161, 1841).
Fucus valentiae Turner (Fuci, vol. 2, p. 17, pl. 78, 1809).
Type from the Red Sea, leg. Lord Valentia, in BM (ex K).
ISRSE SPECIMENS STUDIED : Museri Island: off Mangrove Bay, 14.x.1965, on rock
flat, ca. 1 m below low water (ISRSE 65/3033b-tetrasporic); Camping Bay, 19.x.1965,
on rock flat gently sloping seawards, ca. 30 cm below low water (ISRSE 65/2209tetrasporic, 3085-cystocarpic and tetrasporic).
COMMENTS: Our plants were small, to 5 cm tall, with rather delicate ultimate branchlets
(stichidia). Tetrasporangia were usually borne on the central part of branchlets,
sometimes a little towards the apex, the basal and distal parts being thinner.
Until the confusing interrelationships of Hypnea valentiae, H. cornuta and H.
hamulosa (see Papenfuss 1958; 1968a, p. 88) are disentangled, we prefer to treat H.
valentiae and H. cornuta as separate species.
*HYPNEOCOLAX Børgesen
*Hypneocolax stellaris Børgesen f. orientalis Weber-van Bosse (Siboga-Exped.
Monogr. 59d, p. 393, fig. 143, 1928).
Type from Dobo, Pulao Wokam, Kepulauan Aru (Aru Islands), Indonesia, in L (see
Dawson 1961, p. 234).
Hypnea cornuta from a Chondrophycus papillosus community growing on a shallow
rock flat gently sloping seawards (ISRSE 65/2345i); off Mangrove Bay, 14.x.1965,
on H. cornuta epiphytic on Laurencia on rocks 1 m depth (ISRSE 65/3020g), on H.
cornuta growing on a reef knoll, 1.5 m depth (ISRSE 65/3033d).
DISTRIBUTION: Reported only from the type locality, Japan (Segawa 1941), Hawaii
(Fan 1961), Pacific Mexico (Dawson 1961), Tanzania (Jaasund 1976), Australia
(Womersley 1994) and now the Red Sea. Probably widespread in warm Indo-Pacific
waters.
COMMENTS : This forma differs from the typical form described by Børgesen (1920)
from the Caribbean Sea by producing tetrasporangia rather than bisporangia (Weber-van Bosse 1928, p. 393). Our material was sterile and its assignment to f. orientalis
is based primarily on the geographical distribution of this forma. Womersley (1994,
p. 448) treated the Pacific plants as a distinct subspecies, regarding the difference
between them and the Caribbean ones as “certainly of greater degree than that of
forms”. Many cases of the occurrence of both bi- and tetraspores in the same species
were listed by Bauch (1937, pp. 366-367) in different orders of the Rhodophyta.
Woelkerling (1988, p. 26) cited a few more examples of corallines in which such
23
situation occurs. In some cases bi- and tetraspores were found on the same plants and
even in the same conceptacles, in others on the same plants in different seasons, and
in still others in different localities (Cole & Sheath 1990). Dixon (1973, p. 134)
recommended that bisporangia be interpreted cautiously. We agree and therefore
treat these populations of H. stellaris as belonging to two forms rather than subspecies.
Genetic studies would provide a basis for deciding the appropriate level of taxonomic
separation, if any. This parasite infests several species of Hypnea, namely, H.
musciformis (Børgesen 1920), H. variabilis (Segawa 1941), H. nidifica? (Fan 1961),
H. valentiae (Dawson 1961), H. ramentacea (as H. episcopalis, Womersley 1994, p.
448), and with this report also H. cornuta.
RHODYMENIALES
RHODYMENIACEAE
BOTRYOCLADIA (J. Agardh) Kylin, (nom. cons.)
*Botryocladia skottsbergii (Børgesen) Levring (in Skottsberg, ed., Nat. Hist. Juan
Fernandez, vol. 2(5), p. 645, 1941).
Chrysymenia skottsbergii Børgesen (in Skottsberg, ed., Nat. Hist. Juan Fernandez, vol. 2(3), p. 307,
figs. 49, 50, 1924).
Type from Isla de Pascua (Easter Island), in C (see Womersley & Bailey, 1970).
ISRSE SPECIMENS STUDIED: Entedebir Island: Landing Bay, 12.iii.1962, on dead coral
(E62/20015-cystocarpic), 27.iii.1962 (E62/20181-cystocarpic), 21.x.1965, in shaded
cavelet in coral rock, ca. 50 cm below low water (ISRSE 65/0375, 3134-tetrasporic).
DISTRIBUTION : Warm Indo-Pacific waters.
C OMMENTS : Our plants were to 3.5 cm tall, with stalks 2 or 3 times irregularly
dichotomously branched. Each branch ended with a vesicle. The vesicles, to 7 per
main stalk, were irregular in shape, mostly elongate, 4-7 mm × 3-4 mm. Often
several stalks grew side by side.
The plants growing in shaded habitats were dark red when collected.
The ISRSE samples constitute the first record of B. skottsbergii from the southern Red
Sea (Lipkin 1987), although it seems that the species is not rare in the area, but rather
dwells in a relatively inaccessible habitat. It has also been found on the Sinai coasts in
the northern Red Sea (Lipkin 1979) and probably occurs throughout the Red Sea.
*COELOTHRIX Børgesen
*Coelothrix irregularis (Harvey) Børgesen (Dansk Bot. Ark. 3: 389, 1920).
Cordylecladia ? irregularis Harvey (Nereis Bor. Amer. 2, p. 156, 1853).
Type from Key West, Florida, U.S.A.
ISRSE SPECIMENS STUDIED : Entedebir Island: Landing Bay, 21.x.1965, (ISRSE 65/
3136g, 3141p-tetrasporic).
24
COMMENTS : Our plants were 4-5 cm tall, somewhat irregularly secundly branched.
Their cavity was wider in the younger parts than in the older ones. Gland cells were
not common. Tetrasporangia were present in autumn.
*GELIDIOPSIS Schmitz
The taxonomic placement of Gelidiopsis and its relation to Ceratodictyon are discussed
by Silva et al. (1996, p. 355).
*Gelidiopsis capitata (Kützing) comb. nov.
Acrocarpus capitatus Kützing (Tab. Phycol. 18, p. 12, pl. 35, figs. a, b, 1868).
Type from New Caledonia, leg. Vieillard.
ISRSE SPECIMENS STUDIED : Entedebir Island: Landing Bay, 21.x.1965, in shaded
cavelet in rock, ca. 50 cm below low water, with epiphytic Martensia elegans and
Dictyota (ISRSE 65/3116-tetrasporic).
DISTRIBUTION: New Caledonia and southern Red Sea, probably in the entire Indowest Pacific.
COMMENTS : Our plants were composed of a prostrate system of stolons, ca. 270300 µm in diameter, giving rise to erect, simple or very slightly ramified branches,
ca. 270-300 µm in diameter near their bases and 200-225 µm right below the swollen,
terminal tetrasporangia-bearing stichidia, which were ca. 700 µm in diameter at
their widest part (Fig. 6).
Acrocarpus capitatus was placed tentatively in synonymy with Gelidiopsis intricata
(C. Agardh) Vickers by Dawson (1954, p. 424) and Cribb (1983, p. 56). Norris
(1987, p. 245), while suggesting the merger of Gelidiopsis and Ceratodictyon under
the latter name, followed this view without examining Kützing’s type. In describing
Sphaerocarpus intricatus, however, C. Agardh (1822 [1822-1823], p. 333) indicated
that the fronds were “caespitosa maxime intricata ...”. Our plants, although they may
be regarded as caespitose, formed a rather loose growth which can hardly be
interpreted as intricate. They were much less branched than the specimen illustrated
as A. intricatus by Kützing (1868, pl. 35, figs. d, e). Furthermore, there is a difference
between the two species with regard to the ratio of diameters of a stichidium and the
branch bearing it. For G. intricata the ratio is 2.25-2.5 (Setchell 1924, p. 163, fig.
31; Yamada & Tanaka 1938, p. 73, fig. 6a; Dawson 1954, p. 424, fig. 34c), whereas
in G. capitata it is about 3.5. Mature stichidia of G. capitata are ovate or parabolic
rather than broadly lanceolate like immature stichidia of that species and mature
stichidia of G. intricata.
The plants from Tanzania, reported and illustrated by Jaasund (1976) as G. intricata,
seem to belong also to Gelidiopsis capitata.
25
Fig. 6. Gelidiopsis capitata (ISRSE65/3116): branch with mature stichidia. Scale bar = 1 mm.
*Gelidiopsis repens (Kützing) Weber-van Bosse (Siboga Exped. Monogr., 59(d),
p. 425, 1928).
Gelidium repens Kützing (Tab. Phycol. 18, p. 21, pl. 60: figs. a, b, 1868).
Type from Wagap, New Caledonia.
tunnel in coral reef, ca. 50 cm below low water, in sciaphilic algal community
(ISRSE 65/0383, 3141f).
COMMENTS: Our plants, all sterile, were 3-4 cm tall, terete below and flattened above,
and dichotomously to irregularly branched. They agree best with descriptions and
illustrations of Gelidiopsis scoparia as given by Børgesen (1952, fig. 13) and Phamhoàng (1969). Børgesen later (1954, p. 28) referred his plants to G. acrocarpa, a
species considered by Feldmann (1931, p. 157) to be conspecific with G. repens.
CHAMPIACEAE
CHAMPIA Desvaux
Champia irregularis (Zanardini) Piccone (Nuov. Giorn. Bot. Ital. 16: 313, 1884).
Lomentaria irregularis Zanardini (Flora 34: 34, 1851).
26
Type from the Red Sea (As Suways, Khalij Umm el Ketef [Gulf of Berenice], Al
Hudaydah, Al Mukha), leg. Portier, in herb. Zanardini, Museo Civico di Storia Naturale, Venezia (see De Toni & Levi 1888).
tunnel in coral rock, ca. 50 cm below low water (ISRSE 65/0382, 3126-tetrasporic).
Museri Island: in front of Scopus Ridge, 9.x.1965, in Chondrophycus papillosus
community, on rock flat ca. 20 cm below low water (ISRSE 65/0388, 2347k),
10.x.1965, on rock flat ca. 30 cm below low water (ISRSE 65/2315g-tetrasporic),
Camping Bay, 19.x.1965, on rock flat ca. 30 cm below low water (ISRSE 65/2216,
3080-tetrasporic). Romia Island: 29.iii.1962, (E62/20196-tetrasporic). Harmil Island: 28.iii.1962 (E62/20249).
DISTRIBUTION: Red Sea.
COMMENTS : Our plants were to 8 cm tall in the open habitats, but considerably smaller
in the shade. Tetrasporic plants were obtained both in spring and autumn, cystocarpic
plants were not found. Champia irregularis grew epiphytically on Halimeda discoidea,
Caulerpa racemosa, Chondrophycus papillosus, Hypnea cornuta, Cystoseira myrica,
Digenea simplex and other algae in flat, rocky habitats in shallow water, as well as
epilithically, usually in the shade.
*Champia parvula (C. Agardh) Harvey (Nereis Bor. Amer. 2, p. 76, 1853).
Chondria parvula C. Agardh (Syst. Alg., p. 207, 1824).
Type from Cadiz, Spain, in herb. Agardh, LD 26022 (see Reedman & Womersley
1976).
ISRSE
SPECIMENS STUDIED:
Entedebir Island: Mersat Abiad, 26.iii.1962 (E62/20162).
COMMENTS : A few individuals of this alga were collected in the spring of 1962. The
plant was later found on the Saudi Arabian Red Sea coast at an unspecified locality
(Aleem 1981).
CERAMIALES
CERAMIACEAE
ANOTRICHIUM Nägeli
Anotrichium tenue (C. Agardh) Nägeli (Sitzungsber. Bayer. Akad. Wiss. München
1861(2): 399, 1861).
Griffithsia tenuis C. Agardh (Sp. Alg. 2(1), p. 131, 1828).
Type from Venezia (Venice), Adriatic Sea, leg. Ruchinger, in herb. Agardh, LD
19891 (see Baldock 1976, p. 557).
27
Chondria dasyphylla in Caulerpa racemosa community on a shallow sandy bottom
(ISRSE 65/2341f), in Chondrophycus papillosus community, on a shallow rock flat
gently sloping seawards (ISRSE 65/2355), 10.x.1965, on Halimeda discoidea growing
on a rock flat gently sloping seawards (ISRSE 65/2315k); Camping Bay, 19.x.1965,
on Thalassia hemprichii growing in sand-filled crevices in rock flat at ca. 30 cm
below low water (ISRSE 65/2215).
COMMENTS: This epiphyte was common in various shallow-water habitats of Museri
Island. Our plants were well developed, forming clumps to 6 cm in diameter. No
fertile plants were observed. The species, as Griffithsia tenuis, has hitherto been
recorded in the Red Sea only from its northern part.
CENTROCERAS Kützing
Centroceras clavulatum (C. Agardh) Montagne (Phyceae, in Durieu de Maisonneuve, Expl. Sci. Algérie, Bot., Cryptog., p. 140, 1846).
Ceramium clavulatum C. Agardh (in Kunth, Syn. Pl. 1, p. 2, 1822).
Type from Callao de Lima, Peru, in herb. Agardh, LD (see Womersley & Bailey
1970).
ISRSE SPECIMENS STUDIED: Cundabilu Island: western side of islet, 20.iii.1962, in a
Jania cushion (E62/20101). Museri Island: in front of Scopus Ridge, 9.x.1965, on
Hypnea cornuta among many other epiphytes, the most common of which were
Polysiphonia mollis and Cladophoropsis membranacea, in Chondrophycus papillosus
community, on shallow rock flat gently sloping seawards (ISRSE 65/2337h).
COMMENTS: The plants from Cundabilu had long, well-developed branches. They
formed a sparse growth in the shelter of the thicket of Jania, rather than the massive
cushion of pure Centroceras, as is so often the situation in the Mediterranean or
other regions. The plants from Museri were very small with only two branches
being found, one 5 mm long, the other 3 mm long (see also Lipkin 1972a, p. 428).
The scarcity and small size in the Red Sea of this common species may be attributable
to heavy grazing pressure (Lipkin 1977a). Jania may have provided a shelter for
Centroceras in this case, as the herbivorous fishes seem to avoid calcareous algae
(Lundberg & Lipkin 1979).
CERAMIUM Roth (nom. cons.)
*Ceramium codii (Richards) Mazoyer (Bull. Soc. Hist. Nat. Afrique du Nord 29:
324. 1938).
28
Ceramothamnion codii Richards (Bull. Torrey Bot. Club 27: 264, pls. 21, 22. 1901).
Type from Bermuda.
ISRSE SPECIMENS STUDIED : Museri Island: in front of Scopus Ridge, 10.x.1965 (ISRSE
65/2379e-tetrasporic).
DISTRIBUTION: Widespread in warm waters of the Atlantic, Indian, and Pacific oceans.
COMMENTS : A few plants of this alga were found creeping on a leaf of Thalassia
hemprichii. They were at the end of their tetraspore-shedding season (autumn).
Ceramium diaphanum (Lightfoot) Roth (Catal. Bot. 3, p. 154, 1806).
Conferva diaphana Lightfoot (Fl. Scot., p. 996, 1777).
Type from Scotland.
ISRSE SPECIMENS
sterile).
STUDIED :
Entedebir Island: Landing Bay, 13.iii.1962 (E62/20029-
DISTRIBUTION: Widespread in warm and warm temperate seas.
Ceramium flaccidum (Kützing) Ardissone (Nuov. Giorn. Bot. Ital. 3: 40, 1871).
Hormoceras flaccidum Kützing (Tab. Phycol. 12, p. 21, pl. 69, figs. a-d, 1862).
Type from Kilkee, Co. Clare, Ireland, no. 940,265 ... 55 in L (see Womersley 1978,
p. 236).
ISRSE SPECIMENS STUDIED: Cundabilu Island: western side of islet, 20.iii.1962 (E62/
20075). Um Aabak: 6.iv.1962, 50 cm depth, on Laurencia and a hydroid, the
former growing on the latter (E62/20259). Museri Island: in front of Scopus
Ridge, 8.x.1965, on Caulerpa sertularioides (ISRSE 65/2330a -tetrasporic,
cystocarpic), 9.x.1965, on Chondrophyc us papillosus community (ISRSE 65/
2342a-tetrasporic, cystocarpic), in Caulerpa racemosa community (ISRSE 65/
2351j, 2356-cystocarpic), 19.x.1965 (ISRSE 65/2203-cystocarpic, 3080c tetrasporic, cystocarpic, 3084e-cystocarpic), 16.x.1965 (ISRSE 65/3054). Romia
Island: 29.iii.1962, on Sargassum (E62/20197).
DISTRIBUTION: Widespread in tropical to cold temperate waters.
COMMENTS : This alga was one of the common epiphytes in various habitats in shallow
waters in autumn. Tetrasporophytes and gametophytes were common in autumn but
absent in spring.
Two of our specimens (E62/20075 and E62/20259) resembled Ceramium gracillimum
var. byssoideum Mazoyer, whereas the other specimens resembled C. taylorii Dawson.
Both taxa were considered conspecific with C. flaccidum by Womersley (1978).
This difference in morphology may be seasonal.
29
Ceramium rubrum auctorum
65/2330g-tetrasporic, sparsely on Caulerpa sertularioides).
COMMENTS: The circumscription and correct name of this species, which is commonly
reported throughout the world as Ceramium rubrum (Hudson) C. Agardh, are uncertain
(see Silva et al. 1996, p. 402).
SPYRIDIA Harvey
Spyridia filamentosa (Wulfen) Harvey (in Hooker, Brit. Fl. 2(1) [= J.E. Smith,
Engl. Fl. 5(1)], p. 337, 1833).
Fucus filamentosus Wulfen (Crypt. Aquat., p. 64, 1803).
Type from the Adriatic Sea.
ISRSE SPECIMENS STUDIED : Entedebir Island: Goliath Bay, 3.iv.1962 (E62/20214);
Padina Bay, 12.iii.1962 (E62/20289).
DISTRIBUTION : Widespread in warm and temperate seas.
COMMENTS : Our plants were 10-12 cm tall, sterile, and yellowish in their upper
parts, which were exposed to direct sunlight. It is of interest to note that Spyridia
filamentosa dominates the epiphytic algal community of the lowest parts of Avicennia
marina pneumatophores on Sinai coasts, the northern Red Sea (Dor, 1984; Lipkin,
pers. obs.), but was absent in the mangroves of Dahlak Archipelago visited by the
two expeditions. The lower parts of A. marina pneumatophores in this region were
devoid of macroscopic algal growth, probably because of the lack of wave action.
DELESSERIACEAE
*CALOGLOSSA (Harvey) G. Martens
*Caloglossa leprieurii (Montagne) G. Martens (Flora 52: 234, 237, 1869).
Delesseria leprieurii Montagne (Ann. Sci. Nat., Bot., ser. 2, 13: 196, pl. 5, fig. 1, 1840).
Type from Sinnamary, NW of Cayenne, French Guiana, leg. Leprieur, presumbaly
in PC, isotype: LD 31909-12 (see Post, 1936, p. 52).
ISRSE SPECIMENS STUDIED : Melita Bay in the Gulf of Zula: 2.iv.1962, on Avicennia
marina pneumatophores (E62/20232).
DISTRIBUTION : Widespread in warm and temperate seas, penetrating temperate waters
up to the latitudes of 40-46°.
COMMENTS: Our plants were rather small and had segments 1-1.5 mm wide and 35 mm long. The shape of the segments and the ratio of their length to width
30
corresponded to C. leprieurii var. hookeri Post (1936, p. 53). King & Puttock (1994)
treated C. leprieurii as a morphologically variable species with none of the infraspecific
taxa recognized so far appearing to be more than a range on a continuum of variation.
Kamiya et al. (1993), however, found that var. hookeri was morphologically distinct
and incapable of crossing with other varieties. The formae recognized by Post (1936)
are growth forms or ecophenes.
Of all the mangals that were visited by the two expeditions, only that of Melita Bay
in the Gulf of Zula on the Eritrean coast supported growth of members of the
Bostrychietum. Our plants grew on the upper parts of the pneumatophores of Avicennia
marina, along with barnacles, in the midlittoral zone. On the islands of the Dahlak
Archipelago, the pneumatophores and trunks of mangroves at the same intertidal
level supported other algae, mostly blue-greens. This difference in vegetation may
be attributable to a difference in wave exposure, Melita Bay being highly sheltered.
Brackish water, with which Caloglossa is often associated, is absent both in the Gulf
of Zula and on the islands.
MARTENSIA Hering (nom. cons.)
Martensia elegans Hering (Ann. Mag. Nat. Hist. 8: 92, 1841).
Type from Port Natal, South Africa, leg. Krauss (female plant on mica, lectotypified
by Papenfuss), in HBG.
ISRSE SPECIMENS STUDIED : Entedebir Island: outside Landing Bay to the north,
13.iii.1962, 3 m depth (E62/20031, 20034); Landing Bay, 11.iii.1962 (E62/20001tetrasporic), 21.x.1965, in a shaded cavelet in coral rock (ISRSE 65/0379, 0380,
3131, 3133a); northern shore of Ras Papenfuss, 25.iii.1962 (E62/20148). Museri
Island: in front of Scopus Ridge, 10.x.1965, on Avrainvillea amadelpha (ISRSE 65/
2320h).
COMMENTS : Our plants were 2-3 cm tall, with reticulate parts up to 1.5 cm wide.
Most of them had coherent tissue at the base and one zone of reticulate tissue, with
a narrow coherent, entire margin. Some plants, however, that grew intermingled
with typical M. elegans, had two or three alternating zones of coherent and reticulate
tissue and a denticulate margin. These features are characteristic of Martensia
denticulata Harvey (1855, p. 537) from Western Australia, a species considered
conspecific with M. fragilis Harvey (1854, p. 145) by Millar (1990, p. 418).
According to Svedelius (1908, p. 31), these features may occasionally be observed
in M. elegans.
Plants collected in spring were larger than those obtained in autumn. A few bore
tetrasporangia, whereas all autumn plants were sterile.
This alga prefers habitats well protected from direct sunlight. It grew either in cavelets
or under larger algae on rock flats.
31
DASYACEAE
DASYA C. Agardh (nom. cons.)
*Dasya baillouviana (S.G. Gmelin) Montagne (in Barker-Webb & Berthelot, Hist.
Nat. Iles Canaries 3, p. 165, 1841).
Fucus baillouviana S.G. Gmelin (Hist. Fuc., p. 165, 1768).
Type from the Mediterranean Sea; pl. II in Griselini, Observ. Scolopendre, 1750,
designated lectotype by Dixon & Irvine (1970).
ISRSE SPECIMENS STUDIED : Entedebir Island: Landing Bay, 21.x.1965, in a shaded
cavelet in coral rock, ca. 1 m depth (ISRSE 65/3136a-tetrasporic).
DISTRIBUTION : Widespread in both temperate and tropical waters.
C OMMENTS : Our plants were small (about 5 cm tall), with rather thick axes.
Tetrasporangia were present only in the uppermost parts of the pedicellate stichidia,
their lower locules being already empty.
The alga, for a long time known as D. pedicellata (C. Agardh) C. Agardh, was
been reported from the Red Sea until rather recently (Lipkin 1979, 1987), but it
is not uncommon on the Sinai coasts, where it was obtained with cystocarps in
January (Lipkin 1979 and unpublished data); summer collections were always
sterile.
HETEROSIPHONIA Montagne (nom. cons.)
Heterosiphonia crispella (C. Agardh) Wynne (Cryptog. Algol. 6: 87, 1985).
Callithamnion crispellum C. Agardh (Sp. Alg. 2, p. 183, 1828).
Type from Cádiz, Atlantic coast of Spain, leg. A.N. Cabrera, holotype in LD (44225)
(see Wynne 1985, p. 86).
ISRSE SPECIMENS STUDIED : Entedebir Island: Landing Bay, 21.x.1965, in a shaded
cavelet in coral rock, ca. 50 cm depth, on Chondrophycus parvipapillatus (ISRSE
65/2359a-tetrasporic).
COMMENTS: Our plants were only a few mm tall, yet fertile. This alga was reported
from the northern Red Sea (Nasr 1947 – as H. wurdemannii (Bailey ex Harvey)
Falkenberg), where it is common on coral reefs, as well as in the epiphytic algal
community on seagrass leaves (Lipkin, pers. obs.). It has not been previously reported
from the southern Red Sea.
Our plants were referable to H. crispella var. laxa (Børgesen) Wynne (H. wurdemannii
var. laxa Børgesen, Dansk Bot. Ark. 3: 326, figs. 327, 328, 1919), which appears to
be merely a shaded-habitat ecophene (see Børgesen 1919; Nasr 1947).
32
RHODOMELACEAE (nom. cons.)
ACANTHOPHORA Lamouroux
Acanthophora nayadiformis (Delile) Papenfuss (Israel J. Bot. 17: 96, 1968).
Fucus nayadiformis Delile (Fl. Égypte, Explic. Pl., p. 292, pl. 56, fig. 1, 1813).
Type from Egypt (Al-Iskandariyah [Alexandria]; As-Suways).
ISRSE SPECIMENS STUDIED: Entedebir Island: Goliath Bay, 14.iii.1962, 1 m depth, on
Turbinaria (E62/20042); Mersat Abiad, 26.iii.1962 (E62/20158). Museri Island: in
front of Scopus Ridge, 9.x.1965, in Chondrophycus papillosus community, on shallow
rock flat gently sloping seawards (ISRSE 65/2351).
DISTRIBUTION: Warm waters of the Indian Ocean, Red Sea and Mediterranean Sea.
COMMENTS: This alga is not very common in the southern Red Sea. The plants obtained
in spring were considerably larger, about 16 cm tall, than those collected in autumn,
which measured only 8 cm in height. All of our plants were sterile.
*BOSTRYCHIA Montagne (nom. cons.)
*Bostrychia simpliciuscula Harvey ex J. Agardh (Sp. Alg. 2(3), p. 854, 1863).
Type from Tonga, leg. W.H. Harvey.
ISRSE SPECIMENS STUDIED: Melita Bay, Gulf of Zula: 2.iv.1962, on pneumatophore
of Avicennia marina, along with Caloglossa leprieurii, Calothri x pilosa and
Brachytrichia lloydii (E62/20232a-tetrasporic).
DISTRIBUTION: Widespread in warm seas and estuaries, on mangrove pneumatophores
and trunks, and on rocks.
COMMENTS : Our plants formed a tuft ca. 1 mm thick around the upper part of a pneumatophore over a distance of 3 cm. A few individuals of Caloglossa leprieurii grew
on the tuft, which was partially covered by a mat of Calothrix pilosa 1.5 mm thick
in which mud particles were trapped. Dark patches of Brachytrichia lloydii occurred
on the surface of the pneumatophore immediately below the tuft of Bostrychia.
The creeping lower branches of our plants were ecorticate, with 5 or 6 pericentral
cells half the length of the axial cells. Plants were attached to the pneumatophores by
short holdfasts arising at branching points, usually perpendicular to the plane of the
branches. Erect lateral branches had 4 pericentral cells 85±16 µm in diameter, with
distichous monosiphonous branchlets. The latter usually branched once, sometimes
twice, and were 1.7±0.6 µm long and 41±3.5 µm diameter near the base of the
lateral and 25.5±3.1 µm diameter near its apex. Stichidia were elongate, tapering at
both ends, with a branched continuation above the fertile part. Tetrasporangia were
3 per segment in the center of the stichidium, 1 or 2 near each end.
33
CHONDRIA C. Agardh (nom. cons.)
*Chondria baileyana (Montagne) Harvey (Nereis Bor. Amer. 2, p. 20, pl. 18, fig.
A 1-6, 1853).
Laurencia baileyana Montagne (in Bailey, Amer. J. Sci., ser. 2, 6: 38, 1848).
Type from the North Atlantic coast of the U.S.A. (Newport, Rhode Island; Stonington,
Connecticut; Fort Hamilton, New York), leg. J.W. Bailey, presumably in PC.
ISRSE SPECIMENS STUDIED : Museri Island: in front of Scopus Ridge, 8.x.1965, epilithic
and on many algae growing on a rock flat gently sloping seawards, at ca. 30 cm
below low water (ISRSE 65/0301b-tetrasporic, 2325c), 10.x.1965 (ISRSE 65/2207,
2383a-tetrasporic); Camping Bay, in front of Ras Pandion, 16.x.1965 (ISRSE 65/
3065a-cystocarpic), 19.x.1965 (ISRSE 65/3082a-tetrasporic, male and female
gametophytes).
DISTRIBUTION : Widespread in warm and temperate Atlantic, Mediterranean, and Red
Sea waters.
COMMENTS: Our plants were to 6 cm tall, fertile during autumn.
Chondria collinsiana Howe (in Britton & Millspaugh, Bahama Fl., p. 568, 1920).
Type from Mayaguana, Bahama Islands.
ISRSE SPECIMENS STUDIED : Museri Island: off Mangrove Bay, 14.x.1965 (ISRSE 65/
3039-tetrasporic); Camping Bay, in front of Ras Pandion, 16.x.1965 (ISRSE 65/
3056-tetrasporic).
DISTRIBUTION : Widespread in warm and temperate Atlantic, Mediterranean, Red Sea,
and Indian Ocean waters.
COMMENTS: Our plants were to 5 cm tall and formed tufts on rocky substrates, mostly
coral knolls in shallow water. Practically all individuals collected bore tetrasporangia.
This species has been reported from the northern Red Sea (Nasr 1947). The reports
of the results of the ISRSE are the first records of its occurrence in the southern part.
*Chondria dasyphylla (Woodward) C. Agardh (Syn. Alg. Scand., p. XVIII, 1817).
Fucus dasyphyllus Woodward (Trans. Linn. Soc. London, 2: 239, pl. 23, figs. 1-3, 1794).
Type from Norfolk, England (Cromer; Yarmouth) (see discussion on typification in
Gordon-Mills 1987, p. 246).
ISRSE SPECIMENS STUDIED : Entedebir Island: Landing Bay, 21.x.1965 (ISRSE 65/
3131a). Museri Island: in front of Scopus Ridge, 9.x.1965, the whole population
infected by a parasite (ISRSE 65/2354-tetrasporic); off Mangrove Bay, 14.x.1965
(ISRSE 65/3038-tetrasporic); Camping Bay, in front of Ras Pandion, 16.x.1965
(ISRSE 65/3069-tetrasporic).
34
DISTRIBUTION: Widespread in warm and warm-temperate seas.
COMMENTS : Our plants were to 6 cm tall and very common in shallow waters. This
species was not encountered during spring and might occur only in summer and
autumn.
CHONDROPHYCUS (Tokida et Saito) Garbary et Harper
*Chondrophycus glanduliferus (Kützing) comb. nov.
Chondria glandulifera Kützing (Phycol. Germ., p. 855, 1845).
Type from Trieste, Adriatic Sea.
ISRSE
SPECIMENS STUDIED:
DISTRIBUTION: Mediterranean Sea, warm Indo-west Pacific waters.
COMMENTS : Our plants were 8 cm tall and sterile.
The discovery of this species in the Red Sea fills a gap in its hitherto disjunct
distributional pattern.
Chondrophycus papillosus (C. Agardh) Garbary et Harper (Cryptog. Algol. 19:
195, 1998).
Chondria papillosa C. Agardh (Sp. Alg. 1(2), p. 344, 1822).
Fucus papillosus Forsskål (Fl. Aegypt.-Arab., p. 190, 1775, nom. illeg. (not F. papillosus S.G.
Gmelin, Hist. Fuc., p. 188, 1768).
Type from Al Mukha, Yemen, southern Red Sea, leg. P. Forsskål, in C (see Børgesen
1932, p. 6; Yamada 1931, p. 190, pl. 1a).
ISRSE SPECIMENS STUDIED: Entedebir Island: Padina Bay, 12.iii.1962 (E62/20014);
Goliath Bay, 3.iv.1962, (E62/20212); Landing Bay, 12.iii.1962, 2 m depth (E62/
20028), 21.x.1965 (ISRSE 65/3112-tetrasporic, 3124, 3147). Museri Island: Camping Bay, 19.x.1965 (ISRSE 65/2205, 3082); Camping Bay, off Ras Pandion
16.x.1965 , (ISRSE 65/3071-tetrasporic); in front of Scopus Ridge, 9.x.1965,
dominating the algal community on rock flat ca. 30 cm deep during low water
(ISRSE 65/2337f, 2350), 10.x.196 5, with many epiphytes (ISRSE 65/2370tetrasporic); off Mangrove Bay, 14.x.1965, with some epiphytic Cladophora and
Ceramium (ISRSE 65/3032). Romia Island: 29.iii.1962 (E62/20285). Harmil Island: 28.iii.1962 (E62/20251).
DISTRIBUTION: Widespread in warm and temperate seas.
C OMMENTS : Our plants reached 10 cm in height, but were usually half that size,
brownish yellow, often bearing many epiphytes, and often heavily silted. They bore
tetrasporangia during autumn.
This is one of the most common algae in the entire Red Sea. Around Museri Island
it was the dominant alga in an important plant community, occupying wide areas on
35
rock flats surrounding the island, in the uppermost parts of the subtidal and the
infralittoral fringe of the intertidal zone (Lipkin 1987). It penetrated the lower
midlittoral zone, where it became exposed during low water of every spring tide. In
these relatively high levels the plants were bleached, while in the lower levels of the
rock flat they were yellowish, becoming brownish at depths of 50-60 cm below the
lowest tidal level.
All previous records of this species from the Red Sea were under the name Laurencia
papillosa (C. Agardh) Greville. For a discussion of the nomenclature of this species
see Papenfuss (1968a, p. 99).
*Chondrophycus parvipapillatus (Tseng) Garbary et Harper (Cryptog. Algol. 19:
195, 1998).
Laurencia parvipapillata Tseng (Pap. Michigan Acad. Sci. 28: 204, pl. IV, 1943).
Type from Cape d’Aguilar, Hong Kong, leg. Tseng 2813.
cavelet in coral rock ca. 50 cm depth (ISRSE 65/2359). Museri Island: Camping
Bay, off Ras Pandion, 16.x.1965, (ISRSE 65/0330, 3052); off Mangrove Bay,
14.x.1965 (ISRSE 65/3046, 3047-tetrasporic).
DISTRIBUTION : Widespread in warm Indo-west Pacific waters and the Red Sea.
COMMENTS: This is an easily recognizable, small Chondrophycus . Our plants were
spreading, 2-4 cm in diameter, with flattened branches bearing papillae distichously
on both margins of the branches. Occasionally a few papillae were also encountered
on the flat surfaces, especially near the base of the plant. Peripheral cortical cells in
the distal parts of branches project above the branch surface. Tetrasporophytes were
found in autumn.
The alga grew in habitats protected from direct sunlight, in shallow cavelets or
underneath larger algae on rock flats, but not in heavily shaded habitats; it was dark
red.
DIGENEA C. Agardh
Digenea simplex (Wulfen) C. Agardh (Sp. Alg. 1(2), p. 389, 1822).
Conferva simplex Wulfen (Crypt. Aquat., p. 17, 1803).
Type from Trieste, Adriatic Sea.
ISRSE SPECIMENS STUDIED : Entedebir Island: Goliath Bay, 6.iv.1962 (E62/20256).
community, on rock flat gently sloping seawards (ISRSE 65/2354l), 10.x.1965 (ISRSE
65/2315m). Romia Island: 29.iii.1962 (E62/20199-tetrasporic, cystocarpic and males).
36
COMMENTS : Plants collected in spring were large (to 15 cm tall), well developed and
bushy, whereas those obtained in autumn were small (not more than 2.5 cm tall),
almost unbranched and appeared to be in early stages of development. Plants collected
at Romia Island in spring were fertile, whereas those collected on Entedebir Island,
only a week later, were sterile. This difference may be another expression of the
special ecological conditions on the coasts of Romia Island.
HERPOSIPHONIA Nägeli
Herposiphonia secunda (C. Agardh) Ambronn f. tenella (C. Agardh) Wynne
(Cryptog. Algol. 5: 173, 1985).
Hutchinsia tenella C. Agardh (Sp. Alg. 2(1), p. 105, 1828).
Herposiphonia tenella (C. Agardh) Ambronn (Bot. Zeitung 38: 197, 1880).
The conspecificity of Herposiphonia tenella and H. secunda was proposed by Hollenberg (1968b).
Type from Sicilia (Sicily), Mediterranean Sea.
ISRSE SPECIMENS STUDIED : Museri Island: in front of Scopus Ridge, 9.x.1965, in
Caulerpa racemosa community on Thalassia hemprichii (ISRSE 65/2342h), in
Chondrophycus papillosus community, on Halimeda discoidea, Avrainvillea amadelpha, Caulerpa mexicana, Hypnea cornuta, Caulerpa racemosa, Chondrophycus
papillosus, Acanthophora nayadiformis, Chondria dasyphylla and Hypnea anastomosans (ISRSE 65/2315h, 2343a, 2344g, 2346d, 2347l, 2350g, 2351e, 2353v, 2354m,
2361), 10.x.1965 in Caulerpa racemosa community, on Padina boergesenii, Cystoseira
myrica, Cladophora sp. (ISRSE 65/2366j, 2368p, 2380h); off Mangrove Bay,
14.x.1965, on Halimeda discoidea, Gelidium sp. (ISRSE 65/2318f, 3004e).
COMMENTS : Plants of this common epiphyte were sterile in autumn. They grew on a
variety of host algae in shallow water. Their creeping axes had branchlets of limited
growth on almost every node, and branches of indefinite growth at irregular intervals
of 1-6 nodes, usually 3; certain nodes had neither. The main, creeping axes were
105-130 µm in diameter and the erect branchlets were 50-65 µm in diameter; both
had 7-9 pericentral cells surrounding the central axial cell. Branchlets were 3-5
times dichotomously branched and carried 1-5 trichoblasts at the tips. Rhizoids
emerged from almost every segment, most of them were unbranched, a few ending
with an irregular disc.
LAURENCIA Lamouroux (nom. cons.)
Laurencia obtusa (Hudson) Lamouroux (Ann. Mus. Hist. Nat. [Paris] 20: 130,
1813).
Fucus obtusus Hudson (Fl. Angl., ed. 2, p. 586, 1778).
37
Type from southern England (Hastings, Sussex; Devon) (see Maggs & Hommersand
1993, p. 398).
20076-tetrasporic).
COMMENTS: Our plants were to 12 cm tall and sterile.
Laurencia uvifera (Forsskål) Børgesen (Dansk Bot. Ark. 8(2): 12, figs. 3, 4, pl. 1,
fig. 6, 1932.
Fucus uvifer Forsskål (Fl. Aegypt.-Arab., p. 192, 1775).
Type from the harbor of Constantinople, Turkey, leg. P. Forsskål, in C (Børgesen
1932, p. 12, figs. 3, 4, pl. I, fig. 6).
ISRSE
SPECIMENS STUDIED :
Harmil Island: 28.iii.1962 (E62/20246-tetrasporic).
DISTRIBUTION : Mediterranean Sea, Red Sea.
COMMENTS: Our plants were to 10 cm tall. Longitudinal secondary pit connections
were present between cells of the cortex. Lenticular thickenings were absent in the
walls of the medullary cells, and the cortical cells were not palisade-like, both
characters agreeing with observations on the type specimen made by Børgesen (1932).
Our material also agrees with the illustrations and dimensions given by that author.
*Laurencia venusta Yamada (Univ. Calif. Publ. Bot. 16: 203, fig. H, pl. 6, fig. a,
1931).
Type from Kyushu, Japan (Koshiki-jima, Satsuma Province; Goto, Hizen Province).
ISRSE SPECIMENS STUDIED : Entedebir Island: Goliath Bay, 3.iv.1962 (E62/20215);
Landing Bay, 21.x.1965, on Chondrophycus papillosus (ISRSE 65/3124a). Museri
Island: in front of Scopus Ridge, 10.x.1965 (ISRSE 65/2369a).
DISTRIBUTION : Japan, Korea, Taiwan, Australia, Somalia, Tanzania, Red Sea; probably
widespread in warm Indo-west Pacific waters.
COMMENTS: Our plants were about 5 cm tall, sterile during autumn. In cross section
the walls of the medullary cells showed annular rather than lenticular thickenings.
LEVEILLEA Decaisne
Leveillea jungermannioides (Hering et Martens) Harvey (Trans. Roy. Irish Acad.
22(Sci.): 539, 1855).
Amansia jungermannioides Hering et Martens (Flora 19: 485, figs. 1-4, 1836).
Type from Et Tur, Sinai Peninsula, the northern Red Sea, leg. W. Schimper,
27.iii.1835 (Unio Itiner. no. 472).
38
ISRSE SPECIMENS STUDIED : Entedebir Island: Goliath Bay, 22.iii.1962, on Sarconema
filiforme (E62/20109-tetrasporic and cystocarpic); Padina Bay, 12.iii.1962, on
Chondrophycus papillosus (E62/20268). Romia Island: 29.iii.1962, on Sargassum
(E62/20190). All three samples were cited by Scagel & Chihara (1968).
DISTRIBUTION: Warm waters of the Indo-Pacific region.
COMMENTS: This species is a rather common epiphyte on attached Sargassum and
other Fucales in the Red Sea, but not found on the same species in the floating
masses they often form, except when these masses are composed of plants that have
been recently detached. The length of time a host plant had been floating could be
estimated roughly by the degree of development of the typical animal associations
of floating objects, like goose barnacles or certain bryozoans.
LOPHOCLADIA (J. Agardh) Schmitz
Lophocladia lallemandii (Montagne) Schmitz (Ber. Deutsch. Bot. Ges. 11: 223.
1893).
Dasya lallemandii Montagne (Ann. Sci. Nat., Bot., ser. 3, 12: 289. 1849).
This species, which was originally described from the Red Sea, was reported from
the Dahlak Archipelago by Mazza (1910: 4) but was not encountered by either of the
two expeditions.
POLYSIPHONIA Greville (nom. cons.)
*Polysiphonia mollis J.D. Hooker et Harvey (in Harvey, Nereis Austral., p. 43,
1847).
Type from Georgetown, Tasmania, leg. R. C. Gunn, lectotype (“Gunn 1316”) in BM
(ex K), isotypes in BM (ex K), TCD (see Womersley 1979, p. 477).
ISRSE SPECIMENS STUDIED : Entedebir Island: Landing Bay, 21.x.1965, tetrasporic
(ISRSE 65/3136f); Mersat Abiad, 26.iii.1962 (E62/20175). Museri Island: in front
of Scopus Ridge, 8.x.1965, tetrasporic (ISRSE 65/2325a, 2330b, 2331d, 2333d),
9.x.1965 , tetrasporic, male and female (ISRSE 65/2341e, 2357), 10.x.196 5,
tetrasporic, male and female (ISRSE 65/2315g); off Mangrove Bay, 14.x.1965,
tetrasporic, male and female (ISRSE 65/2318e, 3024a, 3030); Camping Bay, off
Ras Pandion, 16.x.1965, tetrasporic (ISRSE 65/3054e, 3067).
DISTRIBUTION: Widespread in warm and warm-temperate waters.
C OMMENTS : This small and delicate Polysiphonia was one of the most common
epiphytes in autumn. It grew on many algal species in all photophilic communities
in shallow waters. The majority of the plants in every population were reproductive
in autumn and a number of sporelings were observed on various hosts. It is assumed,
therefore, that October is the peak of the reproductive season.
39
Womersley (1979) doubted that P. mollis occurs outside southern Australia. The Red
Sea plants belong to P. mollis as accepted by Weber-van Bosse (1923), Børgesen
(1945), Hollenberg (1961, 1968a), and Islam (1976).
PHAEOPHYTA
ECTOCARPALES
ECTOCARPACEAE
FELDMANNIA G. Hamel
Feldmannia indica (Sonder) Womersley et Bailey (Philos. Trans., Ser. B., 259:
288, 1970).
Ectocarpus indicus Sonder (in Zollinger, Syst. Verz., pp. 2, 3 footnote, 1854).
Type from Pulo Kambing, Bima Bay, Sumbawa (Bima Island), Indonesia, on
calcareous rocks, leg. Zollinger (no. 3428), in MEL (26389) (see Womersley &
Bailey 1970).
ISRSE SPECIMENS STUDIED : Entedebir Island: Landing Bay, 14.iii.1962, on drifting
Turbinaria (E62/20040c).
DISTRIBUTION : Widespread in all warm seas.
COMMENTS : Our plants bore oblong, apically rounded unilocular and plurilocular
sporangia 42-47 µm long and 19-24 µm wide. They grew epiphytically on drifting
Turbinaria side by side with Sphacelaria and other filamentous algae, but were not
observed on attached Turbinaria.
Previous Red Sea records of this species are all from the north (as Ectocarpus duchassaingianus Grunow by Nasr 1947; as Giffordia indica (Sonder) Papenfuss et Chihara
ex Papenfuss by Lipkin 1972a, 1972b and Aleem 1978a, 1978c, 1984).
Feldmannia irregularis (Kützing) G. Hamel (Phéophyc. France, p. XVII, 1939).
Ectocarpus irregularis Kützing (Phycol. Germ., p. 234, 1845).
Type from the Adriatic Sea, on Laurencia obtusa.
Turbinaria (E62/20040).
COMMENTS: Our plants had plurilocular reproductive organs 57-65 µm long and 2627 µm wide at the widest part. They were all sessile, in agreement with the description
of the species given by Hamel (1931, p. 47). For the genus, however, Hamel (1939,
p. 67) noted that plurilocular organs are generally pedicellate.
40
KUETZINGIELLA Kornmann
Kuetzingiella elachistaeformis (Heydrich) Balakrishnan et Kinkar (Seaweed Res.
Utilis. 4(2): 25, figs 20, 98-100, 1981).
Ectocarpus elachistaeformis Heydrich (Ber. Deutsch. Bot. Ges. 10: 470, pl. XXV, fig. 14, 1892).
Type from Madang near Bogia (Hatzfeldthafen), Papua-New Guinea, on Sargassum
cristaefolium var. upolense Grunow.
Turbinaria (E62/20040b).
COMMENTS : Our plants were ca. 3 mm tall, with filaments 13-18 µm in diameter and
bearing plurilocular reproductive organs 85-115 µm long and 20-25 µm wide that
were usually sessile, rarely pedicellate.
This species is probably widespread in the Red Sea but has been previously reported,
as Ectocarpus elachistaeformis, only from the northern part (Nasr 1947; Rayss &
Dor 1963; Aleem 1978a).
SPHACELARIALES
SPHACELARIACEAE
SPHACELARIA Lyngbye
*Sphacelaria brachygonia Montagne (Ann. Sci. Nat., Bot., ser. 2, 20: 305, 1843).
Type from Ilha Santa Catarina, Brazil, leg. Gaudichaud, lectotype in herb. Montagne,
PC (see Sauvageau 1901, p. 134), isotypes in L, PC, TCD (see Prud’homme van
Reine 1982, p. 188).
ISRSE SPECIMENS STUDIED: Museri Island: in front of Scopus Ridge, 8.x.1965, among
other epiphytes on Turbinaria triquetra (ISRSE 65/2333e).
DISTRIBUTION: Brazil, Ghana, Senegal, South Africa (Natal), Red Sea, Mediterranean, Atlantic coast of Europe and north Africa, southern Australia; possibly widespread
in warm and temperate seas.
COMMENTS : Our material was devoid of propagules, but bore numerous plurilocular
reproductive organs that measured 60 µm by 45 µm and were nearly rectangular in
side view, with rounded corners. Filaments were 55 µm in diameter, the length of
segments 18-34 µm, with 3-5 longitudinal divisions in a segment.
Sphacelaria rigidula Kützing (Phycol. General., p. 292, 1843).
Type from Nuweiba, Sinai, Red Sea, on Hormophysa cuneiformis (as Cystoseira triquetra), leg. W. Schimper, holotype in L (no. 937.117-229) (see Prud’homme van
Reine 1982, p. 203).
41
Turbinaria (E62/20040d-with plurilocular reproductive organs); Mersat Abiad,
26.iii.1962, on Sargassum (E62/20171); Goliath Bay, 3.iv.1962, on Turbinaria stalks
(E62/20208). Desta Island: rocks off the northern coast, 26.x.1965, lithophytic
(ISRSE 65/3169h). Museri Island: in front of Scopus Ridge, 8.x.1965, among other
epiphytes on Turbinar ia triquetra (ISRSE 65/2333i-with many plurilocular
reproductive organs), 10.x.1965, on Padina boergesenii (ISRSE 65/2367g).
COMMENTS: Our plants were 3-5 mm tall with a main axis 18.5-21 µm in diameter.
They usually grew on Sargassum and Turbinaria, forming dense velvety growths on
the stalks, at times intermingled with, but less abundant than, S. tribuloides. Propagules
were often present but not abundant in spring and absent in autumn.
Sphacelaria tribuloides Meneghini (Lett. Corinaldi, p. 2, 1840).
Type from La Spezia, Italy, on submerged rocks, leg. Meneghini, lectotype in L
(no. 937.71-699), isotypes in LD, PC (see Prud’homme van Reine, 1982, p. 179).
Turbinaria (E62/20040a). Desta Island: rocks off the northern coast, 26.x.1965,
lithophytic (ISRSE 65/3169h). Museri Island: in front of Scopus Ridge, 8.x.1965,
among other epiphytes on Turbinari a triquetra (ISRSE 65/2333i-with many
plurilocular reproductive organs), 10.x.1965, on Padina boergesenii (ISRSE 65/
2366g).
COMMENTS: Our plants obtained in spring grew intermingled with S. rigidula on the
same branchlets of Turbinaria that carried several ectocarpalean species as well.
They were found on the central parts of the upper surface of the tetrahedral Turbinaria
branches, whereas the ectocarpaleans grew along the margins. The filaments were
about 4 mm long and 24-39 µm in diameter and were clearly distinguishable from
those of S. rigidula by their greater thickness. Spring plants bore many propagules
and a few unilocular sporangia about 40 µm in diameter. Autumn plants lacked
propagules, but bore many plurilocular sporangia.
We have examined two mounts (UC 409138 and 436446) of the plant that Schimper
(1835-37) distributed (Unio itiner. no. 476) under the name Sphacelaria cervicornis
Ag?, and can confirm the generally held view that they are representative of S.
tribuloides.
DICTYOTALES
DICTYOTACEAE
DICTYOTA Lamouroux (nom. cons.)
*Dictyota apiculata var. jedanensis Weber-van Bosse (Siboga-Exped. Monogr.
59a, p. 183, 1913).
42
Type from Jedan Island, Indonesia, NE of Pulau Aru.
ISRSE SPECIMENS STUDIED : Entedebir Island: northern shore of Ras Papenfuss
25.iii.1962 (E62/20147).
DISTRIBUTION: Warm Indo-west Pacific waters.
COMMENTS : Only two small plants of this alga were encountered. They were mostly
creeping, with free tips 1-2 cm in length.
Womersley (1987, p. 196) regarded D. apiculata J.Agardh (type from Port Phillip,
Victoria, Australia) as a growth form of D. dichotoma (Hudson) Lamouroux. The
taxonomic disposition of D. dichotoma var. jedanesis is thus uncertain.
*Dictyota bartayresiana Lamouroux (J. Bot. [Desvaux] 2: 43, 1809).
Type from the Antilles.
ISRSE SPECIMENS STUDIED : Romia Island: 29.iii.1962 (E62/20293)-bearing sporangia.
COMMENTS : Our plants formed little clumps, 2-3 cm high. This alga is one of a group
of species found only on Romia Island.
*Dictyota ceylanica Kützing (Tab. Phycol. 9, p. 11, pl. 25, fig. I, 1859).
Type from Sri Lanka, leg. W.H. Harvey, presumably in L.
ISRSE SPECIMENS STUDIED : Entedebir Island: Mersat Abiad, 4.iv.1962 (E62/20238).
Museri Island: in front of Scopus Ridge, 10.x.1965 (ISRSE 65/2320i).
COMMENTS : Our plants obtained in spring were about 12 cm tall. This alga, like other
Dictyota species, was much less common in autumn. It is probably a winter-spring
seasonal plant in the southern Red Sea, like many Dictyotaceae in the northern Red
Sea and eastern Mediterranean (Lipkin, 1972a).
Dictyota ciliolata Kützing (Tab. Phycol. 9, p. 12, pl. 27, fig. I, 1859).
Type from La Guaira, near Caracas, Venezuela, leg. J. Smith.
Romia Island: 29.iii.1962 (E62/20293a).
COMMENTS : Our plants from Entedebir Island reached 3 cm in height, whereas those
from Romia Island were twice as tall. This size difference may reflect the more
favorable environmental conditions on Romia Island compared with the inner islands
of the archipelago.
43
LOBOPHORA J. Agardh
Lobophora variegata (Lamouroux) Womersley ex Oliveira (Alg. Mar. Bent. Brasil, p. 217, 1977).
Dictyota variegata Lamouroux (J. Bot. [Desvaux] 2: 40, 1809).
Type from the Antilles, leg. Richard, in herb. Lamouroux in CN (see Womersley
1987, p. 255).
ISRSE SPECIMENS STUDIED : Entedebir Island: Landing Bay, 11.iii.1962, ca. 1 m depth
(E62/20005); northern shore of Ras Papenfuss, 25.iii.1962 (E62/20151). Museri
Island: Camping Bay, off Ras Pandion, 16.x.1965 (ISRSE 65/3047d); off Mangrove
Bay, 14.x.1965 (ISRSE 65/3021a); western part of island, x.1965, on big Tridacna
maxima shell (ISRSE 65/2381).
DISTRIBUTION : Widely distributed in tropical and subtropical waters.
COMMENTS: Our plants were relatively small, reaching only about 3 cm in diameter.
The alga, one of the most common in the northern Red Sea (Lundberg & Lipkin
1979), was uncommon in the ISRSE collections.
PADINA Adanson (nom. cons.)
*Padina boergesenii Allender et Kraft (Brunonia 6: 87, fig. 6C, H, I; fig. 7C, D,
1983).
Type from the U.S. Virgin Islands (Hohenacker, Alg. Mar. Sicc., fasc. 11, no. 515,
1862) in MEL (47044).
ISRSE SPECIMENS STUDIED: Sheikh Sa’id Island: 8.iv.1962 (E62/20262). Cundabilu
Island: western end of islet, 20.iii.1962 (E62/20103). Enteraia Island: 24.iii.1962,
lightly lime-encrusted. Entedebir Island: Goliath Bay, 14.iii.1962, 1 m depth (E62/
20045), 23.iii.1962 (E62/20170), 3.iv.1962 (E62/20236); northern shore of Ras
Papenfuss, 25.iii.1962 (E62/20155); Padina Bay, 4.iv.1962 (E62/20242-sporophytic),
(E62/20140-sporophytic and female), lime-encrusted (E62/20141-sporophytic and
female). Museri Island: in front of Scopus Ridge, 9.x.1965, in Chondrophycus
papillos us community, with epiphytic Polysiphonia, Chondria , Cladopho ra,
Enteromorpha and Ceramium (ISRSE 65/0382-males, 2339), in Caulerpa racemosa
community, with epiphytic Cladophora , Polysiphonia, Chondrophycus papillosus
and Hypnea (ISRSE 65/2348), 10.x.1965, with many epiphytes (ISRSE 65/2315,
2366-with a few sporangia); off Mangrove Bay, 14.x.1965, with Bryopsis plumosa
(ISRSE 65/3001-with oogonia, 3013-with sporangia); Camping Bay, off Ras Pandion,
16.x.1965, with Chaetomorpha , Cladophora and epiphytic Polysiphonia (ISRSE
65/3057, 3068); Camping Bay, 19.x.1965, on rock flat ca. 30 cm below low water
(ISRSE 65/2220-with sporangia). Romia Island: 29.iii.1962 (E62/20198a). Harmil
Island: 28.iii.1962, (E62/20245).
DISTRIBUTION : Widespread in warm seas, extending to warm temperate waters.
44
COMMENTS : Our plants were well developed, to 15 cm tall in spring (usually no more
than 10 cm) and to 10 cm in autumn. They were greenish brown and had a very light
(rarely heavy) encrustation of lime. Longitudinal sections showed two layers of
cells marginally and three layers in the rest of the frond. The cells in the ventral
layer were taller than those of the other two layers. Both sporophytes and gametophytes
were collected in spring and autumn. It seemed that a greater proportion of the
population was fertile in autumn.
This alga, only once reported from the Red Sea previously (Por & Lerner-Segev
1966, as P. gymnospora), is nevertheless very common in both the southern and the
northern parts. Many Red Sea records of P. pavonia are probably based on material
of P. boergesenii (see Papenfuss 1968a, p. 34; De Toni & Levi 1888, p. 138; De
Toni 1904).
Padina boryana Thivy (in Taylor, Pacific Sci. 20: 355, fig. 2, 1966).
Type from Tonga Islands, leg. W.H. Harvey (Harvey, Friendly Islands Algae no. 1),
in NY.
ISRSE SPECIMENS STUDIED : Entedebir Island: northern shore of Ras Papenfuss,
25.iii.1962 (E62/20155a).
COMMENTS : Our plant was only 5 cm tall. It consisted of two layers of cells almost to
the base. The ventral layer was composed of large cells, the dorsal of small cells.
The oogonia developed on the dorsal surface in a few concentric rows, each separated
from the next by a dark line on the thallus in which a few hairs occurred.
This alga was reported from the Red Sea earlier as P. commersonii Bory (Hauck,
1887). For a discussion of its nomenclature see Papenfuss (1977, pp. 276-277).
*Padina somalensis Hauck (Hedwigia 26: 45, 1887).
Type from Somalia (Scara, ii.1873; Lasgori [Las Gori, Las Gore, Las Khorai, Las
Khoreh, Laas Qoray]), iii.1873, leg. J.M. Hildebrandt.
ISRSE
male).
SPECIMENS STUDIED :
Romia Island: 29.iii.1962 (E62/20198b-sporophytic and
DISTRIBUTION: Warm waters of the Indian Ocean and Red Sea.
COMMENTS : Our plants were about 10 cm tall, reddish brown, and fertile. Longitudinal sections showed three layers of cells near the margins and four layers throughout
the rest of the plant.
This alga was not encountered during the second ISRSE.
The record of Padina tetrastromatica Hauck from the southernmost Red Sea (Newton, 1953) may be referable to P. somalensis (see discussion on P. tetrastromatica by
Gaillard 1967).
45
STOECHOSPERMUM Kützing
Stoechospermum marginatum (C. Agardh) Kützing (Phycol. General., p. 339,
1843).
Zonaria marginata C. Agardh (Syst. Alg., p. 266, 1824).
Type from the Red Sea, leg. P. Forsskål, in the Agardh herbarium at LD.
ISRSE
SPECIMENS STUDIED :
Romia Island: 29.iii.1962 (E62/20191-fertile).
DISTRIBUTION : India, Sri Lanka, East Africa (extending southward to Natal province,
South Africa), Red Sea. The species is probably widespread in warm waters of the
entire Indian Ocean although it may be uncommon in many places.
COMMENTS: This is one of the group of algae found only on Romia Island, near the
deep waters of the open sea.
SCYTOSIPHONALES
SCYTOSIPHONACEAE
COLPOMENIA (Endlicher) Derbès et Solier
Colpomenia sinuosa (F.C. Mertens ex Roth) Derbès et Solier (in Castagne, Suppl.
Cat. Pl. Marseille, p. 95, 1851).
Ulva sinuosa F.C. Mertens ex Roth (Catal. Bot. 3, p. 327, pl. XII, 1806).
Type from Cádiz, Atlantic coast of Spain.
ISRSE SPECIMENS STUDIED : Entedebir Island: Padina Bay, 11.iii.1962, (E62/20020).
Harmil Island: 28.iii.1962, (E62/20244).
DISTRIBUTION : Widespread in warm and temperate waters, penetrating boreal waters.
COMMENTS: Both lobed as well as unlobed plants were collected during spring, with
both forms being fertile. No plants were obtained in autumn in agreement with the
situation in the northern Red Sea (Lipkin 1972a) and the eastern Mediterranean
(Lipkin & Safriel 1971), where this alga disappears during spring. Small plantlets
appear late in autumn or during early winter.
HYDROCLATHRUS Bory
Hydroclathrus clathratus (C. Agardh) Howe (in Britton & Millspaugh, Bahama
Fl., p. 590, 1920).
Encoelium clathratum C. Agardh (Sp. Alg. 1(2), p. 412, 1823).
Type from unknown locality (see Silva et al. 1996, p. 631).
ISRSE SPECIMENS STUDIED: Entedebir Island: Padina Bay, 11.iii.1962, on Colpomenia
sinuosa (E62/20020a); Goliath Bay, 23.iii.1962, forming growths of about 30 cm in
diameter (E62/20136). Harmil Island: 28.iii.1962, (E62/20250).
46
COMMENTS: Well-developed plants with plurilocular sporangia were collected in March
(spring). No specimens were encountered in October, in agreement with the seasonal
occurrence of this species in the northern Red Sea and the eastern Mediterranean,
where, like C. sinuosa, it is a winter-spring plant that disappears towards the summer
(Safriel & Lipkin 1964; Lipkin & Safriel 1971).
ROSENVINGEA Børgesen
Rosenvingea intricata (J. Agardh) Børgesen (Dansk Bot. Ark. 2(2): 26, 1914).
Asperococcus intricatus J. Agardh (Ofvers. Förh. Kongl. Svenska Vetensk.-Akad. 4: 7, 1847).
Type from Veracruz, Mexico, leg. Liebmann, presumably in the Agardh herbarium
at LD.
ISRSE SPECIMENS STUDIED : Romia Island: 29.iii.1962, (E62/20189). Harmil Island:
28.iii.1962 (E62/20243).
COMMENTS : Our plants formed clumps reaching 10 cm in diameter. This alga was not
encountered in the Archipelago in the autumn of 1965 and is very likely another
member of the winter-spring element of the Red Sea flora.
FUCALES
CYSTOSEIRACEAE
CYSTOSEIRA C. Agardh (nom. cons.)
Cystoseira myrica (S.G. Gmelin) C. Agardh (Sp. Alg. 1(1), p. 53, 1820).
Fucus myrica S.G. Gmelin (Hist. Fuc., p. 88, pl. III, fig. 1, 1768).
Type locality unknown. Gmelin wrote: “Locus Mare mediterraneum? ad Littora
Kamtschatica quoque legit Kraschenninikow”, but clearly this information is incorrect.
ISRSE SPECIMENS STUDIED : Enteraia Island: 24.iii.1962, (E62/20161). Entedebir Island: Mersat Abiad, 26.iii.1962 (E62/20156); Goliath Bay, 3.iv.1962, (E62/20209);
Landing Bay, 21.x.1965, on bottom covered with coral fragments, at 1-2 m depth
(ISRSE 65/3168). Dahlak Kebir Island: 20.iii.1962 (E62/20096 – cited by Papenfuss
& Jensen, 1967, fig. 4). Museri Island: in front of Scopus Ridge, 10.x.1965 (ISRSE
65/2368), 9.x.1965, on rock flat, ca. 30 cm below low water, in Caulerpa racemosa
community (ISRSE 65/0396, 2354j); off the northern coast, 24.x.1965, drift (ISRSE
65/3093).
DISTRIBUTION: Warm waters of the western Indian Ocean and Caribbean Sea.
COMMENTS : Plants obtained in spring were more than 30 cm tall, with well-developed
receptacular parts. The attached specimens collected during autumn were small (about
47
6 cm tall) and composed only of basal vegetative parts. Attached plants obtained in
both spring and autumn carried much epiphytic Jania. Drifting plants collected in
autumn, which were 25 cm long, were intermingled in floating rafts composed mostly
of Sargassum spp. and Turbinaria triquetra. The Cystoseira myrica from these rafts
consisted of well-developed receptacular parts detached at the end of the growing
season. Attached plants were obtained on rocky substrates in well-illuminated habitats
at depths of 20-200 cm below low water level of spring tides.
HORMOPHYSA Kützing
Hormophysa cuneiformis (J.F. Gmelin) P.C. Silva (in Silva, Meñez & Moe,
Smithsonian Contr. Mar. Sci. 27: 81, 1987).
Fucus cuneiformis J.F. Gmelin (Syst. Nat. ed. 13, 2(2), p. 1389, 1792 (new name).
Fucus articulatus Forsskål (Fl. Aegypt.-Arab., p. 191, 1775), not F. articulatus S.G. Gmelin (Hist.
Fuc., p. 77, 1768).
Type from As Suways (Suez), Egypt, leg. P. Forsskål, in C (see Børgesen 1932, p.
11).
ISRSE SPECIMENS STUDIED : Entedebir Island: Landing Bay, 2.iv.1962, cast up (E62/
20204 – cited by Papenfuss 1968b, p. 43, without referring to collection number).
DISTRIBUTION : Widespread in warm waters in the Indo-west Pacific region, spreading
into warm temperate waters.
COMMENTS: Our plants were well developed, reaching 30 cm in height, and bearing
mature conceptacles.
The alga has previously been reported from the Red Sea as H. triquetra (C. Agardh)
Kützing.
SARGASSACEAE
SARGASSUM C. Agardh (nom. cons.)
Sargassum acinaciforme? Montagne (Ann. Sci. Nat., Bot., ser. 3, 13: 239, 1850).
Type from Al Hudaydah (Hodeida), Yemen, southern Red Sea, leg. Arnaud and
Vasière, no. 23, in PC.
ISRSE SPECIMENS STUDIED : Romia Island: 29.iii.1962 (E62/20292). Harmil Island:
28.iii.1962 (E62/20252).
DISTRIBUTION : Red Sea and Iranian coast of the Arabian Gulf.
COMMENTS: Our plants were to 30 cm tall. The lower leaves were lanceolate, 2.5-4
cm long and 4-6 mm wide, with acute tips; the upper leaves were linear and reached
4.5 cm in length and 1-1.5 cm in width. Vesicles were small, 2-3 mm long, 1 mm in
diameter, tapering towards the apex, with a somewhat flattened stalk. A few vesicles
48
bore an apical spine 1-5 mm in length. Receptacles reached 12 mm in length. All the
plants collected were attached.
*Sargassum herbaceum Kützing (Sp. Alg., p. 607, 1849).
Type from Kh[alineover]rk (Karek) Island, Iran.
ISRSE SPECIMENS STUDIED: Entedebir Island: Landing Bay, 2.iv.1962, cast up (E62/
20206b). Museri Island: 1 km off the northern coast, 25.x.1965, in a floating raft
composed of several Sargassum species (ISRSE 65/2310, 2311).
DISTRIBUTION: Red Sea and Arabian Gulf; Indonesia (Sumatra).
COMMENTS : Our plants reached 110 cm in height. The lower leaves were 2 cm long
and 1 cm wide, with sparsely serrulate margins. The upper ones reached 5 cm in
length and 5-7 mm in width and had serrate margins. The plants obtained in 1965
were from a floating raft which, to judge from the associated animals, had been
floating only a short time.
Sargassum latifolium (Turner) C. Agardh (Sp. Alg. 1(1), p. 13, 1820).
Fucus latifolius Turner: Fuci, vol. 2, p. 66, pl. 94, 1808.
Type from the Red Sea, leg. Lord Valentia, 1807, in BM (ex K).
ISRSE SPECIMENS
20205).
STUDIED:
Entedebir Island: Landing Bay, 2.iv.1962, cast up (E62/
DISTRIBUTION: Red Sea and northern Indian Ocean.
COMMENTS : Our plants reached 40 cm in height, the leaves reaching about the same
size in the lower and in the upper parts of plants: 5 cm in length and 1 cm in width,
with serrate margins. The vesicles were nearly spherical, 5-6 mm in diameter.
Sargassum latifolium C. Agardh f. polycarpum (Figari et De Notaris) Grunow
(Verh. Zool.-Bot. Ges. Wien, 66: 28, 1916).
Sargassum polycarpum Figari et De Notaris (Mem. Reale Accad. Sci. Torino, ser. 2, 13: 147, 1853).
Type from the northern Red Sea (As Suways [Suez], Al ’Aqabah [Akaba].
ISRSE SPECIMENS STUDIED: Museri Island: 1 km off the northern coast, 25.x.1965, in
a floating raft composed of several Sargassum species (ISRSE 65/3190).
COMMENTS : Our plants reached 50 cm in height and were fertile. The leaves were
similar to those of S. latifolium, but a little narrower and with less prominent teeth.
Many goose-barnacles were present on the plants, indicating that they had been
afloat for at least 3-4 months.
49
Sargassum notarisii Zanardini (Mem. Reale Ist. Veneto Sci. Lett. Arti 7: 232, 1858).
Sargassum crispum var. notarisii (Zanardini) Grunow (Verh. Zool.-Bot. Ges. Wien 65: 438, 1915).
Sargassum forsskalii var. notarisii (Zanardini) Papenfuss (Israel J. Bot. 17: 51, 1968).
Type from northern Red Sea (As Suways, Al’Aqabah), leg. Portier.
ISRSE SPECIMENS STUDIED : Entedebir Island: Landing Bay, 21.x.1965, attached (ISRSE
65/3165). Museri Island: 1 km off the northern coast, 25.x.1965, in a floating raft
composed of several Sargassum species (ISRSE 65/2307, 2308).
DISTRIBUTION : Red Sea, African coast of the Indian Ocean, Andaman Islands.
COMMENTS: Our plants reached 40 cm in length. Leaf shape varied between nearly
orbicular to obovate. The lower leaves were smaller and had almost entire margins
whereas those on the upper parts had the delicate serrulation typical of the species.
All previous records of this alga from the Red Sea have been from the northern part
(Papenfuss 1968a). The complicated nomenclature of this species is discussed in
Silva et al. (1996, p. 688).
*Sargassum portierianum Zanardini var. mombassiense Grunow (Verh. Zool.Bot. Ges. Wien, 66: 145, 1916).
Type from Mombasa, Kenya, leg. J.M. Hildebrandt.
ISRSE
SPECIMENS STUDIED :
Entedebir Island: Mersat Abiad, 26.iii.1962 (E62/20157).
DISTRIBUTION : Type locality and Red Sea.
COMMENTS: Our plants reached 60 cm in height.
The nominal variety (type from Quseir [Koseir], Egypt, is not known from the
southern Red Sea.
Sargassum steinitzii nom. nov.
Replaced name: Sargassum tenue J. Agardh, (Sp. Alg. 1, p. 303, 1848), not S. tenue
(Kützing) Endlicher (Gen. Pl. Suppl. 3, p. 31, 1843).
Type from India, Wight 239, presumably in the Agardh herbarium at LD.
ISRSE SPECIMENS
20206a).
STUDIED :
Entedebir Island: Landing Bay, 2.iv.1962, cast up (E62/
DISTRIBUTION : Warm waters of the Indian Ocean and Red Sea.
COMMENTS: Our plants were drifting fragments up to 25 cm long that agreed well
with Agardh’s description of S. tenue.
Reinbold (1907) reported this alga from the Red Sea as S. tenue but this name is
untenable because it is a later homonym of S. tenue (Kützing) Endlicher. Sargassum
gracile Greville (1849 [1848-1849], p. 504, pl. XI, upper figs. 1-3) is a generally
50
accepted heterotypic synonym, but this name is also untenable, being a later homonym of Sargassum gracile J. Agardh (1848, p. 310). Silva et al. (1996, p. 661)
adopted the name S. capillare Kützing (1843, p. 361), which Grunow (1915,
p. 374) treated as a questionable synonym of S. tenue J. Agardh. In view of
the uncertain synonymy, however, it seems appropriate to propose a substitute
name. The new name honours the late Prof. Heinz Steinitz, the leader of the first
ISRSE.
Sargassum subrepandum (Forsskål) C. Agardh (Sp. Alg. 1(1), p. 8, 1820).
Fucus subrepandus Forsskål (Fl. Aegypt.-Arab., p. 192, 1775).
Type from Et Tur (Tor), Sinai Peninsula, northern Red Sea, leg. P. Forsskål, in C
(see Børgesen 1932, p. 12).
Museri Island: 1 km off the northern coast, 25.x.1965, in a floating raft composed
of several Sargassum species (ISRSE 65/3191).
COMMENTS : Our plants reached 40 cm in height.
*Sargassum virgatum C. Agardh (Sp. Alg. 1(1), p. 10, 1820).
Fucus virgatus Mertens (Mém. Mus. Hist. Nat. [Paris] 5: 177, 1819), not F. virgatus Gunnerus (Fl.
Norveg., p. 45, 1766).
Type from the Indian Ocean.
ISRSE SPECIMENS STUDIED: Entedebir Island: Landing Bay, 2.iv.1962, cast up (E62/
20206); Goliath Bay, 3.iv.1962 (E62/20210).
COMMENTS : Our plants reached 45 cm in height.
TURBINARIA Lamouroux
*Turbinaria papenfussii W.R. Taylor (J. Linn. Soc. Bot. 58: 477, pl. 1, figs. 1-9,
1964).
Type from Romia Island, leg. G.F. Papenfuss, E62/20200, 29.iii.1962, in UC (see
Papenfuss 1968a, p. 62).
ISRSE
SPECIMENS STUDIED :
as above.
DISTRIBUTION: Known only from the type locality.
C OMMENTS : No specimens of this alga were encountered during ISRSE 1965. It
seems to be one of the group of species found only on Romia Island.
51
Turbinaria triquetra (J. Agardh) Kützing (Sp. Alg., p. 621, 1849).
Turbinaria vulgaris var. triquetra J. Agardh (Sp. Alg. 1, p. 267, 1848).
Type from the Red Sea, leg. Fontanier, in PC (see Taylor, 1964, p. 479, figs. 1-4).
ISRSE SPECIMENS STUDIED : Sheikh Sa’id Island: 8.iv.1962 (E62/20261). Enteraia
Island: 24.iii.1962, about 3 m long (E62/20144). Entedebir Island: Goliath Bay,
14.iii.1962, at 1.5 m depth (E62/20044 – erroneously cited by Taylor, 1964, p. 480
from Massawa), 3.iv.1962 (E62/20207); Landing Bay, 21.x.1965, 3 m depth (ISRSE
65/3149, 3163). Museri Island: Camping Bay, 19.x.1965, on rock flat, ca. 30 cm
below low water (ISRSE 65/2224); Camping Bay, off Ras Pandion, 25.x.1965, part
of a floating raft (ISRSE 65/3200); 1 km off the northern coast, 25.x.1965, floating
in a Sargassum raft (ISRSE 65/2299), 24.x.1965, drift (ISRSE 65/3095, 3177); off
Mangrove Bay, 8.x.1965 (ISRSE 65/2333), 14.x.1965 (ISRSE 65/2399); Sepia Bay,
17.x.1965 (ISRSE 65/3075).
DISTRIBUTION : Warm waters of the Red Sea and the western Indian Ocean (see Taylor,
1964, p. 479).
COMMENTS: This species provided the largest marine plants encountered during the
two expeditions. They were often 1-2 (-3) m long. Attached plants were smaller in
autumn than in spring.
The specimens on which this species is based were seen by J. Agardh when he visited
Paris. He left a manuscript in the Muséum d’Histoire Naturelle, which was cited by
Decaisne (1841, p. 145) as the source of the name Turbinaria triquetra J. Agardh,
for which, however, Decaisne did not provide a description. When Agardh eventually
published this taxon (1848, p. 267), he decided that it was merely a variety of T.
vulgaris J. Agardh. Kützing (1849, p. 621) treated this taxon under the name Turbinaria
triquetra “J. Ag. Ms” and repeated Agardh’s diagnosis, presumably taken from
Agardh’s “Species Algarum”, which he did not cite.
CHLOROPHYTA
ULVALES
ULVACEAE
ENTEROMORPHA Link (nom. cons.)
Ulva as conceived by Linnaeus (1753) encompassed present-day species of
Enteromorpha as well as Ulva. The tubular form of the thallus in certain species was
used to separate Enteromorpha from Ulva early in the 19th century, and the two
genera continued to be recognized by almost all subsequent authors with the notable
exception of Le Jolis (1863). Recently, Blomster et al. (1999), on the basis of sequence
data from rRNA ITS and 5.8S genes, concluded that the two genera constitute a
single monophyletic clade, within which there are several well-identified clades.
Few of these clades correspond to morphological species that have been described
within these genera. It seems likely that a proposal to merge the two genera will be
forthcoming.
52
Enteromorpha compressa (Linnaeus) Nees (Horae Phys. Berol., index, 1820).
Ulva compressa Linnaeus (Sp. Pl., p. 1163, 1753).
Type from northern Europe.
ISRSE SPECIMENS STUDIED : Dahlak Kebir Island: shore near Das Go village, 13.iii.1962
(E62/20071). Museri Island: off Scopus Ridge, 8.x.1965, on Caulerpa racemosa, C.
sertularioides and Thalassia hemprichii (ISRSE 65/2326c, 2330d, 2332c), 9.x.1965,
epilithic (ISRSE 65/2341a), 10.x.1965, on Halimeda discoidea in Chondrophycus
papillosus community (ISRSE 65/2315f); off Mangrove Bay, 14.x.1965 (ISRSE
65/3041d); Camping Bay, 19.x.1965 (ISRSE 65/2217).
DISTRIBUTION: Widespread in warm and temperate seas, penetrating boreal waters.
COMMENTS : Plants collected in autumn were 2-5 cm in height, whereas those from
spring reached 12 cm.
*Enteromorpha stipitata Dangeard (Botaniste 42: 42, figs. 16, 17, 1959).
Type from Rabat, Morocco.
Chondrophycus papillosus community growing on rock flat ca. 20 cm below low
water, on Padina boergesenii (ISRSE 65/0387).
DISTRIBUTION: Atlantic Morocco, Mediterranean and Red Sea.
COMMENTS : Plants reached 3 cm in height, with a stipe 1-2 mm long, 1-1.5 cm wide
approximately 2 cm from the base, and bearing a number of branches proximally,
each several cells in width. Cells were rectangular or quadrangular, arranged in
transverse rows in the lower parts and in longitudinal rows throughout, 13 × 13 µm
to 26 × 10 µm with 2 or 3(-4) pyrenoids.
Our plants agree in morphology and number of pyrenoids with Enteromorpha stipitata
(Dangeard, 1959, fig. 16, plant on left; fig. 17, E) but the cells are larger than those
of Dangeard’s material, which were 8-9 × 17-18 µm. In this respect, our plants are
closer to E. stipitata var. linzoides Bliding (1960), the cells of which are 17-21 ×
12-14 µm. The latter taxon, however, is reported to have 1-5(-8) pyrenoids (“fully
grown cells often have 3-7 pyrenoids” according to Bliding, 1963, p. 100), whereas
in our plants the maximum number was 4. Since we believe that pyrenoid number is
less variable than cell size, we prefer to assign our plants to E. stipitata rather than
to E. stipitata var. linzoides.
ULVA Linnaeus (nom. cons.)
Ulva lactuca sensu lato nec Linnaeus (see Papenfuss 1960)
ISRSE SPECIMENS
(E62/20108).
STUDIED :
Entedebir Island: Goliath Bay, 22.iii.1962, on Laurencia
53
DISTRIBUTION : Reported from all temperate and tropical seas.
COMMENTS: Our plants were light green, translucent, up to 20 cm long and 10 cm
wide, and sterile.
This species, along with U. reticulata, seems to be a winter species in the southern
Red Sea, as it is in the northern part of this sea and in the eastern Mediterranean.
Observations on the eastern Mediterranean coast (Lipkin, unpublished) suggest that
Ulva requires a periodical intermixing of fresh water, which is provided in this arid
region only by rains in the winter. In localities where freshwater is supplied artificially,
such as sewage outfalls, Ulva thrives all year long.
The name Ulva lactuca has been used by many authors for the Red Sea and eastern
Mediterranean non-reticulate Ulva (see Papenfuss 1968a, p. 12). Föyn (1955) showed
that plants of the Mediterranean and southern European Atlantic coasts were
reproductively isolated from U. lactuca of England and northern European Atlantic
coasts, the provenance of U. lactuca L. He indicated other differences between the
northern and southern European populations and suggested the name U. thuretii for
the latter. This name is invalid, however, since he failed to provide a Latin description
or designate a type specimen. No other name has been proposed for these populations.
Föyn remarked that a thorough description of his U. thuretii was given by Thuret
(Thuret & Bornet 1878). Thuret’s description was based on plants from Cherbourg
which differ in several respects from the eastern Mediterranean and Red Sea plants.
Bliding (1969, p. 554) showed that U. thuretii Föyn is conspecific with U. rigida C.
Agardh. He stated, however, that one of the typical characteristics of Ulva rigida is
the presence of very small marginal teeth. Since our specimens lack such marginal
teeth, it is possible that the eastern Mediterranean and Red Sea plants may represent
still another species. The relationships between the different Ulva populations of
this extensive region should be clarified before any new names are proposed.
Ulva reticulata Forsskål (Fl. Aegypt.-Arab., p. 187, 1775).
Type from the southern Red Sea (Al Qunfudhah, Saudi Arabia; Al Mukha, Yemen),
leg. P. Forsskål, in C (see Børgesen 1932).
Romia Island: 29.iii.1962 (E62/20273). Harmil Island: 28.iii.1962 (E62/20272).
DISTRIBUTION : Warm waters of the Indian and western Pacific oceans.
COMMENTS: Our plants reached 20 cm in length. This conspicuous species has been
reported from only one locality in the northern part of the Red Sea, Al Qusayr
(Kosseir). It is more common in the southern Red Sea. In the central part, Aleem
(1978c) recorded the plant from several localities in the vicinity of Al Judda (Jedda),
all below the Tropic of Cancer; from the northern part it has been reported only
once, from Al Qusayr (Kosseir) (see Papenfuss 1968a). It has never been reported
from the northernmost reaches of the Red Sea.
54
CLADOPHORALES
CLADOPHORACEAE (nom. cons.)
CHAETOMORPHA Kützing (nom. cons.)
Chaetomorpha aerea (Dillwyn) Kützing (Sp. Alg., p. 379, 1849).
Conferva aerea Dillwyn (Brit. Conferv., fasc. 11, pl. 80, 1806).
Type from Cromer, Norfolk, England, leg. D. Turner, in BM (ex K) (see Womersley
1984, p. 174).
ISRSE SPECIMENS STUDIED : Bay of Archico: 9.iv.1962 (E62/20278). Entedebir Island: Goliath Bay, 21.iii.1962 (E62/20097), 22.iii.1962, forming extensive floating
masses (E62/20111); Padina Bay, 12.iii.1962 (E62/20277); Landing Bay, 21.x.1965,
on Cystoseira myrica (ISRSE 65/3168a). Museri Island: off Mangrove Bay, 14.x.1965
(ISRSE 65/3020b, 3035b); Camping Bay, off Ras Pandion, 16.x.1965, on Chondria
(ISRSE 65/2317f, 3056d, 3063); 19.x.1965, epilithic (ISRSE 65/3082h); in front
of Pandion Cliff, 24.x.1965, from floating rafts of Sargassum (ISRSE 65/0381,
3096). Harmil Island: 28.iii.1962 (E62/20276).
COMMENTS : Our plants were composed of long filaments, sometimes 20 cm long or
longer, consisting of cylindrical cells, 650-950 µm long and 340-480 µm in diameter.
The ratio of length to diameter was 1.6-2.8, usually around 2.0-2.2. Isodiametric
cells were not encountered.
Luxuriant growths of this species occurred in floating masses, frequently associated
with Sargassum rafts. It grew less luxuriantly as an epiphyte.
The relationship between Chaetomorpha aerea and C. linum is uncertain. The former
was described from attached plants in England, whereas the latter was described
from floating masses in Denmark. Christensen (1957) concluded that they are growth
forms of the same species. Kornmann (1972), however, on the basis of culture
experiments, believed that the two entities were distinct. Burrows (1991), after
reviewing arguments for and against the conspecificity of these two entities,
recognized only one species, but without strong conviction. We follow Kornmann
(1972) and Blair (1983) in recognizing separate species.
Chaetomorpha linum (O.F. Müller) Kützing (Phycol. Germ., p. 204, 1845).
Conferva linum O.F. Müller, (Fl. Dan., 5(13), p. 7, pl. 771 (2), 1778).
Type from Nakskov Fjord, Lolland, Denmark (see Womersley 1984, p. 176).
ISRSE SPECIMENS STUDIED : Cundabilu Island: western end of beach, 15.iii.1962 (E62/
20056a). Entedebir Island: Mersat Abiad, 26.iii.1962, entangled with Acanthophora
(E62/20163a), 4.iv.1962 (E62/20240); Landing Bay, 21.x.1965 (ISRSE 65/3119).
Museri Island: in front of Scopus Ridge, 8.x.196 5, on Caulerpa selago, C.
sertularioides and Turbinaria triquetra (ISRSE 65/0301d, 2325, 2330i, 2333d),
9.x.1965, on many algae in Chondrophycus papillosus community (ISRSE 65/2341c),
55
in Caulerpa racemosa community (ISRSE 65/2352) , 10.x.19 65, on Padina
boergesenii (ISRSE 65/2367k), 19.x.1965 (ISRSE 65/0347, 3076c); off Mangrove
Bay, 14.x.1965 (ISRSE 65/2398c, 3023); Camping Bay, in front of Ras Pandion,
16.x.1965 (ISRSE 65/3065b, 3068c). Romia Island: 29.iii.1962 (E62/20274). Harmil
Island: 28.iii.1962 (E62/20275).
DISTRIBUTION : Almost cosmopolitan.
COMMENTS : The plants, which we assign to this species, were more slender than
those assigned to C. aerea, the filament diameter in spring being a minimum of 200230 µm in one population (E62/20056a) and a maximum of 340-385 µm in another
(E62/20163a). The cell length to width ratio was 0.5-2 and 1-1.5, respectively.
Plants collected in autumn were thinner (minimum diameter of 95-100 µm and
maximum of 140-145 µm) than those obtained in spring. Cell length varied from
175 to 310 µm. Cell length to width ratio was 1.2-2.
CLADOPHORA Kützing (nom. cons.)
*Cladophora lehmanniana (Lindenberg) Kützing (Phycol. General., p. 268, 1843).
Conferva lehmanniana Lindenberg (Linnaea 14: 179, pl. II, 1840).
Type from Helgoland, leg. Binder, isotypes in L (937,281...47) and in herb. Agardh,
LD (9341-9343) (see van den Hoek 1963, p. 182).
65/2325b), 9.x.1965, on Avrainvillea amadelpha (ISRSE 65/2343b), 10.x.1965
(ISRSE 65/2383), 19.x.1965, on rock flat ca. 30 cm deep, epilithic and on many
algae (ISRSE 65/2210, 2211, 3078, 3079); off Mangrove Bay, 14.x.1965 (ISRSE
65/2398); Camping Bay, off Ras Pandion 16.x.1965 (ISRSE 65/3069b).
DISTRIBUTION : Warm seas of eastern Atlantic and Indian oceans, Mediterranean, Red
Sea, southern Australia.
COMMENTS: Our plants reached 12 cm in height and were very common as epiphytes
in shallow waters.
This species is very similar to Cladophora vagabunda in habit. It differs from the
latter by the gradual tapering of axes towards the apex, with neighboring cells being
of almost the same diameter, whereas in C. vagabunda there is a marked decrease in
diameter of the terminal cells. It is possible, therefore, that at least part of the records
of C. vagabunda (as C. crystallina and C. fascicularis) from the Red Sea, represent
C. lehmanniana.
Previous Red Sea records of Cladophora lehmanniana are all from the north.
Cladophora vagabunda (Linnaeus) van den Hoek (Rev. Europ. Sp. Cladophora, p.
144, 1963).
Conferva vagabunda Linnaeus (Sp. Pl., p. 1167, 1753).
56
Type from a salt marsh on Selsey Island, Sussex, England, in herb. Dillenius, OXF
(see van den Hoek 1963, p. 144).
ISRSE SPECIMENS STUDIED: Cundabilu Island: 20.iii.1962 (E62/20099). Museri Island: in front of Scopus Ridge, 19.x.1965 (ISRSE 65/3077).
DISTRIBUTION: Widespread in warm and warm-temperate waters.
COMMENTS : Our plants were 5 to 10 cm tall. The main axis in plants from Cundabilu
Island were 50-100 µm in diameter, whereas it measured 100-150 µm diameter in
plants from Museri Island. The two sets of plants were very similar in habit. The
species, which was not common, has been previously reported from three stations in
the northern Red Sea (as C. crystallina and C. fascicularis, see Papenfuss, 1968a,
p. 14).
SIPHONOCLADACEAE (nom. cons.)
BOERGESENIA J. Feldmann
Boergesenia forbesii (Harvey) J. Feldmann (Compt. Rend. Hebd. Séances Acad.
Sci. [Paris] 206: 1504, 1938).
Valonia forbesii Harvey (Proc. Amer. Acad. Arts 4: 333, 1860).
Type from Ryukyu-retto, Japan, leg. C. Wright, in TCD.
ISRSE SPECIMENS STUDIED : Enteraia Island: 15.iii.1962, eastern coast, lower intertidal
(E62/20047), 24.iii.1962 (E62/20142). Entedebir Island: Goliath Bay, 16.iii.1962
(E62/20059), 6.iv.1962, intertidal (E62/20265). Romia Island: 29.iii.1962 (E62/
20187, 20266).
COMMENTS : The plants were large, with vesicles reaching 4 cm in length and ca. 1
cm in width. They were obtained in spring, but not in autumn. Plants consisted of
groups of 4 to 10 large vesicles with a great number of smaller basal vesicles. Some
of the plants obtained on Romia Island had vesicles that were once or twice
dichotomously branched rather than the usual ones, which may be basally proliferous
but are never branched (Fig. 7). Prof. Eric Coppejans informs us that he has observed
such plants at many sites in the Indian Ocean.
CLADOPHOROPSIS Børgesen (nom. cons.)
Cladophoropsis herpestica (Montagne) Howe (Mem. Torrey Bot. Club 15: 31,
1914).
Conferva herpestica Montagne (Prodr. Gen. Phyc., p. 15, 1842).
Type from the Bay of Islands, New Zealand, in herb. Montagne, PC (see Womersley
1984, p. 184).
57
Fig. 7. Boergesenia forbesii (E62/20266): unusual branched vesicle from Romia Island. Scale bar =
10 mm.
intermingled with Cladophoropsis membranacea, Chondria, Chaetomorpha and
Ceramium (ISRSE 65/3222).
DISTRIBUTION : Widespread in the warm Indo-west Pacific and Mediterranean.
COMMENTS: This species has previously been reported from the Red Sea under the
name Cladophoropsis zollingeri (Kützing) Børgesen (see Papenfuss 1968a, p. 16).
Cribb (1960) and Papenfuss (loc. cit.) pointed to the possible conspecificity of C.
zollingeri, described from Java (Kützing 1849, p. 415), and C. herpestica. The two
taxa were said to differ in filament diameter and cell wall thickness, but Cribb
(1960) showed that these characters vary enough so that the distinction between the
species cannot be maintained.
In our plants the filaments were ca. 260 µm in diameter; they had thick, striated
walls (14-40 µm), and typical basal rhizoids (see Kützing, 1854, pls. 64, 66). Empty
reproductive cells were present in autumn.
The plants formed cushions intermingled with Cladophoropsis membranacea in the
upper midlittoral zone. In the southern Red Sea this species appeared to be much less
common than C. membranacea.
Cladophoropsis membranacea (Hofman Bang ex C. Agardh) Børgesen (Overs.
Kongel. Danske Vidensk. Selsk. Forh. 1905: 289, legends to figs. 8-13, 1905).
Conferva membranacea Hofman Bang ex C. Agardh (Syst. Alg., p. 120, 1824).
58
Type from St. Croix, U.S. Virgin Islands, presumably in the Agardh herbarium at LD.
ISRSE SPECIMENS STUDIED : Entedebir Island: Padina Bay, 13.iii.1962, forming more
or less circular cushions (E62/20025); Goliath Bay, 16.iii.1962, forming extensive
mats in shade of overhanging ledge (E62/20061), growing in clumps, coarser than
E62/20061 (E62/20062), 3.iv.1962 (E62/20216); Landing Bay, 21.x.1965, (ISRSE
65/3142); northern shore of Ras Papenfuss, 25.iii.1962, forming mats (E62/20294).
Desta Island: rocks off the northern coast, 26.x.1965, on Chondrophycus papillosus
(ISRSE 65/3169b). Museri Island: in front of Scopus Ridge, 8.x.1965, with Caulerpa
racemosa and C. serrulata (ISRSE 65/2326e, 2330h), 9.x.1965, on Avrainvillea
amadelpha, Hypnea cornuta, Chondrophycus papillosus, Cystoseira myrica and
Chaetomorpha (ISRSE 65/2337b, 2343g, 2344e, 2350c, 2352c), 10.x.1965, epilithic
and on Cystoseira myrica (ISRSE 65/2320c, 2368e, 2382), 19.x.1965 (ISRSE 65/
3084i); off Mangrove Bay, 14.x.1965, on Avrainvillea amadelpha (ISRSE 65/2317g);
Camping Bay, 19.x.1965, rock flat 30 cm below low water (ISRSE 65/2212), off
Ras Pandion, 16.x.1965 (ISRSE 65/3065).
C OMMENTS : Our plants grew on rocks in the intertidal zone, where they formed
dense cushions which sometimes formed extensive mats, usually towards the lower
limits of the belt in which the alga grew extensively (see below). In deeper habitats
they usually grew epiphytically, forming sparse growths on their hosts. The filaments
were about 180 µm in diameter and had thin walls.
This was the dominant alga in a belt in the upper midlittoral where it filled small pits
in the lower part of the concave, eroded, fossil reef wall, shaded by the overhanging
ledge typical of the rocky shores of the islets in the archipelago (see Wainwright
1965, figs. 3, 4; Lipkin 1987, figs. 3, 4), and in the upper part of the rock flat
(Lipkin 1987).
This species was first reported from the Red Sea by Lundberg & Lipkin (1979), who
found it in the stomach of a herbivorous fish (Siganus) in the Gulf of Elat.
DICTYOSPHAERIA Decaisne ex Endlicher
Dictyosphaeria cavernosa (Forsskål) Børgesen (Dansk Bot. Ark. 8(2): 2, pl. 1, fig.
1, 1932).
Ulva cavernosa Forsskål (Fl. Aegypt.-Arab., p. 187, 1775).
Type from the southern Red Sea (Al Qunfudhah, Saudi Arabia; Al Mukha, Yemen),
leg. P. Forsskål, in C (see Børgesen 1932).
ISRSE SPECIMENS STUDIED: Sheikh Sa’id Island: 8.iv.1962 (E62/20270). Entedebir
Island: 12.iii.1962 (E62/20013); eastern coast, 15.iii.1962, lower intertidal (E62/
20049); Goliath Bay, 22.iii.1962 (E62/20110). Museri Island: off Mangrove Bay,
14.x.1965, on dead coral (ISRSE 65/2394). Harmil Island: 28.iii.1962 (E62/20253).
DISTRIBUTION: In most warm seas.
59
COMMENTS: Many small plants, 1 cm in diameter, as well as large ones, up to 5 cm in
diameter, were present in spring. During autumn only large plants were collected.
Those from Museri Island grew on solid substrate of coral-limestone at a depth of 1
m in well-lluminated habitats.
PHYLLODICTYON J.E. Gray
Phyllodictyon anastomosans (Harvey) Kraft et Wynne (Phycol. Res. 44: 129. 1996).
Cladophora ? anastomosans Harvey (Phycol. Austral. 2, pl. CI, 1859). Struvea anastomosans
(Harvey) Piccone et Grunow ex Piccone (Croc. Corsaro Madera e Canarie, p. 20, 1884).
Type from Fremantle, Western Australia, leg. W.H. Harvey, in TCD (Alg. Austral.
Exsicc. no. 582a).
ISRSE SPECIMENS
(E62/20050).
STUDIED :
Entedebir Island: Landing Bay, 15.iii.1962, 3 m depth
DISTRIBUTION : Widespread in most warm waters and warm-temperate seas.
COMMENTS: Our specimens were small, only 12 mm tall, but fully developed.
This species has been known for more than a century as Struvea anastomosans .
Recently, however, Kraft & Wynne (1996) showed that in S. plumosa Sonder, the
type of the generic name, development of the thallus regularly involves segregative
division whereas in S. anastomosans septa are produced by the centripetal deposition
of cell-wall material. Retaining the name Struvea for the type and one other species
(S. elegans Børgesen) for which segregative division has been reported, Kraft &
Wynne resurrected the generic name Phyllodictyon J.E. Gray (J. Bot. [London] 4:
69. 1866) for eight species previously assigned to Struvea but lacking regular
segregative division.
This species has been reported only from the northern part of the Red Sea (Nasr
1947). It is common on the Sinai coasts (Lipkin, unpublished), however, and is
probably present throughout the Red Sea.
SIPHONOCLADUS Schmitz
*Siphonocladus tropicus (P. et H. Crouan) J. Agardh (Lunds Univ. Års-Skr., Afd.
Math. och Naturvetensk. 23(2): 105, 1887).
Apjohnia tropica P. et H. Crouan, in Schramm & Mazé (Essai Alg. Guadeloupe, p. 47, 1865).
Type from Guadeloupe (Moule; Basse-Terre; Marie Galante, Grand-Bourg).
ISRSE
SPECIMENS STUDIED:
COMMENTS: Our plants grew in shallow water. This alga, hitherto not reported from
the Red Sea, has subsequently been found at a depth of 45 m off Elat in the north
(Lipkin, pers. obs.). It was never obtained in the Elat area in shallow water.
60
VALONIACEAE
VALONIA C. Agardh
Valonia aegagropila C. Agardh (Sp. Alg. 1(2), p. 429, 1823).
Type from the lagoons of Venezia, Adriatic Sea, in the Agardh herbarium, LD 15977.
ISRSE SPECIMENS STUDIED: Entedebir Island: Landing Bay, 11.iii.1962, ca. 1 m depth
(E62/20006), 21.x.1965 (ISRSE 65/3142a); Goliath Bay, 16.iii.1962 (E62/20060).
Museri Island: in front of Scopus Ridge, 9.x.1965 (ISRSE 65/2343k), 10.x.1965
(ISRSE 65/2214), 19.x.1965 (ISRSE 65/3085f); off Mangrove Bay, 14.x.1965
(ISRSE 65/3035, 3043a); Camping Bay, off Ras Pandion, 16.x.1965 (ISRSE 65/
3053).
COMMENTS : The plants formed cushions 4-8 cm in diameter, sometimes of somewhat
irregular shape, and 1-1.2 cm thick. They grew in shallow water on rocks,
accumulating sediment in the spaces between the vesicles.
*VALONIOPSIS Børgesen
*Valoniopsis pachynema (G. Martens) Børgesen (Biol. Medd., Kongel. Dansk.
Vidensk. Selsk. 11(6): 10, figs. 1, 2, 1934).
Bryopsis pachynema G. Martens (Preuss. Exped. Ost-Asien, Tange, p. 24, pl. IV, fig. 2, 1868).
Type from Sumatra, Indonesia (near Bengkulu; Pulau Tikus), leg. Eduard von Martens,
zoologist of the Expedition, son of the author of the species.
ISRSE
SPECIMENS STUDIED:
COMMENTS: The plants formed cushions 2-6 cm in diameter and 10-15 mm thick.
The main filaments were 550-800 µm in diameter and the ultimate ones about
500 µm.
This alga is reported here for the first time from the southern Red Sea. It is fairly
common on the coasts of the Sinai Peninsula in the northern Red Sea (Lipkin,
unpublished) and is probably present throughout the Red Sea.
BRYOPSIDALES
BRYOPSIDACEAE
BRYOPSIS Lamouroux
Bryopsis plumosa (Hudson) C. Agardh (Sp. Alg. 1(2), p. 448, 1823).
Ulva plumosa Hudson (Fl. Angl., ed. 2, p. 571, 1778).
Type from Exmouth, Devon, England (see Womersley 1984, p. 282).
61
ISRSE SPECIMENS STUDIED : Cundabilu Island: eastern end of beach, 15.iii.1962 (E62/
20055). Entedebir Island: Goliath Bay, 23.iii.1962 (E62/20135). Museri Island: off
Mangrove Bay, 14.x.1965, on rocks 1 m depth (ISRSE 65/2396).
COMMENTS: Our plants reached 4 cm in height, the ones obtained during autumn
being a little larger than those obtained during spring. The latter had ramuli only on
the distal 3-5 (-10) mm of the branches, whereas the branches of the former had
ramuli along most of their length.
DERBESIA Solier
Derbesia tenuissima (Moris et De Notaris) P. et H. Crouan (Fl. Finistère, p. 133,
1867).
Bryopsis tenuissima Moris et De Notaris (Mem. Reale Accad. Sci. Torino, ser. 2, 2 (Cl. Sci. Fis. e
Mat.): 259, pl. VI, fig. III, 1839).
Type from Isola di Capraia, ca. 70 km SW of Livorno, Italy, presumably in TO.
Silva et al. (1996, p. 810) erred in citing the type locality as Isla Cabrera, Islas
Baleares.
ISRSE SPECIMENS STUDIED : Entedebir Island: Goliath Bay, 23.iii.1962, forming low
cushiony growths (E62/20132).
COMMENTS: Our plants had filaments that were 30-70 µm in diameter and irregularly
subdichotomously branched. Sporangia occurred in considerable numbers; they varied
in shape, being pyriform, obovate, or clavate and were 60-90 µm in diameter and
110-130 µm long.
This is the first report of this alga from the southern Red Sea; hitherto it has been
reported only from its northern part (Al Ghurdaqah). Halicystis parvula Schmitz ex
G. Murray, the gametophytic stage of this species, has not yet been recorded from
the Red Sea.
CODIACEAE
CODIUM Stackhouse
Codium arabicum Kützing (Tab. Phycol. 6, p. 35, pl. 100, fig. II, 1856).
Type from Et Tur (Tor), Sinai Peninsula, Red Sea, leg. G.W. Schimper, 26.iii.1835
(Unio Itiner. no. 469), L 910.187-1920.
ISRSE SPECIMENS
20074).
STUDIED :
Cundabilu Island: eastern end of islet, 20.iii.1962 (E62/
DISTRIBUTION : Warm waters of the Indo-west Pacific region.
COMMENTS: Our plants were small, to 3 cm in diameter.
62
CAULERPACEAE
CAULERPA Lamouroux
Most ISRSE specimens of this genus were identified by W.R. Taylor, who published
on those of the first expedition (Taylor 1967). The ecology of Red Sea Caulerpa
was discussed by Lipkin (1974).
*Caulerpa fastigiata Montagne (Ann. Sci. Nat., Bot., ser. 2, 8: 353, 1837).
Type from Cuba, leg. Ramon de Sagra, in PC (see Weber-van Bosse 1898, p. 262).
ISRSE SPECIMENS STUDIED: Entedebir Island: Padina Bay, 12.iii.1962 (E62/20014);
Goliath Bay, 3.iv.1962 (E62/20211).
COMMENTS: Our plants were well developed, to 5 cm tall, forming dense tufts.
Caulerpa lentillifera J. Agardh (Mus. Senckenberg. 2: 173, 1837).
Type from the coast of Eritrea, southern Red Sea, leg. Rüppell, in herb. Agardh, LD
16851 (see Womersley & Bailey, 1970, p. 275).
ISRSE SPECIMENS STUDIED : Entedebir Island: Landing Bay, 21.x.1965 (ISRSE 65/
3144). Museri Island: in front of Scopus Ridge, 10.x.1965 (ISRSE 65/2364, 2375).
DISTRIBUTION: Widespread in warm waters of the Indian and central Pacific oceans.
COMMENTS: Our plants were large, with assimilators reaching 10 cm in height and
stolons extending to 35-40 cm in length. They were of a translucent-green color in
the shade, tending to bleach in brighter light, the tips of the assimilators becoming
yellowish.
On Museri Island the alga grew in crevices ca. 20 cm wide in a rock flat in a
somewhat shaded habitat, together with C. racemosa which was a little etiolated (the
so-called var. occidentalis).
Caulerpa mexicana Sonder ex Kützing (Sp. Alg., p. 496, 1849).
Type from the Atlantic coast of Mexico, in L.
20056) , 20.ii i.1962 (E62/20098 – erroneously cited by Taylor, 1967 from
15.iii.1962). Museri Island: in front of Scopus Ridge, 9.x.1965 (ISRSE 65/0390),
10.x.1965 (ISRSE 65/2236). Harmil Island: 28.iii.1962, (E62/20247a).
COMMENTS: Our plants had relatively short, thin, leaf-like assimilators 3-5 cm long
and slender stolons 1.5-2 mm in diameter. At Museri Island they grew on a rock flat
overlain by a layer of calcareous sand a few mm thick, and were covered, even at
63
low tides, by 20-50 cm of water. The alga occurred in both the Chondrophycus
papillosus and the Caulerpa racemosa communities, which together covered much
of the rock flat surrounding the island (Lipkin 1987).
Caulerpa peltata Lamouroux (Nouv. Bull. Sci. Soc. Philom. Paris, 1: 332, 1809).
Type from the Antilles.
ISRSE SPECIMENS STUDIED : Cundabilu Island: 14.iii.1962, on Tridacna shell at 3 m
depth (E62/20052), 20.iii.1962 (E62/20152). Entedebir Island: Landing Bay,
21.x.1965 (ISRSE 65/3115, 3140, 3146). Nocra Island: south side opposite village,
16.iii.1962, 1-2 m depth (E62/20063). Museri Island: off Mangrove Bay, 14.x.1965
(ISRSE 65/3002).
DISTRIBUTION : Widespread in tropical and subtropical seas.
COMMENTS: (Sample E62/20064 was erroneously cited by Taylor (1967) under
both C. peltata and C. racemosa var. turbinata; it is C. racemosa.)
COMMENTS: Our plants were rather large for this species, forming loose dark-green
mats reaching 20 cm in diameter. They grew in shaded habitats, such as small caves
and overhangs, attached to the rock by short rhizoids. The peltate ramuli, which
usually occurred singly on stalks but sometimes were in small groups (up to five),
were thin and reached 5 mm in diameter. Where more light was available, as at the
entrance to caves, the peltate ramuli were thicker and smaller, while in the highest
light intensity the species can stand, they were replaced by pyriform ramuli.
This alga was reduced to a variety of Caulerpa racemosa by Eubank (1946), and this
treatment has been followed by many authors. While it is clear that the shape of the
assimilators in Caulerpa is determined by light intensity and temperature, it is not
clear what name or names should be given to populations with peltate ramuli. We
believe that the similarity between the pyriform ramuli of the bright-light ecophene
of C. peltata and those of C. racemosa is only superficial. The low-light ecophenes
of the two species are very different, as shown by culture experiments (Lipkin,
1974, p. 165).
Using cross-gradient cultures, Enomoto & Ohba (1987), Ohba & Enomoto (1987),
and Ohba et al. (1992) showed that peltata-type ramuli are formed under conditions
of low light intensity and high temperatures while ramuli characteristic of C.
racemosa var. laetevirens (clavate with obtuse heads) are formed under conditions of
high light intensity and low temperatures. The results were the same whether the
inoculating stock bore peltate or clavate ramuli. Those authors concluded that var.
peltata and var. laetevirens are ecophenes of the same species. They were ambiguous
regarding the name of this species, but presumably they meant C. racemosa. While
granting the conspecificity of these two ecophenes, we suggest that this species is
distinct from C. racemosa.
64
Caulerpa racemosa (Forsskål) J. Agardh (Lunds Univ. Års-Skr., Afd. Math. och
Naturvetensk. 9(8): 35, 1873).
Fucus racemosus Forsskål (Fl. Aegypt.-Arab., p. 191, 1775).
Type from As Suways (Suez), Egypt, northern Red Sea, leg. P. Forsskål, in C (see
Børgesen 1932, p. 9; Papenfuss & Egerod 1957, p. 88).
ISRSE SPECIMENS STUDIED : Cundabilu Island: 20.iii.1962, juvenile form (E62/20077).
Enteraia Island: 24.iii.1962 (E62/20143). Entedebir Island: Landing Bay, 11.iii.1962,
ca. 1 m depth (E62/20002); Goliath Bay, 16.iii.1962 (E62/20068), 23.iii.1962 (E62/
20133 – erroneously cited by Taylor, 1967 as 22.iii.1962), 30.iii.1962 (E62/20195
– erroneously cited by Taylor, 1967 as E62/10195). Nocra Island: southern side
opposite village, 16.iii.1962, 1-2 m depth (E62/20064). Um Aabak: by the Nocra
Channel, 22.iii.1962 (E62/20113). Museri Island: in front of Scopus Ridge, 8.x.1965
(ISRSE 65/2326), 9.x.1965 (ISRSE 65/0394, 2244, 2340, 2347, 2362 – from different plant communities 10.x.1965 (ISRSE 65/2230, 2231, 2232, 2263, 2384 – from
different plant communities); off Mangrove Bay, 14.x.1965 (ISRSE 65/2390, 3017,
3034); Camping Bay, 19.x.1965 (ISRSE 65/2218), off Ras Pandion, 16.x.1965
(ISRSE 65/3061). Romia Island: 29.iii.1962 (E62/20264). Harmil Island: 28.iii.1962,
(E62/20247).
COMMENTS : Many of the formae and varieties described for this species are merely
ecophenes (see Peterson 1972; Lipkin 1974; Calvert 1976), their morphology being
governed mainly by light intensity, length of exposure during low tides, degree of
exposure to wave action, or sometimes temperature (Lipkin & Friedmann 1967). At
many localities individuals may extend into more than one habitat, resulting in different phenotypes on a single plant, sometimes even on the same stolon. These parts
of the plant, if separated (as they often are in herbarium material), would suggest
different formae or varieties. The varietal epithets may be useful in referring to
different ecophenes. Among the ISRSE specimens, the turbinata-ecophene (with
obconical ramuli) grew in bright light on rock flats in shallow water, sometimes
exposed during low water of spring tides. The occidentalis-ecophene (with
subspherical ramuli) grew in partial sunlight in crevices about 20 cm wide and to
40 cm deep in the rock flat, where a dense growth of C. racemosa and C. lentillifera
was present. Some stolons, extending into narrow fissures where the light intensity
was low, developed ramuli typical of the lamourouxii-ecophene (few or widely
scattered, pyriform or clavate).
The degree of morphological plasticity within a population varies from population
to population, suggesting taxonomic boundaries within the species as presently
circumscribed (Calvert 1976; Carruthers et al. 1993). Furthermore, Benzie et al.
(1997) showed that the genetic distance between some varieties of C. racemosa in
Queensland is as large as that between species of the genus. The same situation may
obtain in the Mediterranean Sea (see Verlaque et al. 2000).
It is clear that a single sampling of C. racemosa can not be assigned with certainty to
a particular infraspecific taxon. We have therefore put all racemosa-like samples
65
from the Dahlak Archipelago under the name C. racemosa without recognizing
infraspecific taxa. Culture experiments and genetic studies would help to shed light
on the taxonomic status of various morphs and populations currently assigned to this
vexatious species.
Caulerpa selago (Turner) C. Agardh (Syn. Alg. Scand., p. XXIII, 1817).
Fucus selago Turner (Fuci, vol. 1, p. 122, pl. 55, 1808).
Type from the Red Sea, leg. Lord Valentia.
ISRSE SPECIMENS STUDIED : Entedebir Island: Goliath Bay, 22.iii.1962 (E62/20106);
Mersat Abiad, 22.iii.1962 (E62/20166). Museri Island: Braathen Bay, 7.x.1965, on
calcareous sand, ca. 50 cm below low water (ISRSE 65/0300, 0398), 8.x.1965 (ISRSE
65/0301).
DISTRIBUTION : Indian Ocean and western Pacific.
COMMENTS: Our plants had erect branches (assimilators) to 13 cm long. In his study
of spring samples, Taylor (1967, p. 15, pl. I) described and illustrated assimilators in which the lower part of the axis was roughened by the stub bases of discarded
ramelli. Similar assimilators were found among plants collected in the autumn of
1965. The shedding of ramuli thus seems to be due to senescence rather than to
seasonality.
This alga, one of the larger species of Caulerpa in the Dahlak area, is also one of the
least common. On Museri Island it was restricted to the shallow water of sand-filled
depressions in the fossil reef flat. Morphological variation among our plants was
correspondingly restricted and was probably due to differences in age rather than in
environment.
Caulerpa serrulata (Forsskål) J. Agardh (Mus. Senckenberg. 2: 174, 1837).
Fucus serrulatus Forsskål (Fl. Aegypt.-Arab., p. 189, 1775).
Type from Al Mukha, Yemen, southern Red Sea, leg. P. Forsskål, in C under Fucus
laminosus (see Børgesen 1932, p. 5).
ISRSE SPECIMENS STUDIED : Cundabilu Island: western end of beach, 15.iii.1962 (E62/
20058). Entedebir Island: Padina Bay, 11.iii.1962 (E62/20007); northern shore of
Ras Papenfuss, 25.iii.1962 (E62/20150); Landing Bay, 6.iv.1962, exposed in the
intertidal zone (E62/20254 – erroneously cited by Taylor, 1967 as 20354), 21.x.1965
(ISRSE 65/3123). Museri Island: off Mangrove Bay, 14.x.1965 (ISRSE 65/2392,
3012); off Ras Pandion, 16.x.1965, (ISRSE 65/3055). Museri trough, 40°23’E
15°32’30"N, 19.x.1965, dredged from 120 m, on soft bottom of greenish-light brown
mud with a few shell fragments and larger particles (ISRSE 65/2303); 40°17’E
15°32’N, 22.x.1965, dredged from 110 m, on soft bottom of light brown mud with
a few shell fragments (ISRSE 65/2301, 2302). Romia Island: 29.iii.1962 (E62/
20263). Hawakil (Howakil, Ouachil) Bay, stn. no. 4, 40°19’E 14°58’N, 17.x.1965,
66
dredged from 9-11 m, on soft mud flat, with a few filamentous epiphytes (ISRSE
65/3167).
COMMENTS : This species is the second most common Caulerpa in the southern Red
Sea, and in certain stations it is the most common. Like the most common species,
C. racemosa, it shows considerable morphological variation in different habitats. It
seems that light intensity is the most important factor influencing its morphology.
The bright-light ecophene has dichotomously branched, spirally twisted assimilators
that are serrulate at their margins. Ramuli of the assimilators tend to be terete or
nearly so. In the low-light ecophene, the so-called var. boryana, the assimilators
remain dichotomously branched and serrulate, but lose their spiral twisting, with the
ramuli becoming flattened and band-shaped rather than terete. Nasr (1947) reported
that plants in aquaria lose the serration if no water movement occurs. Such plants,
with entire margins, were not encountered in the field. The low-light ecophene, or
“boryana-ecophene”, was obtained from horizontal surfaces in deep water, in crevices
and other shaded habitats in shallow water, on mud flats at ca. 10 m, where water
movement presumably occurs, and at 110-120 m, where the water is presumably
still.
Caulerpa sertularioides (S.G. Gmelin) Howe (Bull. Torrey Bot. Club 32: 576,
1905).
Fucus sertularioides S.G. Gmelin (Hist. Fuc., p. 151, pl. XV, fig. 4, 1768).
Type from tropical America.
ISRSE SPECIMENS STUDIED : Enteraia Island: 24.iii.1962 (E62/20138 – cited by Taylor,
1967, p. 16). Museri Island: in front of Scopus Ridge, 8.x.1965 (ISRSE 65/2330),
9.x.1965, in Caulerpa racemosa community on rock flat sloping gently seawards, at
ca. 20 cm below low water (ISRSE 65/0393), in Caulerpa racemosa community, on
calcareous sand, ca. 15 cm below low water (ISRSE 65/2243, 2338), 10.x.1965 on
rock flat at 30 cm below low water (ISRSE 65/2237, 2238, 2373); Camping Bay,
off Ras Pandion, 16.x.1965, on rock flat, at 50 cm below low water (ISRSE 65/
2229).
DISTRIBUTION: Widespread in tropical waters.
COMMENTS : This is another of the less common species of Caulerpa.
Our specimens were identified by W.R. Taylor as var. sertularioides f. brevipes.
However, culture experiments (Calvert 1976) have shown that in this species as in
other species of Caulerpa, “variety” and “forma” morphologies are not genetically
controlled, but are markedly influenced by environmental conditions, light intensity
in particular. Thus it appears that infraspecific names in this species connote ecophenes
rather than infraspecific taxa.
Unlike our plants of C. racemosa and C. serrulata, those of C. sertularioides were
restricted to horizontal surfaces in shallow water. This narrow ecological niche was
reflected in morphological uniformity.
67
*CAULERPELLA Prud’homme van Reine et Lokhorst
*Caulerpella ambigua (Okamura) Prud’homme van Reine et Lokhorst (Nova
Hedwigia 54: 114, pls. 1-4, 1992).
Caulerpa ambigua Okamura (Bot. Mag. (Tokyo), 11: 4, pl. 1, figs. 3-12, 1897).
Type from Ogasawara-jima, Ogasawara-gunto (Bonin Islands), Japan.
ISRSE SPECIMENS STUDIED : Cundabilu Island: 14.iii.1962, on Tridacna shell at 3 m
depth (E62/20052a), 20.iii.1962 (E62/20073b, 20075a-erroneously cited by Taylor
1967 as E62/70045a, 20105).
DISTRIBUTION : Most warm seas.
C OMMENTS : A detailed description of our plants was given by Taylor (1967, as
Caulerpa ambigua). He observed that some axes developed clusters of lobed structures,
which he interpreted as starch-storing organs. In a study of Caulerpa ambigua from
the Cape Verde Islands, Prud’homme van Reine & Lokhorst (1992) observed similar
clusters and noted that the axes terminated in what they considered to be discharge
pores. In addition, they observed a transverse wall at the base of each cluster, which
were devoid of trabeculae. Interpreting these clusters as zoidangia, they pointed out
that the non-holocarpic reproduction in this species contrasted with the holocarpic
reproduction in all other species of Caulerpa for which sexual reproduction is known.
They thus established the genus Caulerpella to accommodate Caulerpa ambigua.
Silva et al. (1996, p. 813) preferred to retain Caulerpa ambigua in Caulerpa because
it shares the highly specialized trabeculate structure of all other species of the genus.
Assuming that future studies confirm the alleged function of the lobed structures
observed by Prud’homme van Reine & Lokhorst, the existence of specialized
reproductive organs would seem to distinguish this species generically from Caulerpa,
in which numerous discharge papillae arise directly from the vegetative thallus.
Despite the presence of a wall separating the reproductive clusters from the remainder
of the plant, reproduction is probably a form of holocarpy in which most but not all
available cytoplasm participates. The contrast between the dense dark-green contents
of the developing reproductive clusters and the pale vegetative fronds from which
the cytoplasm has moved was described and shown photographically by Kemperman
& Stegenga (1983) in material from Costa Rica that they thought was representative
of a new species (C. biloba).
HALIMEDACEAE
HALIMEDA Lamouroux (nom. cons.)
Halimeda discoidea Decaisne (Ann. Sci. Nat., Bot., ser. 2, 18: 102, 1842).
Type in PC, said by author to be from Kamchatka, but true provenance unknown
(see Hillis, 1959, p. 353; Hillis-Colinvaux, 1980, p. 138).
ISRSE SPECIMENS STUDIED : Enteraia Island: 24.iii.1962 (E62/20146). Entedebir Island: Goliath Bay, 30.iii.1962 (E62/20194); Landing Bay, 21.x.1965 (ISRSE 65/
68
2314, 2316). Nocra Island: southern side opposite village, 16.iii.1962 (E62/20065).
community on rocky flat, ca. 20 cm below low water (ISRSE 65/0395, 2349),
10.x.1965, with many epiphytes (ISRSE 65/2315), 14.x.1965 (ISRSE 65/2317);
off Mangrove Bay, 14.x.1965, with many epiphytes (ISRSE 65/ 2318, 3022); Camping Bay, off Ras Pandion, 16.x.1965 (ISRSE 65/2319,2321).
DISTRIBUTION: Widespread in all warm seas.
COMMENTS : Plants collected in March and October were similar in size (to 12 cm
tall) and appearance, although there were more plants with young, small, dark-green
peripheral segments in spring than in autumn. As is usually true of Halimeda, our
plants were covered with many epiphytes, especially on the lower, older parts.
Halimeda opuntia (Linnaeus) Lamouroux (Hist. Polyp. Corall. Flex., p. 308, 1816).
Corallina opuntia Linnaeus (Syst. Nat., ed. 10, 1, p. 805, 1758).
Type from Jamaica (see Hillis 1959, p. 361).
ISRSE SPECIMENS STUDIED : Entedebir Island: Little Bay, 14.iii.1962, 1 m depth (E62/
20043); Goliath Bay, 30.iii.1962 (E62/20193). Nocra Island: south side opposite
village, 16.iii.1962, 1-2 m depth (E62/20066). Museri Island: Camping Bay, off
Ras Pandion, 16.x.1965, on rock flat 1 m below low water (ISRSE 65/0346), 1.5 m
below low water, with gametangia (ISRSE 65/2322); off Mangrove Bay, 14.x.1965,
with gametangia (ISRSE 65/2323, 2324); off Scopus Ridge, 12.x.1965 , with
gametangia (ISRSE 65/2385a).
DISTRIBUTION: Widespread in all warm waters.
COMMENTS : Our plants formed hemispherical clumps to 15 cm in diameter both in
spring and in autumn. They grew on rock from about mean sea level down to 2 m.
Reproductive plants were found only in the autumn. This species was more commonly
encountered than H. discoidea.
UDOTEACEAE
AVRAINVILLEA Decaisne
Avrainvillea amadelpha (Montagne) A. et E.S. Gepp (Trans. Linn. Soc. London,
Bot. 7: 178, 1908).
Udotea amadelpha Montagne (Ann. Sci. Nat., Bot., ser. 4, 7: 136, 1857).
Type from Agalega Islands, NE of Madagascar, leg. Le Duc, in PC (see OlsenStojkovich 1985, p. 36).
Caulerpa racemosa community on on rock flat ca. 20 cm below low water (ISRSE
65/0397, 2343), 10.x.1965 (ISRSE 65/2320); off Mangrove Bay, 14.x.1965 (ISRSE
65/3043). Romia Island: 29.iii.1962 (E62/20186).
DISTRIBUTION: Warm waters of western Indian Ocean and Red Sea.
69
COMMENTS: Plants collected in spring were 1.5-3.5 cm tall, thin, and green to greyish
green, whereas those collected in autumn were 3.5-5.5 cm tall, spongy, and brownish
green. The fact that the latter had many more epiphytes than the former implies that
new flabellae begin to grow from a perennial base in late winter and end their
growth in autumn.
The plants from Museri Island grew in a Chondrophycus papillosus community at
depths of about 20-40 cm during low water on a rock flat overlain with calcareous
sand and sloping gently seaward (Lipkin 1987, as Laurencia papillosa community).
They were associated with Valonia aegagropila and Halimeda opuntia as well as C.
papillosus and often covered with epiphytes, mostly Cladophora, Enteromorpha
and Polysiphonia .
*CHLORODESMIS Harvey et Bailey
*Chlorodesmis papenfussii Ducker (Phycologia 8: 17, figs. 2, 3, 1969).
Type from Romia Island, leg. G.F. Papenfuss, 29.iii.1962 (E62/20184), in UC.
ISRSE
SPECIMENS STUDIED:
as above.
DISTRIBUTION : Known only from the type locality.
COMMENTS: Ducker (1967, p. 156) tentatively assigned the type collection to Chlorodesmis baculifera (J. Agardh) Ducker, an Australian species, prior to recognizing
that it represented an undescribed species.
DASYCLADALES
POLYPHYSACEAE
ACETABULARIA Lamouroux (nom. cons.)
Most recent papers treat the small Acetabularia-like species, along with the larger A.
peniculus (R. Brown ex Turner) C. Agardh, as belonging to the genus Polyphysa
Lamarck (see Berger & Kaever 1992). Olsen et al. (1994), on the basis of molecular
analyses, concluded that Acetabularia parvula Solms-Laubach and A. exigua SolmsLaubach are generically distinct from other species of Acetabularia, including A.
peniculus, the type of Polyphysa. Silva et al. (1996, pp. 890-891) pointed out that
the Polyphysa clade recognized by Olsen et al. (1994). excludes the type of Polyphysa
and thus represents a genus that lacks a name. We follow Silva et al. (1996) in
recognizing only one genus until the taxonomic situation has been clarified.
Acetabularia parvula Solms-Laubach (Trans. Linn. Soc. London, Bot. 5: 29, pl.
2, figs. 3, 5, 1895).
Type from Makasar, Sulawesi (Celebes), Indonesia, leg. Weber-van Bosse; (Herb.
Weber-van Bosse no. 927) (see Valet, 1969, p. 622).
70
ISRSE SPECIMENS
20073).
STUDIED :
Cundabilu Island: western end of islet, 20.iii.1962 (E62/
COMMENTS : Plants collected in 1962 were 3-4 mm high and with a form typical of
this species. In about 10% the caps were incompletely developed. In about 50% the
caps were mature, with their rays filled with cysts. Mature caps had (12) 15-20 (21)
rays and measured (2.5) 3.75-4.75 (5.75) mm in diameter. Acetabularia stalks were
collected in 1965 on rocks off the northern coast of Desta Island (26.x.1965, ISRSE
65/3169g). These agreed with those of A. parvula, but identification remains uncertain
because no caps were present. It may be that A. parvula in the southern Red Sea
initiates caps in winter, reaching reproductive maturity in spring.
Previous reports of this alga from the Red Sea, except that of Lundberg & Lipkin
(1979), were made under the name Acetabularia moebii Solms Laubach, a synonym
of A. parvula (see Valet 1969, p. 622).
ANTHOPHYTA
HELOBIAE
CYMODOCEACEAE
THALASSODENDRON den Hartog
Thalassodendron ciliatum (Forsskål) den Hartog (Verh. Kon. Ned. Akad. Wetensch.,
Afd. Natuurk., Tweede Sect. 59(1): 188, 1970).
Zostera ciliata Forsskål (Fl. Aegypt.-Arab., p. 157, 1775).
Type from Al Mukha, Yemen, southern Red Sea, leg. P. Forsskål, in BM (see den
Hartog 1970, p. 183).
ISRSE
SPECIMENS STUDIED:
DISTRIBUTION: Warm waters of Indo-west Pacific.
COMMENTS: This plant is very common in the northern Red Sea (Lipkin 1977b),
much less common in the central and southern parts.
HYDROCHARITACEAE
HALOPHILA Du Petit Thouars
Halophila ovalis (R. Brown) J.D. Hooker (Fl. Tasman. 2, p. 45, 1858).
Caulinia ovalis R. Brown (Prodr., p. 339, 1810).
Type from tropical northeastern Australia, locality not specified; leg. R. Brown (5816),
in BM (see Dandy 1971).
ISRSE SPECIMENS
(E62/20041).
STUDIED :
Entedebir Island: Goliath Bay, 14.iii.1962, 1 m depth
71
DISTRIBUTION : Widespread in warm Indo-west Pacific waters.
COMMENTS: Our plants were sterile.
This plant is rather common on Sinai coasts in the northern Red Sea (Lipkin 1977b).
Halophila stipulacea (Forsskål) Ascherson (Sitzungsber. Ges. Naturf. Freunde Berlin,
1867: 3, 1867).
Zostera stipulacea Forsskål (Fl. Aegypt.-Arab., p. 158, 1775).
Type from Al Mukha, Yemen, southern Red Sea, leg. P. Forsskål, presumably in C.
ISRSE SPECIMENS
20107).
STUDIED :
Entedebir Island: Goliath Bay, 14.iii.1962, sterile, (E62/
DISTRIBUTION : Warm waters of the western Indian Ocean and Red Sea. This species
migrated about 130 years ago into the Mediterranean, where it presently occurs
mostly in the eastern basin (see Lipkin 1975a, b).
COMMENTS: This is the most common seagrass on the Sinai coasts in the northern
Red Sea (Lipkin 1977b).
THALASSIA Banks ex König
Thalassia hemprichii (Ehrenberg) Ascherson (Mitth. Justus Perthes’ Geogr. Anst.
17: 242, 1871).
Schizotheca hemprichii Ehrenberg (Abh. Königl. Akad. Wiss. Berlin, 1832(1): 429, 1834).
Type from Mesewa (Massaua), Ethiopia, southern Red Sea, leg. C.G. Ehrenberg, in
BM (also ex K), L, P (den Hartog 1970, p. 234).
ISRSE SPECIMENS STUDIED : Enteraia Island: 24.iii.1962, (E62/20145). Museri Island: in front of Scopus Ridge, 8.x.1965, in Caulerpa racemosa-Thalassia hemprichii
community, with many epiphytes (ISRSE 65/0304), 9.x.1965 (ISRSE 65/0307),
10.x.1965 (ISRSE 65/2379).
DISTRIBUTION : Widespread in warm Indo-west Pacific waters.
COMMENTS: This seagrass, occurring in the entire Red Sea, seems to be more common
in its southern part.
Discussion and summary
Whereas the benthic marine flora of the northern Red Sea has been studied intensively
both in earlier years (see Papenfuss 1968a) and in the last three decades (e.g., Lipkin
1972a, 1972b, 1977a, 1977b, 1977c, 1979, in Mergner and Schuhmacher 1974;
Mergner and Svoboda 1977; Aleem 1978a, 1978b, 1978c, 1980, 1981, 1984, 1990;
Lundberg & Lipkin 1979; Khoja 1987; Natour et al. 1979a, 1979b), the southern
72
Red Sea has been neglected. In the 38-year period since the first ISRSE visited that
area in 1962, only two papers (Klausewitz 1967; Simonsen 1968) have been published
containing information about its marine flora, and that information is minimal. The
results of the two Israeli expeditions, therefore, expand considerably our knowledge
of the marine flora of this poorly known part of the Red Sea. No fewer than 73 of
the species collected during the expeditions (about 56%) were not previously known
from the southern part of this sea (see Table 3). Furthermore, among the 131 identified
species collected by the two Israeli expeditions, 47 (ca. 36%) are new records for the
entire Red Sea (Table 2). Three species known only from the Red Sea are new to
science: Cladophoropsis papenfussii Ducker, Pterocladiella microscopica Lipkin et
Papenfuss, and Turbinaria papenfussii W.R. Taylor. This unexpectedly high percentage
of new records from the Red Sea implies that our knowledge of its marine flora is
still far from complete even after more than two hundred years of collecting.
A species of Hypnea, widely distributed in warm waters but lacking a legitimate
name, is described as a new species, Hypnea anastomosans Papenfuss, Lipkin et
Silva. Sargassum steinitzii is proposed as a new name for S. tenue J. Agardh. The
combination Gelidiopsis capitata is made for Acrocarpus capitatus Kützing, which
we regard as being distinct from G. intricata (C. Agardh) Vickers. The combination
Chondrophycus glanduliferus is made for Chondria glandulifera Kützing.
Genera reported here that were not known from the Red Sea prior to the collections
of the two Israeli expeditions are the following: Bostrychia, Brachytrichia, Caloglossa,
Caulerpella (recently segregated from Caulerpa), Chamaecalyx, Chlorodesmis,
Coelothrix, Gelidiopsis, Gracilariophila , Hypneocolax, Pterocladiella (recently
segregated from Pterocladia), Valoniopsis, and Wurdemannia. Subsequent to the
first expedition, Brachytrichia was reported from the northernmost Red Sea at the
Gulf of Elat (Safriel & Lipkin 1964) and Bostrychia much later from the same area
(Natour et al. 1979b).
Mangroves are common in the Dahlak Archipelago and several sites were visited
(Sheikh Sa’id I., Museri I., Desta I., Seil Anber, Romia I.). However, although a
special search was made for members of the Bostrychietum on mangrove
pneumatophores, only two were found (Bostrychia simpliciuscula and Caloglossa
leprieurii) and only at the highly sheltered locality of Melita Bay on the Eritrean
coast in the Gulf of Zula (Lipkin 1987). They were growing with two blue-greens
(Calothrix pilosa and Brachytrichia lloydii).
The bulk of the collections made during the expeditions came from the central part
of the archipelago, where the two base camp islands are located (Table 3). This is an
area of shallow water, usually with calm seas and little wave action, and thus subject
to exaggerated fluctuations in environmental conditions in response to changes in
the weather. Sea water may reach extreme values of temperature, salinity, O2 and
CO2 concentrations, and other factors. These fluctuations and extreme values render
the environment hostile to many marine algae. The impoverishment of the marine
flora in the central part of the archipelago was underscored by a short visit to Romia
I., an outer island (Fig. 1) where environmental fluctuations and extremes are
ameliorated by the swells that come from the main channel of the Red Sea and wash
73
the island’s shores. A relatively large number of species (29) was found here, including
10 that were not seen elsewhere in the archipelago (Table 3). A similar brief visit
was paid to Harmil I., another outer island, where 16 species were collected, including
one that was not seen elsehere in the archipelago. The lower number of species on
Harmil I. compared with Romia I. probably may be explained by the fact that the
side facing the mainland, where the collections were made, is fairly sheltered from
the swell. Brief visits to inner islands yielded far fewer species: Enteraia I., nine;
Nocra I., six; Desta I., five; Um Aabak, Sheikh Sa’id I., and Dahlak Kebir I., two
each. The amount of time devoted to collecting complicates comparisons. Three
days were spent on Cundabilu I., an inner island which yielded 16 species, including
four not seen elsewhere in the archipelago. The largest number of species was found
on the base camp islands, where many different habitats were visited and much more
time was devoted to collecting. Entedebir I. yielded 78, including 32 collected only
here, while Museri I. yielded 70, including 23 not collected elsewhere. When the
amount of time devoted to collecting is taken into consideration, the diversity and
uniqueness of the algal flora of Romia I. is striking despite the low number of
species compared with the base camp islands.
Although well within the tropics (Fig. 1), the Dahlak Archipelago supports a marine
flora that is subject to marked seasonal fluctuations (Lipkin 1987). Of the 131 species
identified from the two expeditions, only 46 (35%) were found in both spring and
autumn (Table 4). This number may not be accurate, however, since some species
found only in one season may have come from habitats and communities not visited
during the alternate season. These species include Sargassum spp., Boergesenia
forbesii, Valoniopsis pachynema, Codium arabicum, Galaxaura rugosa, and Gelidiella
spp. Of the 46 species that were found in both spring and autumn, seven reproduced
during both seasons, among them the highly opportunistic Acrochaetium gracile,
Sphacelar ia rigidula , and Padina boergesenii. In three of the seven species,
reproductive phases were seasonal. In Botryocladia skottsbergii, for example, cystocarpic plants were found only in spring, tetrasporangial plants only in autumn. Six
species were found reproductive only in spring: Cystoseira myrica, Hormophysa
cuneiformis, Jania adhaerens, Gelidiella lubrica, Sarconema filiforme, and Martensia
elegans. Four species were found reproductive only in autumn: Halimeda opuntia,
Sphacelaria tribuloides, Ceramium flaccidum, and Chondrophycus papillosus. It is
of note that Halimeda opuntia reproduces in this tropical region at the same season
as H. tuna in the temperate eastern Mediterranean. Most of the 46 species (26 or
57%) were not found reproductive in either season. Nonetheless, many were
represented by different stages of growth in the two seasons.
Of the 52 species found only in spring, 16 (31%) had reproductive thalli, while
among the 33 species found only in autumn, 16 (48%) had reproductive thalli. The
occurrence of Colpomenia sinuosa, Hydroclathrus clathratus, Dictyota spp., and
Ulva spp. only in the spring agrees with their seasonality in the temperate northern
Red Sea and eastern Mediterranean (Lipkin 1987). While most species found only in
autumn have perennial crustose bases that are difficult to detect in spring, but some,
like Hypnea anastomosans , H. valentiae, Coelothrix irregularis, Chondria spp., and
Chondrophycus parvipapillatus, may be true summer-autumn species.
74
Acknowledgments
The late Prof. G.F. Papenfuss played a major role in the development of this paper – as collector,
identifier, describer, and organizer. At his death, he left us a manuscript that we have updated and
rewritten, thus introducing opinions that may or may not have met his approval.
We are indebted to the late Prof. H. Steinitz and the late Dr. O. Oren, leaders of ISRSE 1962, and the
late Prof. C. Lewinsohn and Prof. L. Fishelson, leaders of ISRSE 1965, for their help and encouragement during work in the field and thereafter. The expeditions were financially supported by
the Government of Israel, Hebrew University, Tel Aviv University, the U.S. National Science
Foundation, and the U.S. Office of Naval Research (Contract no. 62558-4556 to Tel Aviv University).
Study of the collections at Berkeley was supported by the U.S. National Science Foundation (Grant
BG 1656). We are grateful for the assistance of several specialists. Prof. Young Meng Chiang, in
collaboration with Prof. G.F. Papenfuss, identified much of the material from the first expedition. Dr.
S.C. Ducker identified the E62/ Chlorodesmis (describing a new species). Dr. L. Hillis-Colinvaux
ldentified the E62/ Halimeda. The late Prof. W.R. Taylor identified the Turbinaria from the first
expedition (describing a new species) as well as Caulerpa from both expeditions.
Dr. Richard L. Moe has been generous with his time and expertise in helping us manage the electronic
files.
Table 2: Number of genera and species and of new records of marine algae and seagrasses known
from the Red Sea previous to the Israel South Red Sea Expeditions of 1962 and of new records
resulting from the expeditions.
75
Table 3: Distribution of marine algae and seagrasses in the Dahlak Archipelago (spring of 1962 and
autumn of 1965).
76
77
Table 3: continuation
78
Table 4: Occurrence of marine algae and seagrasses in the Dahlak Archipelago in spring (1962) and
autumn (1965) with indication of reproductive plants.
79
Table 4: continuation
80
81
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Received 21 March 2001, acepted in revised form 18 December 2001.
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Marine algae and seagrasses of the Dahlak Archipelago, southern

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