Pacific Science (1973), Vol. 27, No.1, p. 37-43 Printed in Great Britain

Marine Algae of the Smithsonian-Bredin Expedition to the Society and Tuamotu Islands I

WM. RANDOLPH TAYLOR2

ABSTRACT: Short lists of the marine algae collected by the Smithsonian-Bredin Expedition to the Society and Tuamotu Islands are given. A detailed discussion and description of Giffordia indica (Sond.) Papenf. & Chihara is given.

THE UNITED STATES NATIONAL MUSEUM has in Ch/orodesmis and Bood/ea. I included T!Jrbinaria well earned its success in supplementing its records in reviewing that genus (Taylor, 1964: budget-financed field study expeditions with 475). Setchell (1926: 61), also dealing with support from private sponsors, whereby great , gives us the chief, though surely very enrichment of our national collections has re­ incomplete, source of information on the algae sulted. One ofthese was the Smithsonian-Bredin of the area and, like myself, he found difficulty Expedition of April and May 1957 to the Tua­ with the Weber localities. General features of motu Archipelago and the the vegetation on Raroia in the are (specifically the lIes sous Ie Vent and Moorea), alluded to by Doty and Morrison (1954) and by French mandates just north of the Tropic of Guilcher, Denizot, and Berthois (1966: 851) as Capricorn in the southern Pacific Ocean. The being present on Mopelia Atoll in the Society generous sponsor, J. Bruce Bredin of Wilming­ Islands, but they deal primarily with the crus­ ton, Delaware has also supported work in the tose lithothamnia of the reef, which are entirely Atlantic, such as his 1960 expedition to Yucatan lacking from the Smithsonian-Bredin collec­ (Taylor, 1972: 34). tions. There are in fact, numerous papers men­ The marine algae collected were few, but not tioning algal components of the reef structure, without interest, for there is singularly little but they do not have comprehensive taxonomic known of the marine vegetation of either of coverage. these Pacific island groups. They figure in the The writer is particularly grateful to Dr. report of "Captain Beechey's Voyage...in Waldo L. Schmitt and the United States Na­ H.M.S. Blossom" (Hooker and Arnott, 1841: tional Museum for submitting this material for 77), in a general review by von Martens (1866: study and report. Dr. W. F. Pmd'homme van 1) and in Svedelius's important phytogeogra­ Reine loaned from the Rijksherbarium, Leiden, phic paper (1924: 1). Mme. Weber-van Bosse's the uniquely valuable specimens from Mme. Tahiti paper(1910: 1) should bear directly, since Weber-van Bosse's herbarium and others in Tahiti is very near Moorea, but only one of the their collection. Professor George F. Papenfuss localities, Tearei (as Tearia), suggests Tahiti, and Dr. Paul Silva gave help from the Univer­ while two, the village Rikitea and the lIe Tara­ sity of California. Many others contributed in vai, appear on maps of the far less direct fashion. away at the southeast end of the Tuamotu Archipelago. Only Cau/erpa racemosa appears both in her short list and in this one. Butters SOCIETY ISLANDS (1911: 161) mentions two Liagoras from Tahiti, The expedition with which this paper is con­ and Brand (1911a: 138; 1911b: 611) cites species cerned made algal collections in two regions in the group. At Moorea, 10 miles northwest of 1 This study with others was supported by grant 3186 the better-known Tahiti, particularly on the from the National Science Foundation. Manuscript reef at Pointe Haure (or Haum) on the north­ received 10 May 1972. 2 Department of Botany, University of Michigan, east corner of the island, on Toatane Reef west Ann Arbor, Michigan 48104. of the entrance to Paopao Bay (Cook Bay), and 37 38 PACIFIC SCIENCE, Volume 27, January 1973 a little to the east, but west of Irihonu Pass at Rhodophyceae Maharepa Bay on 8-10 May 1957. Other collec­ Gelidiella acerosa (Forssk.) Borg. tions had been made at about 100 miles Island, Motu , Station 62-57. to the west in the Leeward Group, specifically Jania pumila Lamx., prox. Moorea Island, on the northern island Huahine Nui, north of Pointe Haure, on Turbinaria, Station 105-57. Baie Fare between Pointe Teffaao (or Teffaar) Jania rllbens Lamx. , Motu and Pointe Teopape (or Teopapa) (2 May), and Tapu, Station 62-57. Huahine Nui, Passe along the outer reef south of Passe Avamoa Avamoa, Station 95-57. near the bay and village of Fare (3 May). Still Amphiroa rigida Lamx. Huahine Nui, Passe earlier, good samples were retained from Avamoa, Station 95-57. There were very few gatherings made on Bora Bora Island, taken instances of femur-ended segments in this from the lagoon side of the ocean reef west of material. Motu Tapu (25 April). IfJ'Pnea pannosa J. Ag., prox. Bora Bora Island, Motu Tapu, Station 62-57. Chloropqyceae Dictyosphaeria cavernosa (Forssk.) Borg. Bora Bora Island, Motu Tapu, Station 62-57. TUAMOTU ISLANDS Boodlea composita (Harv.) Brand. Bora Bora Island, Motu Tapu, Station 62-57, a poor frag­ This archipelago extends in a southeasterly mentary sample. Huahine Nui, Passe Avamoa, direction for a great distance from those islands Station 95-57. north of the Society group where this expedi­ Caulerpa pickeringii Harv. & Bail. Moorea tion made its algal collections. Island Island, Maharepa Bay, Station 116-57. This was is perhaps the closest, and the algae were col­ a generous collection of a plant not commonly lected north of the pier in Temao Harbor on reported. Its habit is quite different from that of 16 April 1957. Tikahau Island is just a little to other Caulerpas. It forms close colonies with the east. The material available came from a sparingly branched erect axes a very few centi­ depth of 15 m on a patch of reef in the lagoon meters tall. These resemble those of a small Co­ south ofthe deep water pass, at Mareva anchor­ dium, having the same texture, but Codiums do age on 12 April. A third sample came from the not commonly form dense plant carpets. The lagoon shore of Maiai Islet at Tikahau on 13 ramelli are so very densely crowded that the April. surface appears quite continuous, but on dissec­ Ulva rigida C. Ag. Makatea Island, Temao tion the typical detailed structure with trabecu­ Harbor, Station 37-57. lae characteristic of the genus is evident. Microdictyon okamurai Setch. Tikahau, Maiai Caulerpa racemosa (Forssk.) J. Ag. Bora Bora Islet, Station 26-57. Island, Motu Tapu, Station 62-57, in a dwarf Cladophoropsis sundanmsis Reinb. Makatea Is­ form. land, Temao Harbor, Station 37-57. Chlorodesmis comosa Harv. & Bail. Bora Bora Halimeda opuntia (L.) Lamx., f. elongata (Bart.) Island, Motu Tapu, Station 62-57. Moorea Bart. Tikahau, Mareva anchorage, Station 18­ Island, Pointe Haure, Station 105-57. 57. Halimeda lacunalis W. R. Taylor. Tikahau, Phaeophyceae Mareva anchorage, Station 18-57. Ectocarpus breviarticulatus Ag. Moorea J. Phaeophyceae Island, Toatane Reef, Station 103-57. Huahine Nui Island north of Baie Fare, Station 90-57. Ectocarpus breviarticulatus J. Ag. Makatea The plants from Huahine Island bore plurilocu­ Island, Temao Harbor, Station 37-57. lar gametangia in abundance. Giffordia indica (Sond.) Papenf. & Chihara. Turbinaria condensata Sond. Bora Bora Island, Under one name or another Ectocarpus indicus Motu Tapu, Station 62-57. Moorea Island, Sonder (1854: 3, from Bima Bay on the north Pointe Haure, Station 105-57. coast of Sumbawa Island near the eastern end) Algae of the Smithsonian-Bredin Expeclition-TAYLOR 39

is recognized to be a widespread and important Since I have recognized E. duchassaingianus (Tay­ alga of the western Pacific and Indian oceans. lor, 1960: 207 as Giffordia duchassaingiana) from Unfortunately, it seems generally to have been numerous Atlantic and Caribbean stations, I reported very casually, without thorough ac­ have been able to make measurements of fila­ companying description or figures and has been ments and of plurilocular gametangia from associated in synonymy with various other en­ these and from the several Pacific stations repre­ tities. Askenasy (1888: 19,20, pI. 5, fig. 2, 10) sented in the Michigan collections. The average quoted from the original description where the maximum filament diameter for the Atlantic­ gametangia are said to be secund, which cer­ Caribbean material I find to be 29 pm, thelargest tainly has not generally been an emphasized gametangia to average 23.5 pm in diameter and feature, but the statement may have affected 156 pm in length, with an extreme of 248 pm. Setchell's interpretation. The figure ofE. indicus The gametangia do not, then, often reach ex­ which Mme. Weber-van Bosse (1913: 130) pro­ treme lengths. While minor differences seem to vided from Sonder material shows plurilocular appear in some specimens they do not seem gametangia erect on the ascending filaments, constant, and so it appears that on the present not secund near the bases of lateral branchlets, evidence I must accept Giffordia indica (Sonder) and, admitting that the plant she figured was Papenfuss & Chihara (Papenfuss 1968: 30) over very small, the branching is erect, nothing like E. duchassaingianus. the figures given by Setchell (1924: 169). B0r­ This leads to a consideration ofthe Tuamotu gesen (1937: 4) suggested that E. indicus was material brought back by this expedition. It was only a form ofE. duchassaingianus Grunow. Still generous in quantity, but not well-enough pre­ later (1941: 16) he discussed E. indicus at length served to show the chromatophores. The lower and in this case reduced E. duchassaingianus to parts were heavily infiltrated with calcareous synonymy under E. indicus, which Dawson mud and probably little effort had been made to (1956: 43) accepted. However, his figures re­ get any creeping parts or initial organs of at­ semble E. mitchellae somewhat, and my own tachment (Fig. 1-11) to the underlying coral (Taylor, 1928, pI. 14, fig. 11) of E. duchassain­ rock, so these were very scarce and of limited gianus are not entirely convincing, when it is value. Nevertheless the material justified ex­ considered synonymous with E. indicus. Simi­ tended study and comparison with other Pacific larly Okamura's figure (1936: 137) suggests E. and Atlantic materials available. mitchellae. 3 Trono (1969: 25) reported very small A condensed description based strictly on the plants somewhat similar to those of Mme. Tuamotu material would be as follows: G. Weber-van Bosse from the Caroline Islands, but indica (Sonder) Papenfuss & Chihara: Plants with unexpectedly short plurilocular gametan­ crowded colonial, 2-5 cm tall, the bases some­ gia. Womersley and Bailey (1970: 288) on ex­ what decumbent and entangled, the erect fila­ amination of Sonder material found that the ments loosely and repeatedly alternately bran­ figure given by Setchell did not at all agree with ched, for the most part 18-26 pm diameter, the the type, but rather with Giffordia mitchellae cells cylindrical, 1.25-2.50diameters long, some­ (Harv.) Hamel, which is quite my impression what more slender toward the tips where 10­ from Setchell's figure and from material I have 12 pm diameter, 30-70 pm long; extensive re­ seen. Womersley and Bailey refer indica to gions, apical andintercalary, including axes with Hamel's genus FeldJJlannia, a genus for which I branchlets, composed of short cells distinctly see no particular need. Like B0rgesen, Trono, turgid and with very dense contents, not dif­ and Papenfuss, Womersley and Bailey (1969: fering greatly from adjacent regions in di­ 437) reduced duchassaingianus to synonymy under ameter, but the cells only half as long, or barely indica without giving many supporting data. as long, as broad (Fig. 1-7-10); plurilocular gametangia scattered along the filaments (Fig. 1-1-3), ascending to erect, sessile, or rarely on 3 In Kuckuck's reJiquae (1963) he is seemingly clear as to E. duchassaingianus but he had not come to 1-2-stalk cells, occasionally terminating a a final decision on its relations with E. ;,tdicus and branchlet (Fig. 1-4-5), subcylindrical or a E. amicoruJ11 Harv. little clavate, blunt at the tips, 11-31 pm di- 40 PACIFIC SCIENCE, Volume 27, January 1973

\ 7

8

; : 3 6 11 ~ Algae of the Smithsonian-Bredin Expedition-TAYLOR 41 ameter, 75-260 j1,m long, with loculi about 7.5­ simply minute ones as inferred by Setchell. The 8.0 j1,m diameter. TUAMOTU ISLANDS, Tikahau largest filaments in Leiden specimens reached Atoll, Maiai Island, along the shore on the 34-39 j1,m in diameter. The plurilocular game­ lagoon side. Collected Smithsonian-Bredin Ex­ tangia generally ranged in diameter from 21­ pedition, Station 26-57, 13 April 1957. 36 j1,m and in length to 200-300 j1,m, but in the Sonder's diagnosis (1854: 3; Anon., 1855: material of the type collection from the island 44) does not provide detailed measurements of of Sumbawa, near Pulu Kambing, in Bima Bay the plants. Askenasy does, and both in filament (Zollinger no. 3428), it commonly exceeded diameter and plurilocular gametangial dimen­ 300 j1,m and actually reached 403 j1,m. The sions the figures are considerably below the Tuamotu material is, then, on all accounts of average. B0rgesen (1914: 159, under E. duchas­ quite conservative dimensions. One should saingianus) cites dimensions comparable with note that in one Leiden specimen from the those from the Pacific, as does Jaasund (1969: Celebes the gametangia seemed mature, but 256) from Tanzania and Nasr (1947: 64) from were only one-two cells and to 16 j1,m in the Red Sea. I have mentioned the measure­ diameter, while to 170 j1,m in length, which ments I have averaged from Bermudian and suggests dimorphism among the plurilocular West Indian Atlantic material, but did not note plants. unilocular sporangia there myself. B0rgesen The largest plurilocular gametangia seem to did, and reports them as reaching 70 j1,m di­ occur in the upper central parts of the tufts: ameter and 110 j1,m in length. I did see them, those formed eatly near the base matured while together with pluriloculars, in material from relatively small, whereas those near the filament Hawaii (collector Doty, no. 19280), where they tips were immature. A peculiarity found in the were smaller, only 26 j1,m diameter and 85 j1,m Tuamotu material concerned these plurilocular in length. The materials available to me from gametangia. In numerous instances near the the Marshall Islands, Caroline Islands, and bases of the filament tufts these proliferated Singapore in structural dimensions are within freely (Fig. 1-6), but the individual outgrowths the range I find from the Tuamotus. Mme. were not seen to exceed three-five cells in Weber-van Bosse (1913: 130) reports similar length. Their propagative potential in a soft, filament diameters, but longer (to 300 j1,m) sandy, or muddy environment is, however, pluriloculars, while her measurements of uni­ evident. locular sporangia agree with those of B0rgesen, The matter of the reputed growth zones calls rather than what I found in Hawaiian material. for some comment. In most ectocarpi familiar On examining a small portion of her Makassar to me these are of modest extent, usually basal material (no. 936,303-77) I found the larger uni­ to terminal "hairs." In G. indica they are very locular sporangia often to be a little obovate, long, even several scores of cells, and there and to reach 39-47 j1,m in diameter, 85-125 j1,m seems to be no distinctive point in the series in length, but none to approach the diameter of where particularly short cells initiate activity. 60 j1,m which she cites. The cells are half as long as broad or subequal, Opportunity to examine E. indicus material in in some collections very turgid and very dense the Rijksherbarium, Leiden, including that of content (as in the Tuamotu material), but in formerly in the possession of Mme. Anna others less distinctive. These series may be ter­ Weber-van Bosse, served importantly to extend minal (Fig. 1-7-9), where they are relatively my familiarity with the species. Some clearly short, or intercalary and more often long, in­ showed the short entangled basal holdfast fila­ cluding lateral branchlets themselves similarly ments. A Sonder type specimen included was distinctive (Fig. 1-10). Instances were seen well over 4 cm tall, indicating that she had at where small branch systems of many divisions least one well-developed plant to study, not were completely in this state.

FIG. 1. GifJordia indica. 1-3, Plurilocular gametangia lateral on leading axes; 4, 5, plurilocular gametangia terminal on branchlets; 6, plurilocular gametangium with the loculi germinating to filaments in situ instead of producing swarmers; 7-10, terminal and intercalary filament areas considered to be growth zones; 11, an initial hapteron. 42 PACIFIC SCIENCE, Volume 27, January 1973

Chnoospora minima (Her.) Papenfuss. Makatea HOOKER, W. J., and G. A. W. ARNOTT. 1841. Island, Temao Harbor, Station 37-57. The botany of Captain Beechey's voyage; ... Turbinaria condensata Sond. Makatea Island, in His Majesty's ship Blossom...in the years Temao Harbor, Station 37-57. 1825,26,27, and 28. ii+485 p., 99 pI. Treut­ tel and Wurtz, London. Rhodop!?yceae JAASUND, E. 1969. Marine algae in Tanzania I. Bot. Marina 12 (1-4): 255-274, 10 textfig. Pterocladia nana Okam.,prox. Makatea Island, KUCKUCK,P. T.,edit.P. KORNMANN.1963. Ecto­ Temao Harbor, Station 37-57. carpaceen-Studien VIII; Einige Arten aus warmen Meeren. Helgoland. wiss. Meere­ sunters. 8(4): 361-382, 10 textfig. LITERATURE CITED MARTENS, G. VON. 1866. Die Tange. In: Die ANONYMOUS. 1855. Florae Aechipelagi Indici, preussische Expedition nach Ost-Asien, nach species novae. Natuurk. Tijdschr. Ned.-Ind. amtlichen Quellen, Bot. Theil. P. 1-152, pI. 8: 44-52. 1-8. ASKENASY, E. 1888. Algen, mit Unterstutzung NASR, A. H. 1947. Synopsis ofthe marine algae der Herren E. Bornet, A. Grunow, P. Hariot, of the Egyptian Red Sea coast. Bull. Fac. Sci. N. Moebius, O. Nordstedt. Forschungsreise Egypt Univ. 26: 1-155,25 textfig., 14 pI. S.M.S. "Gazelle," IV Theil, Botanik. P. 1­ OKAMURA, K. 1936. Nippon kaiso-shi [Marine 58, pI. 1-12. algal flora of Japan]. Published by the author, B0RGESEN, F. 1914. The marine algae of the Tokyo. 964 p., 427 fig. Danish West Indies. Dansk bot. Ark. 2 (2): PAPENFUSS, G. F. 1968. A history, catalogue 1-66+2,44 fig. and bibliography of Red Sea benthic algae. ---. 1937. Contribution to a south Indian Israel ]. Bot. 17 (1, 2): 1-118. marine flora. ]. Indian Bot. Soc. 16 (1,2): 1­ SETCHELL, W. A. 1924. American Samoa: Part 56, 33 textfig., 1 pI. I, Vegetation ofTutuila Island. Publ. Carneg. ---. 1941. Some marine algae from Mauri­ Instn. 341. Vol. 20: 1-188,20 pI. tius, II. Phaeophyceae. Det K. Danske Vi­ ---. 1926. Tahitian algae and Tahitian densk. Selsk., BioI. Medd. 16 (3): 1-81, 24 spermatophytes collected by W. A. Setchell, textfig., 8 pI. C. B. Setchell and H. E. Parks. Tahitian algae. BRAND, F. 1911a. Uber einige neue Grunalgen Univ. Calif. Publ. Bot. 12 (5): 61-143, pI. 7­ aus Neuseeland und Tahiti. Ber. dtsch. bot. 22. Ges. 29 (3): 138-145, pI. 7. SONDER, O. G. 1854. Algae, p. 1-4. In: H. Zol­ ---. 1911b. Uber die Siphoneengattungen linger, Systematisches Verzeichnis der im ChlorodestJlis. Ber. dtsch. bot. Ges. 29 (9): Indischen Archipel in den Jahren 1842-1848 606-611, 7 textfig. gesammelten sowie in der aus Japan emp­ BUTTERS, F. K. 1911. Notes on the species of fangenen Pflanzen. 2 Hefte. xii + 160 p., 1 pI. Liagora and Galaxatlra of the central Pacific. Zurich. Minn. Bot. Stud. 4 (2): 161-184, pI. 24. SVEDELIUS, N. 1924. On the discontinuous geo­ DAWSON, E. Y. 1956. Some marine algae of the graphical distribution of some tropical and southern Marshall Islands. Pacif. Sci. 10 (1): subtropical marine algae. Ark. Bot. 19 (3): 1­ 25-66, 66 textfig. 70, 14 fig. DoTY, M. S., and J. P. E. MORRISON. 1954. TAYLOR, W. R. 1928. The marine algae of Interrelationships of the organisms on Florida, with special reference to the Dry Raroia aside from man. Atoll Res. Bull. 35: Tortugas. Publ. Carneg. Instn. 379: Papers 1-61. from the Tortugas Lab. 25: v+219, 3 text­ GUILCHER, A., M. DENIZOT, and L. BERTHOIS. fig., 37 pI. 1966. Sur la constitution de la crete externe ---. 1960. Marine Algae of the eastern de l'Atoll de Mopelia ou Maupihaa (tIes de la tropical and subtropical coasts of the Ameri­ Societe) et quelques autres recifs voisins. cas. Univ. Michigan Press, Ann Arbor. xi+ Cah. Oceanogr. 18: 851-856, 3 pI. 870 p., 14 photographs in text, 80 pI. Algae of the Smithsonian-Bredin Expedition-TAYLOR 43

---. 1964. The genus Turbinaria in eastern Caulerpa de la Nouvelle-Hollande. Ann. Inst. seas. J. Linn. Soc. (Bot.) 58 (375): 475-490, oceanogr. Monaco 21 (1): 1-8, pI. 1,2. 8 textfig., 3 pI. ---. 1913. Liste des algues du Siboga. I, ---. 1972. Marine algae of the Smithsonian­ Myxophyceae, Chlorophyceae, Phaeophyceae Bredin Expedition to Yucatan-1960. Bull. avec Ie concours de M. Th. Reinbold. Siboga Mar. Sci. 22 (1): 34-44. Exped. 59a: 1-186, 52 textfig., 5 pI. TRoNa, G. c., JR. 1969. The marine benthic WOMERSLEY, H. B. S., and A, BAILEY.. 1969. algae of the Caroline Islands, II. Phaeophyta The marine algae ofthe Solomon Islands and and Rhodophyta. Micronesica 5 (1): 25-119, their place in biotic reefs. Phil. Trans., B, 11 pI. 255: 433-442. WEBER-VAN BOSSE, A. 1910. Note sur les ---. 1970. Marine algae of the Solomon Caulerpa de l'Ile Tahiti et sur un nouveau Islands. Phil. Trans., B,259 (830): 257-352, 10 textfig., pI. 24-27.