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SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES • NUMBER 17

The Genus Caulerpa
from Central Visayas,
Philippines

Ernani G. Meñez
and Hilconida P. Calumpong

SMITHSONIAN INSTITUTION PRESS
City of Washington
1982

ABSTRACT

Meñez, Ernani G., and Hilconida P. Calumpong. The Genus Caulerpa from Central Visayas, Philippines. Smithsonian Contributions to the Marine Sciences, number 17, 21 pages, 2 figures, 3 plates, 1982.—This taxonomic study of Caulerpa shows 20 taxa occurring in Central Visayas, Philipines, including Caulerpa reyesii, new species. Of these, three are newly reported from the Philippines and seven taxa are new records from Central Visayas.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution’s annual report, Smithsonian Year. SERIES COVER DESIGN: Seascape along the Atlantic coast of eastern North America.

Library of Congress Cataloging in Publication Data

Menez, Ernani G.

The genus Caulerpa from central Visayas, Philippines

(Smithsonian contributions to the marine sciences ; no. 17)

Bibliography: p.

1. Caulerpa—Classification. 2. Botany—Philippines—Visayan Islands—Classification. I. Calumpong, Hilconida P. II. Title III. Series.

QK569.C37M46    1982    589.4′7    82–5976    AACR2

The Genus Caulerpa
from Central Visayas,
Philippines

Ernani G Meñez
and Hilconida P. Calumpong

Ernani G. Meñez, Smithsonian Oceanographic Sorting Center, Smithsonian Institution, Washington, D.C. 20560. Hilconida P. Calumpong, Department of Biology, Silliman University, Dumaguete City 6501, Philippines.

Introduction

During the summers of 1978 and 1979, Smithsonian Institution conducted biological collecting expeditions in Central Visayas, Philippines, in order to collect marine plant and animal organisms for purposes of systematic, ecological, and distributional studies. Included in the plant collection was a large number of Caulerpa species, which forms the basis of the present study, taken from different parts of this region.

Fifty-three taxa of Caulerpa have been previously reported from the Philippines. Of these, 14 occurred in Central Visayas. These were Caulerpa brachypus, C. cupressoides, C. fastigiata, C. lentillifera, C. peltata, C. racemosa, C. racemosa var. laetevirens, C. racemosa var. macrophysa, C. serrulata, C. serrulata var. typica f. lata, C. sertularioides, C. taxifolia, C. urvilliana, and C. verticillata. A few of these taxa were not found in our collections.

The principal collecting areas are in Central Visayas, in the middle of the Philippine Archipelago (Figure 1). The area consists of the islands of Negros, Cebu, Bohol, Siquijor, and their satellite islets. Except for the southwestern portion of Negros Island, which is exposed to a major body of water, the Sulu Sea, the rest of Central Visayas is sheltered. Because it is practically closed off by other surrounding islands, the greater portion of this area is uniquely protected from intense wave actions and currents, thus, Central Visayas may represent a major local phytogeographic boundary in the Philippines. In an unpublished thesis by Calumpong (1981), she recorded 216 taxa of benthic marine algae from Central Visayas. Of these, 50 taxa are newly reported from the Philippines.

The first detailed study on Philippine Caulerpa was by Gilbert (1942), who reported 10 taxa but only Caulerpa serrulata var. typica f. lata was recorded as occurring in Mactan and Cebu Islands, Central Visayas. Later, Taylor (1966) listed two varieties of Caulerpa racemosa from Cebu Island. However, the bulk of the work on algae from Central Visayas was done by Reyes (1972, 1976), who collected from the islands of Negros and Siquijor and described 11 taxa of Caulerpa from this region.

The purpose of this study is to identify the Caulerpa species occurring in Central Visayas, Philippines, and to provide a sound reference and a simple, workable key for the identification of these species. Twenty taxa are described, including Caulerpa reyesii, new species. Of these, three are newly reported from the Philippines and seven taxa are new records from Central Visayas. The taxa new to the Philippines are designated with one asterisk (*) and those from Central Visayas are marked with two asterisks (**) in the systematic section. Basionyms and synonyms are included in the list. References pertinent to the study of Caulerpa, general collection data, and distribution records in the Philippines are included. Collector is designated as EM for E.G. Meñez. The numbers preceding the collector correspond to the year of collection. The collecting site number appears after the collector and this is followed by specimen numbers in parentheses. Specimens are being deposited in the U.S. National Herbarium, Smithsonian Institution, the Silliman University Herbarium in Dumaguete City, Philippines, and the Philippine National Herbarium in Manila, Philippines.

FIGURE 1.—Map of Central Visayas, Philippines; inset is general map of Philippines showing location of Central Visayas (black squares = collecting sites).

ACKNOWLEDGMENTS.—Ernani G. Meñez acknowledges the financial support of the Smithsonian Institution Fluid Research Fund in the Philippines for travel and research. Hilconida P. Calumpong is grateful for the one-year study leave and travel assistance provided by the Silliman University Faculty and Staff Development Program through Dr. Angel C. Alcala. Ms. Calumpong’s research in the USA was made possible through a fellowship award from the Smithsonian Institution Educational Outreach Program.

We are indebted to Dr. William J. Gilbert, Department of Biology, Albion College, to Dr. Robert T. Wilce, Department of Botany, University of Massachusetts, and to Dr. Roy T. Tsuda, The Marine Laboratory, University of Guam, for their critical review of this paper.

We appreciate the help of Dr. Harold E. Robinson, Department of Botany, Smithsonian Institution, in the Latin description of Caulerpa reyesii, new species. We also wish to thank Dr. Paul C. Silva, Curator, University of California (Berkeley) Herbarium for the use of the Index Nominum Algarum card file and his assistance with problems of literature citations.

The loan of herbarium material from the U.S. National Herbarium, Smithsonian Institution and Silliman University Herbarium, Philippines, through the kindness of Dr. James Norris and Prof. Alfredo Reyes, respectively, is gratefully acknowledged.

The bulk of the material used for this study was provided by the Smithsonian Oceanographic Sorting Center. The material was collected during the 1978 and 1979 Smithsonian Institution marine biological expeditions to Central Visayas, Philippines. The expeditions were organized by the Smithsonian Oceanographic Sorting Center and supported by the Smithsonian Scholarly Studies Program and Dreyfuss Foundation.

Description of Collecting Sites

78    EM-1:Liloan Point, southern Cebu (9°25´15˝N, 123°18´10˝E). Algae collected from seagrass bed and coral reef, 0.5–3.0 m, 8 May 1978.

78    EM-3: Paliton, San Juan, Siquijor Island (9°10´25˝N, 123°28´15˝E). Algae collected from an extensive seagrass bed and from a back reef with isolated clumps of corals, 0.2–6.5 m, 9 May 1978.

78    EM-5: Solong-on, Siquijor town, Siquijor Island (9°13´00˝N, 123°27´30˝E). Algae collected from seagrass bed, 0.5–3.0 m, 10 May 1978.

78    EM-7: Tambobo, Siaton, Negros Oriental (9°03´45˝N, 123°07´33˝E). Algae collected from isolated limestone rocks inside the bay, 1.5–3.0 m, 12 May 1978.

78    EM-9: Bantayan Beach, Dumaguete City, Negros Oriental (9°22´30˝N, 123°15´45˝E). Algae collected from isolated rocks and seagrasses, 1.0–2.0 m, 15 May 1978.

78    EM-10: Lag-it, Okiot, Bais, Negros Oriental (9°35´46˝N, 123°09´24˝E). Algae collected from seagrass beds and corals, 1.0–2.0 m, 17 May 1978.

78    EM-22: Linaon, about 8 km south of Sojoton Point, Negros Occidental (9°57´06˝N, 122°26´24˝E). Algae collected from seagrass bed, dead staghorn corals and isolated limestone rocks, 0.5–5.0 m, 28 May 1978.

78    EM-23: Actin, Basay, Negros Oriental (9°25´00˝N, 122°36´12˝E). Algae collected from Sargassum bed, rocks and stony corals, 0.5–5.0 m, 29 May 1978.

78    EM-24: Cabcab, Zamboanguita, Negros Oriental (9°05´30˝N, 123°11´15˝E). Algae collected from seagrass bed and back reef, 0.2–5.0 m, 2 June 1978.

78    EM-25: Banilad, Dumaguete City, Negros Oriental (9°16´45˝N, 123°18´15˝E). Algae collected from isolated rocks, seagrass bed and back reef, 0.5–5.0 m, 3 June 1978.

78    EM-26: Tapon-Norte, San Jose, Negros Oriental (9°25´00˝N, 123°14´15˝E). Algae collected from rocky outcrops, seagrass bed and corals, 0.5–4.0 m, 5 June 1978.

78    EM-27: Suyac Island, off old town of Sagay, Negros Occidental (10°57´15˝N, 123°27´00˝E). Algae collected from coral rubbles, 0.3–6.0 m, 8 June 1978.

78    EM-28: Rocky area between Ilacaon Island and Cadiz, Negros Occidental (11°00´45˝N, 123°15´52˝E). Algae collected from rocks, 2.0 m, 9 June 1978.

78    EM-29: Ilacaon Island, Negros Occidental (11°02´22˝N, 123°12´00˝E). The substrate consists of rocks, corals and seagrass, 0.3–6.0 m, 9 June 1978.

78    EM-30: Sumilon Island (9°27´00˝N, 123°23´24˝E). Algae collected from limestone rocks and corals, 0.3–6.0 m, 15 June 1978.

78    EM-31: Ayungon, Negros Oriental (9°50´30˝N, 123°08´30˝E). Rocky platform, seagrass and coral rubbles, 0.3–3.0 m, 17 June 1978.

78    EM-32: Apo Island (9°04´55˝N, 123°16´30˝E). Algae on limestone rocks and dead staghorn corals, 0.5–3.0 m, 22 June 1978.

78    EM-33: Apo Island, northern end (9°05´09˝N, 123°16´00˝E). Rocks and stony corals provide excellent substrate for algae, 0.4–3.0 m, 23 June 1978.

78    EM-34: Apo Island, western end (9°04´40˝N, 123°16´00˝E). Algae collected from limestone rocks and corals, 0.5–3.0 m, 23 June 1978.

78    EM-37: Bantayan Beach, Dumaguete City, Negros Oriental (9°22´30˝N, 123°15´45˝E). Seagrasses and corals provide good substrate for a rich algal flora, 1.0–2.0 m, 23 June 1978.

78    EM-39: Calindagan Reef, off Dumaguete City, Negros Oriental (9°21´00˝N, 123°16´00˝E). Algae collected by local diver using SCUBA, 26 m, 6 June 1978.

78    EM-40: Bacolod City, Negros Occidental (10°41´15˝N, 122°56´45˝E). Algae collected nearshore from sand substrate, 8 June 1978.

79    EM-8: Pescador Island, Moalboal, Cebu (9°55´30˝N, 123°20´36˝E). Few algae collected from crevices on rocks and corals, 0.3–4.5 m, 7 May 1979.

79    EM-9: Pinagsamahan Beach, Moalboal, Cebu (9°56´12˝N, 123°23´25˝E). Algae abundant in seagrass bed and corals, 1.0–4.5 m, 8 May 1979.

79    EM-11: Northern end of Apo Island, Dauin, Negros Oriental (9°05´20˝N, 123°16´10˝E). Fore-reef slope with few algae, 12.0–36.5 m, 10 May 1979.

79    EM-12: Northwestern portion of Apo Island, Dauin, Negros Oriental (9°04´45˝N, 123°16´00˝E). Algae sparse on rocks and rubbles, 0.5–1.5 m, 10 May 1979.

79    EM-13: Southern end of Apo Island, Dauin, Negros Oriental (9°04´10˝N, 123°16´00˝E). Algae collected from rocks exposed to heavy surf, 0.3–3.5 m, 11 May 1979.

79    EM-14: Southeastern portion of Apo Island, Dauin, Negros Oriental (9°04´30˝N, 123°16´15˝E). Algae collected from staghorn corals, 0.6–4.5 m, 11 May 1979.

79    EM-15: Malo, Siaton, Negros Oriental (9°03´00˝N, 122°59´00˝E). Algae attached to rocks at 3.0 m, 14 May 1979.

79    EM-16: Polopantao, Jimalalud, Negros Oriental (9°59´00˝N, 123°12´06˝E). Algae attached to rocks and corals near river mouth, 1.0–3.0 m, 15 May 1979.

79    EM-18: Barrio Dumanhog, northwestern side of Siquijor Island (9°12´57˝N, 123°30´14˝E). Algae depauperate in seagrass bed and rubbles, 4.5 m, 14 May 1979.

79    EM-21: Barrio Dapdap, northwestern side of Siquijor Island (9°13´21˝N, 123°29´05˝E). Few algae collected from rocks, 4.5 m, 15 May 1979.

79    EM-22: Barrio Cang-aluang, northwestern side of Siquijor Island (9°13´24˝N, 123°28´48˝E). Algae collected from coral reef, 4.5 m, 15 May 1979.

79    EM-23: Barrio Sabang, Larena, northwestern side of Siquijor Island (9°14´27˝N, 123°34´00˝E). Algae collected from coral reef, 4.5 m, 16 May 1979.

Class CHLOROPHYCEAE

Family CAULERPACEAE

Genus Caulerpa

Caulerpa Lamouroux, 1809a:136

Thallus a branched coenocyte, differentiated into a cylindrical, prostrate, creeping stolon and erect assimilative branches; attached by branched rhizoids; erect branches simple or branched, usually bearing terete, flat or compressed branchlets. Thallus internally traversed by extensions (trabeculae) of the cell wall. Reproduction sexual or asexual (anisogamous), when sexual, biflagellated gametes are liberated through branch papillae. Plants are widely distributed in tropical and subtropical seas.

Key to Species of Caulerpa from Central Visayas

*Caulerpa arenicola Taylor

PLATE 1H

Caulerpa arenicola Taylor, 1950:55, 204, pl. 28: fig. 2.

DESCRIPTION.—Plants small, forming loose mats on sand, to 15 mm tall; the creeping stolon branched, terete, to 0.5 mm in diameter, bearing very short, slender, few descending branches with filiform rhizoids at the ends; the substipitate, erect foliar branches simple or forked once or twice, with terete rachis, to 300 µm in diameter, bearing oppositely-pinnate, cylindrical, filiform pinnules, 250–300 µm in diameter, to 1.5 mm long, slightly upcurved to straight, distal end conical, apiculate with basal end somewhat narrower; the foliar branches of even margin, or interrupted occasionally by shorter pinnules.

SPECIMENS STUDIED.—78 EM-9(126–130, 178); 78 EM-25(7); 79 EM-16(50).

PHILIPPINE DISTRIBUTION.—Visayas (Siquijor, Negros Oriental).

Caulerpa brachypus Harvey

PLATE 1I

Caulerpa brachypus Harvey, 1860:333.—Gilbert, 1961:435.—Domantay, 1962:279.—Taylor, 1966:350; 1977:4.—Trono, 1972:95; 1973:217.—Cordero, 1974:6; 1977:25, figs. 2, 3.—Reyes, 1976:153, pl. 7: fig. 4.

DESCRIPTION.—Plants of moderate size, forming dense, long and spreading mats on sand-mud substrate among seagrasses, to 50 mm tall; the prostrate, sparingly branched, terete stolon to 1 mm in diameter, with few descending branches bearing filiform rhizoids at the ends; the substipitate, erect foliar branches proliferous occasionally, ligulate, 5–40 mm long, 2–4 mm wide, with serrate margins or almost entire in a few specimens, irregularly disposed, 2–15 mm distance apart on upper side of stolon.

SPECIMENS STUDIED.—78 EM-5(29–33, 298–316); 79 EM-22(1, 2).

PHILIPPINE DISTRIBUTION.—Luzon (Pangasinan); Visayas (Siquijor, Negros Oriental, Negros Occidental); Mindanao (Zamboanga, Basilan, Sulu).

**Caulerpa crassifolia (C. Agardh) J. Agardh

Plate 1D,E

Caulerpa taxifolia var. crassifolia C. Agardh, 1823:436.

Caulerpa crassifolia (C. Agardh) J . Agardh, 1873:13.—Gilbert, 1942:12; 1946:78; 1961:436.—Meñez, 1961:53.

DESCRIPTION.—Plants forming loose clusters on sand, at 10 m, to 43 mm tall; the creeping stolon terete, slender, no more than 200 µm in diameter, bearing descending branches with filiform, laxly branched rhizoids at the ends; the stipitate, erect foliar branches simple, with compressed rachis, 1–1.5 mm wide, bearing oppositely-pinnate, flat to compressed ramelli, 0.5–1 mm wide, 1.5–3.5 mm long, almost with an even width, except for slight attenuation near the rounded tip, longer ramelli slightly curved; ramelli at mid-portion of foliar branch longer than basal and distal portions; foliar branches very few, distantly spaced on upper side of prostrate stolon, with stipes rarely exceeding 5 mm in length.

SPECIMENS STUDIED.—HPC-6 (material collected by H.P. Calumpong).

PHILIPPINE DISTRIBUTION.—Luzon (Pangasinan); Visayas (Negros Oriental).

REMARKS.—Specimens were collected from sandy bottom, 10 m deep, from Ajong, Sibulan, Negros Oriental on 12 November 1979. Plants examined are smaller compared to those described by Gilbert (1942:12) from West Java.

Caulerpa cupressoides (West in Vahl) C. Agardh

Plate 1B,C

Fucus cupressoides West in Vahl, 1802:38.

Caulerpa cupressoides (West in Vahl) C. Agardh, 1823:441.—Reyes, 1972:142; 1976:155, pl. 8: fig. 7.

DESCRIPTION.—Plants of moderate size, forming loose mats on rocks and seagrasses, to 60 mm tall; the creeping, terete, naked, sparingly branched stolon to 2 mm in diameter, bearing descending branches with thick, branched, filiform rhizoids at the ends; the stipitate, erect, foliar branches simple or branched, with terete or slightly compressed rachis, to 1.5 mm in diameter in larger plants, bearing multiseriate or occasionally pinnate, upcurved, compressed, seminaviculate, ovoid, conical, sometines linear branchlets, to 2 mm long, not exceeding 0.5 mm at the widest portion, with cuspidate or mucronate tips.

SPECIMENS STUDIED.—78 EM-3(62–65); 78 EM-5(243–245); 78 EM-9(131, 179, 243); 78 EM-18(75–79);78 EM-25(8–11).

PHILIPPINE DISTRIBUTION.—Visayas (Negros Oriental, Siquijor).

Caulerpa fastigiata Montagne

PLATE 1A

Caulerpa fastigiata Montagne, 1837:353.—Gilbert, 1942:9; 1961:436.—Reyes, 1976:154, pl. 8: figs. 1, 2.

DESCRIPTION.—Plants filiform, forming thick mats on rocks and sand, to 30 mm tall; the terete, creeping stolon to 300 µm in diameter, with laxly branched, filiform rhizoids borne at the ends of descending filaments; the ascending filaments morphologically similar to stolons, with subopposite, alternate, dichotomous or occasionally sub verticillate branches, to 1 mm long, 150–225 µm in diameter, with obtuse or rounded apices.

SPECIMENS STUDIED.—78 EM-5(215–225); 78 EM-27(29A); 78 EM-31(25); 78 EM-34(643); 78 EM-37(16).

PHILIPPINE DISTRIBUTION.—Luzon (Oriental Mindoro); Visayas (Negros Oriental, Siquijor, Suyac, Apo); Mindanao (Zamboanga, Basilan).

Caulerpa lentillifera J. Agardh

PLATE 1F,G

Caulerpa lentillifera J. Agardh, 1837:173.—Gilbert, 1942:23, 1961:436.—Domantay, 1962:277.—Trono, 1972:96; 1973: 217; 1975:33; 1978:4.—Reyes, 1976:155, 162, pl. 8: fig. 6.—Trono and Tuazon, 1978:3.

DESCRIPTION.—Plants small, in loose mats on coral rubbles and rocks, to 30 mm tall; the terete stolon widespread, to 2 mm in diameter, bearing long, descending branches, to 20 mm long, with laxly branched, filiform rhizoids at the ends; the few, shortly stipitate, erect branches simple or subsimple, with terete rachis, 1–1.5 mm in diameter, bearing spherical translucent ramelli, to 1.5 mm in diameter, supported by stalks, 0.5–1.0 mm long; ramelli conspicuously constricted at the base, imbricately arranged or in rows of four.

SPECIMENS STUDIED.—78 EM-5(34, 227, 228); 79 EM-18(62, 63); 79 EM-23(12).

PHILIPPINE DISTRIBUTION.—Luzon (Oriental Mindoro, Pangasinan, Sorsogon, Catanduanes, Marinduque); Visayas (Negros Oriental, Siquijor); Mindanao (Zamboanga, Basilan, Sulu).

**Caulerpa microphysa (Weber-van Bosse) Feldmann

PLATE 2L

Caulerpa racemosa var. clavifera f. microphysa Weber-van Bosse, 1898:361, pl. 33: fig. 5.

Caulerpa microphysa (Weber-van Bosse) Feldmann, 1955: 430.—Taylor, 1966: 350.

DESCRIPTION.—Plants small, forming dense clusters on corals and rocks, to 15 mm tall; the prostrate, terete, naked stolon less than 1 mm in diameter, bearing descending branches with branched, filiform rhizoids at the ends; the ascending branches on the upper side of the stolon shortly stipitate, simple or occasionally branched, with terete rachis bearing crowded, spherical ramelli, 1–1.5 mm in diameter, supported by stalks, 0.5 mm long, constricted at the apical end.

SPECIMENS STUDIED.—78 EM-5(34A); 78 EM-33(545–547); 78 EM-30(10–12); 78 EM-34(137–139); 79 EM-8(10–15); 79 EM-11(11); 79 EM-12(7–9, 105–107); 79 EM-18(63–67); 79 EM-21(16, 17); 79 EM-22(15–17).

PHILIPPINE DISTRIBUTION.—Visayas (Negros Oriental, Sumilon, Apo, Pescador, Siquijor); Mindanao (Zamboanga).

Caulerpa racemosa (Forsskål) J. Agardh

Fucus racemosus Forsskål, 1775:191.

Caulerpa racemosa (Forsskål) J. Agardh, 1873:35.—Gilbert, 1946:78.—Velasquez, 1953:100.—Meñez, 1961:51.—Taylor, 1966:350.—Reyes, 1972:142.—Trono, 1972:96; 1973:218.—Velasquez, Cornejo, Santiago, and Baens-Arcega, 1973:14, pl. 4: fig. 20.—Trono, 1975:33; 1978:5.—Velasquez, Trono, and Doty, 1975:131.—Trono and Tuazon, 1978:3.—Cordero, 1978:280.

DESCRIPTION.—Plants highly variable, forming loose to dense, sometimes entangled colonies, consisting of few to many erect branches, simple or branched, bearing clavate, turbinate, spherical, hemispherical or discoid ramelli, which are stipitate or substipitate, having obtuse or flattened tips; ramelli distichously, multiseriately or imbricately arranged on terete rachis; creeping stolon terete, with short or long descending branches, giving off branched rhizoids at the ends.

**Caulerpa racemosa var. clavifera (Turner) Weber-van Bosse

PLATE 2A

Fucus clavifer Turner, 1808:126.

Caulerpa clavifera (Turner) C. Agardh, 1817:xxiii.—Dickie, 1877:244.—Howe, 1932:169.

Caulerpa racemosa var. clavifera (Turner) Weber-van Bosse, 1898:361.—Gilbert, 1946:78; 1961:437.—Galutira and Velasquez, 1964:496.

DESCRIPTION.—Plants up to 50 mm tall, with irregularly spaced erect branches, simple or occasionally branched, bearing ramelli that are gradually expanded into a round end, to 2 mm in diameter; ramelli distichously or irregularly arranged on terete rachis.

SPECIMENS STUDIED.—78 EM-32(53); 78 EM-33(548–552); 79 EM-12(13–18); 79 EM-23(11, 12).

PHILIPPINE DISTRIBUTION.—Luzon (Batanes, Babuyan Island, La Union, Ilocos Norte, Pangasinan, Bataan, Batangas, Quezon, Oriental Mindoro); Visayas (Panay, Negros Oriental, Apo); Mindanao (Zamboanga, Misamis Oriental, Basilan, Sulu).

Caulerpa racemosa var. macrophysa (Kützing) Taylor

PLATE 2C

Chauvinia macrophysa Kützing, 1857:6.

Caulerpa racemosa var. macrophysa (Kützing) Taylor, 1928:101.—Gilbert, 1942:19; 1961:439.—Domantay, 1962:277.—Taylor, 1966:351.

DESCRIPTION.—Plants large, to 100 mm tall; erect branches simple or occasionally branched, few and distantly spaced on stout, creeping stolon; branches bearing ramelli expanded into a hemispherical end and arranged irregularly on terete rachis.

SPECIMENS STUDIED.—78 EM-5(1–23, 260–263, 270–297); 79 EM-18(68–71, 80, 81); 79 EM-21(6–11); 79 EM-22(64).

PHILIPPINE DISTRIBUTION.—Luzon (Pangasinan, Oriental Mindoro); Visayas (Cebu, Negros Oriental, Siquijor); Mindanao (Basilan, Sulu).

**Caulerpa racemosa var. occidentalis (J. Agardh) Børgesen

PLATE 2B

Caulerpa chemnitzia var. occidentalis J. Agardh, 1873:37.

Caulerpa racemosa var. occidentalis (J. Agardh) Børgesen, 1907:379.—Domantay, 1962:278.—Taylor, 1966:351.

DESCRIPTION.—Plants up to 50 mm tall; creeping, terete stolon giving off erect branches, simple or rarely branched, bearing ramelli abruptly expanded into subspherical ends, reaching 4 mm in diameter, distichously or radially arranged on terete rachis.

SPECIMENS STUDIED.—78 EM-1(13, 14, 50, 51, 77, 78); 78 EM-9(132); 78 EM-10(80, 81); 78 EM-23(183); 78 EM-24(2–5); 78 EM-26(1, 2); 78 EM-28(53); 78 EM-29(24–29); 78 EM-30(1–5); 78 EM-31(1, 2); 78 EM-34(131); 78 EM-37 (1–5);78 EM-38(383, 390, 392, 394); 79 EM-8(17); 79 EM-12(13–18, 108–117); 79 EM-14(6–12); 79 EM-21(5).

PHILIPPINE DISTRIBUTION.—Luzon (Oriental Mindoro, Pangasinan, Sorsogon, Catanduanes, Marinduque); Visayas (Negros Oriental, Siquijor); Mindanao (Zamboanga, Basilan, Sulu).

**Caulerpa racemosa var. peltata (Lamouroux) Eubank

PLATE 2K

Caulerpa peltata Lamouroux, 1809b:332.—Gilbert, 1942:22.

Caulerpa racemosa var. peltata (Lamouroux) Eubank, 1946:421.—Gilbert, 1961:439.

DESCRIPTION.—Specimens belonging to this species exhibit several different forms, but consistently show the characteristic of having peltate disks at the distal ends of short stalks. The same plant may bear peltate and spherical to turbinate ramelli. The discoid ramelli may be borne on ascending branches, arranged alternately in several ranks or produced directly from the dorsal side of the creeping stolon and supported by terete stipes. Plants found in sheltered areas seem to be the largest, having terete stolons, to 1 mm in diameter, bearing single, stipitate, peltate disks, 1–5 mm in diameter. The depauperate forms were mostly found in rocky, exposed areas and these showed a high degree of polymorphism. They were smaller; slender stolon to 300 µm in diameter, bearing single, stipitate, peltate disks, which are less than 2 mm in diameter; disks sometimes several on erect branches; from margins of disks, occasionally other branches are produced. The stolon bears short, descending branches, with branched filiform rhizoids at the ends.

SPECIMENS STUDIED.—78 EM-25(1–6); 78 EM-33(384, 386–389, 397–399, 401, 402, 542–544); 78 EM-34(124–130, 132, 140–145); 78 EM-30(6); 78 EM-32(24); 79 EM-8(16); 79 EM-11(16, 17); 79 EM-12(10–12, 118–120); 79 EM-15(311); 79 EM-18(59–61); 79 EM-21(9–12); 79 EM-22(11–14).

PHILIPPINE DISTRIBUTION.—Luzon (Oriental Mindoro); Visayas (Negros Oriental, Siquijor, Apo, Sumilon, Pescador); Mindanao (Zamboanga).

**Caulerpa racemosa var. uvifera (Turner) J. Agardh

PLATE 2D

Fucus uvifer Turner, 1819:81, pl. 230.

Caulerpa racemosa var. uvifera (Turner) J. Agardh, 1873:35.—Gilbert, 1961:440.

DESCRIPTION.—Plants simple, erect branches to 30 mm tall, bearing stalked ramelli, imbricately disposed, closely crowded, gradually expanded to swollen tips; length of stalk supporting ramelli about the same as diameter of swollen ends.

SPECIMENS STUDIED.—79 EM-22(60–63).

PHILIPPINE DISTRIBUTION.—Visayas (Siquijor); Mindanao (Zamboanga).

Caulerpa serrulata (Forsskål) J. Agardh

PLATE 2E

Fucus serrulatus Forsskål, 1775:189.

Caulerpa serrulata (Forsskål) J. Agardh, 1837:174.—Gilbert, 1942:14; 1946:78; 1961:440.—Domantay, 1962:278.—Meñez, 1961:53.—Taylor, 1966:351.—Reyes, 1972:142.—Trono, 1972:96; 1973:218; 1975:33.—Cordero, 1973:19.—Velasquez, Cornejo, Santiago, and Baens-Arcega, 1973:15, pl. 5: fig. 21.—Trono and Tuazon, 1978:3.

DESCRIPTION.—Plants moderately large, widespread, in dense colonies on rocks and coral rubbles, to 50 mm tall; the creeping, terete stolon with few, short, descending branches having branched rhizoids at the ends; ascending branches many, flat or compressed, twisted, once to several times dichotomously or irregularly branched; branches regularly with serrate margins, to 30 mm long, 3 mm broad, supported by terete stalks, 3–7 mm long.

SPECIMENS STUDIED.—78 EM-7(1); 78 EM-9(114–120); 78 EM-25(21); 78 EM-29(6–10); 78 EM-30(11); 78 EM-31(4–9); 79 EM-8(6–9); 79 EM-9(18); 79 EM-15(197–200); 79 EM-16(48, 49).

PHILIPPINE DISTRIBUTION.—Luzon (Ilocos Sur, Pangasinan, Bataan, Batangas, Catanduanes, Sorsogon, Marinduque, Oriental Mindoro); Visayas (Leyte, Siquijor, Sumilon, Apo, Negros Oriental); Mindanao (Zamboanga, Basilan).

**Caulerpa serrulata var. boryana f. occidentalis (Weber-van Bosse) Yamada and Tanaka

PLATE 2F

Caulerpa freycinetii var. boryana f. occidentalis Weber-van Bosse, 1898:315, pl. 25: fig. 11.

Caulerpa serrulata var. boryana f. occidentalis (Weber-van Bosse) Yamada and Tanaka, 1938:62.—Gilbert, 1942:15; 1961:441.

DESCRIPTION.—Erect branches to 30 mm tall, 1–3 times dichotomously branched, flat or compressed, supported by a long, terete stalk, to 7 mm long; branches not twisted.

SPECIMEN STUDIED.—78 EM-24(1).

PHILIPPINE DISTRIBUTION.—Palawan (Culion); Visayas (Negros Oriental); Mindanao (Sulu).

Caulerpa sertularioides (Gmelin) Howe

PLATE 3F

Fucus sertularioides Gmelin, 1768:151.

Caulerpa sertularioides (Gmelin) Howe, 1905:576.—Blanco, 1938:512.—Gilbert, 1946:78; 1961:441.—Domantay, 1962:278.—Meñez, 1961:52.—Taylor, 1966:351.—Reyes, 1972:143.—Velasquez, Cornejo, Santiago, and Baens-Arcega, 1973:14, pl. 4: fig. 19.—Trono, 1972:96.—Cordero, 1978:280.

DESCRIPTION.—Plants moderately large, widespread, forming loose colonies on rocks and corals, to 60 mm (rarely 100 mm) tall; stolon terete, to 1 mm in diameter, bearing few, short, descending branches with branched rhizoids at the ends; the erect, simple or occasionally branched, featherlike branches with long stalks extending into a terete rachis, to 1 mm in diameter, bearing cylindrical, pinnately arranged pinnules, commonly 8 mm long, 200 µm in diameter, upcurved with mucronate tips.

SPECIMENS STUDIED.—78 EM-5(35–38, 317–323); 78 EM-22(2, 3, 14); 78 EM-40(1); 79 EM-18(72–74); 79 EM-23(9, 10).

PHILIPPINE DISTRIBUTION.—Luzon (Babuyan Island, Pangasinan, Fafaan, Batangas, Manila Bay, Oriental Mindoro, Sorsogon); Visayas (Siquijor, Aklan, Negros Oriental, Negros Occidental); Mindanao (Sulu).

Caulerpa urvilliana Montagne

PLATE 3D,E

Caulerpa urvilliana Montagne, 1845:21.—Taylor, 1966:352.—Trono, 1972:97; 1973:219.—Reyes, 1976:154, pl. 7: fig. 5.

DESCRIPTION.—Plants large, commonly forming loose colonies on sand, to 150 mm tall; the terete stolon to 3.5 mm in diameter, bearing few to many cylindrical, descending branches with long, branched rhizoids; the terete, erect branches several times dichotomously or alternately branched, with mamillate, multiseriately disposed protuberances and mucronate tips; these erect branches, 20–140 mm tall, supported by terete stalk, 10–60 mm long.

SPECIMENS STUDIED.—78 EM-5(39–46, 246–259); 79 EM-21(13–15); 79 EM-22(56–59, 95).

PHILIPPINE DISTRIBUTION.—Luzon (Sorsogon); Palawan; Visayas (Siquijor, Negros Oriental, Negros Occidental); Mindanao.

*Caulerpa webbiana Montagne

PLATE 2G–J

Caulerpa webbiana Montagne, 1838:129.

DESCRIPTION.—Plants small, in thick cushions on exposed rocks, to 15 mm tall; the terete, creeping stolon beset with few, short, determinate branchlets, divided once to few times; the terete, descending branches on the ventral side of the stolon, naked or with few determinate branchlets, bearing laxly branched rhizoids at the ends; ascending branches from stolon many, simple or branched, 10–15 mm tall, having many whorls, sometimes distichous determinate branchlets 1–5 times divided, with mucronate tips. Plants in sheltered habitat have fewer, but longer branches, occasionally with distichous determinate branchlets.

SPECIMENS STUDIED.—78 EM-32(54); 78 EM-33(132–144, 541).

PHILIPPINE DISTRIBUTION.—Visayas (Apo).

Caulerpa verticillata J. Agardh

PLATE 3A–C

Caulerpa verticillata J. Agardh, 1847:6.—Gilbert, 1961:442.—Trono, 1973:219.—Reyes, 1976:154, pl. 8: fig. 3.

DESCRIPTION.—Plants filiform, attached to rocks and corals, to 50 mm tall; the creeping, naked terete stolon with numerous short, descending branches, terminated by laxly branched, filiform rhizoids; the terete, erect branches, simple or irregularly branched, bearing abundant tufts of whorled, determinate branchlets, to 180 µm in diameter, to 8 times dichotomously branched; the filiform branchlets constricted at the base, with 2–3 teeth at the apices.

SPECIMENS STUDIED.—79 EM-12(3–6).

PHILIPPINE DISTRIBUTION.—Palawan; Visayas (Siquijor, Apo); Mindanao (Sulu, Basilan).

*Caulerpa reyesii, new species

FIGURE 2; PLATE 3G–I

DESCRIPTION.—Plantae mediocriter robustae in coloniis laxis late effusae. Stolones repentes teretes ramosi ad 2.5 mm in diametro; rami descendentes ad 60 mm longi in apicem rhizoideis filiformis ferentes; rami ascendentes 10–70 mm alti simplices vel interdum frondiformes; frondes basi vel ad medio ramosae basi breviter terete stipitatae, stipitis in rachidis compressis 1 mm latis continuis, ramulis subsessilibus complanatis obovatis clavatis adaxialiter incurvalis ad 7 mm longis et in partibus latissimus 2 mm latis plerumque as apicem mucronatis vel cuspidatis distichis vel variabiliter tetrastichis et subimbricatis.

FIGURE 2.—Habit of Caulerpa reyesii, new species. Holotype: 78 EM-5(330), Solong-on, Siquijor Island, 10 May 1978.

Plants moderately robust, widespread, forming loose colonies. The creeping stolon terete, branched, to 2.5 mm in diameter, bearing long, terete, descending branches, to 60 mm long, with filiform rhizoids at the ends. The ascending branches from stolon, 10–70 mm tall, simple or if branched, secondary branching produced from the mid-portion or the lower margin of the compressed rachis; erect branches supported by short, cylindrical stalks that extend into a compressed rachis, 1 mm broad, bearing subsessile, flat, overlapping, obovate, clavate ramelli, which are adaxially incurved, tetrastichous, distichous, to 7 mm long and 2 mm at the widest portion, mostly mucronate or cuspidate at the tips.

HOLOTYPE.—78 EM-5(330), on coral rubbles and sand in seagrass bed, at Solong-on, Siquijor Island, Philippines, 10 May 1978; deposited in the U.S. National Herbarium, Smithsonian Institution, Washington, D.C, U.S.A. Isotypes deposited in the Philippine National Herbarium, Manila, Philippines and Silliman University Herbarium, Dumaguete City, Philippines.

PARATYPES STUDIED.—78 EM-5(24–28).

REMARKS.—It is with great pleasure to name this species of Caulerpa after our colleague and friend, Professor Alfredo Reyes, Department of Biology, Silliman University, Dumaguete City, Philippines, for his unselfish efforts to advance phycological studies in Central Visayas. Professor Reyes (1976:157, pl. 9: figs. 6, 7) previously recorded and described this species as Caulerpa species. His specimens were collected from “dead coral-sandy bottom at the lower intertidal zone” at Siquijor Island, Philippines.

Literature Cited

Agardh, C.A.

1817. Synopsis algarum Scandinaviae, adjecta dispositione universali algarum. xl + 135 pages. Lund.

1823. Species algarum rite cognitae cum synonymis, differentis specificis et descriptionibus succinctis. Volume 1, part 2, pages 169–521. Lund.

Agardh, J.G.

1837. Novae species algarum, quas in itinere ad oras maris rubri collegit Eduardus Rüppell; cum observationibus. Museum Senckenbergianum, Abhandlungen aus dem Gebiele der Beschreibenden Naturgeschichte, 2:169–174.

1847. Nya alger frän Mexico. Öfversigt af Kongliga Velenskaps Akademiens Förhandlingar, 4:5–17.

1873. Till algernes systematik, I: Caulerpa., II: Zonaria. III: Sargassum. Lunds Universitets Årsskrift, 9(8): 71 pages.

Blanco, G.J.

1938. Fisheries of Northeastern Luzon and the Babuyan and Batanes Islands. The Philippine Journal of Science, 66(4):501–521, 5 plates.

Børgesen, F.

1907. An Ecological and Systematic Account of the Caulerpas of the Danish West Indies. Kongelige Danske Videnskabernes Selskab Skrifter, Naturvidenske og Mathematik Afd, 4(5):337–392, 31 figures.

Calumpong, H.P.

1981. The Systematics and Distribution of the Benthic Marine Algae of Negros and Adjacent Islands, Philippines. Masters thesis, Silliman University, Dumaguete City 6501, Philippines.

Cordero, P.A., Jr.

1973. On the Marine Algae of Biliran (Leyte) and Vicinities, Central Philippines. Leyte-Samar Studies, Divine Word University, 7(1):15–37.

1974. Distribution of Caulerpa brachypus Harvey (Chlorophyta) in Japan and Adjacent Regions. Nankiseibutu: Nanki Biological Society, 16(1):6–9.

1977. Phycological Observations, V: Gross Morphological Polymorphisms in Caulerpa brachypus (Caulerpales, Chlorophyta) from the Philippines, with Notes on Their Taxonomy. Bulletin of the Japanese Society of Phycology, 25:25–30, 3 figures.

1978. Phycological Observations, VI: Mangrove-associated Algae from Aklan, Philippines. Kalikasan, Philippine Journal of Biology, 7(3):275–296, 5 figures.

Dickie, G.

1877. Contributions to the Botany of the Expedition of H.M.S. Challenger. Journal of the Linnean Society [of London] (Botany), 15:235–246.

Domantay, J.S.

1962. An Ecological Survey of the Marine Vegetation of the Hundred Islands and Vicinity. The Philippine Journal of Science, 90(2):271–295.

Eubank, L.L.

1946. Hawaiian Representatives of the Genus Caulerpa. University of California Publications in Botany, 18(18):409–432, figures 1, 2, plate 22.

Feldmann, J.

1955. Les plantes des Caulerpa et leur valeur systematique. Revue General de Botanique, 62:422–431, figures 1, 2.

Forsskål, P.

1775. Flora Aegyptiaco-Arabica. 219 pages. Copenhagen.

Galutira, E.C., and G.T. Velasquez

1964. Taxonomy, Distribution and Seasonal Occurrence of Edible Marine Algae in Ilocos Norte, Philippines. The Philippine Journal of Science, 92(4):483–522, figure 1, plates 1–9.

Gilbert, W.J.

1942. Notes on Caulerpa from Java and the Philippines. Papers of the Michigan Academy of Sciences, Arts and Letters, 27:7–26, 5 figures.

1946. Studies on Philippine Chlorophyceae, II: Survey of Literature and List of Recorded Species Prior to 1940. Bulletin of the Torrey Botanical Club, 73(1):73–79.

1961. An Annotated Checklist of Philippine Marine Chlorophyta. The Philippine Journal of Science, 88(4):413–449, 1 plate.

Gmelin, S.G.

1768. Historia fucorum. 239 pages, 33 plates. St. Petersburg.

Harvey, W.H.

1860. Characters of New Algae, Chiefly from Japan and Adjacent Regions, Collected by Charles Wright in the North Pacific Exploring Expedition under Captain John Rodgers. Proceedings of the American Academy of Arts and Sciences, 4:327–335.

Howe, M.A.

1905. Phycological Studies, II: New Chlorophyceae, New Rhodophyceae, and Miscellaneous Notes. Bulletin of the Torrey Botanical Club, 32:563–586, plates 23–29.

1932. Marine Algae from the Islands of Panay and Negros (Philippines) and Niuafoou (between Samoa and Fiji). Journal of the Washington Academy of Science, 22(7):167–170, figure 1.

Kützing, F.T.

1857. Tabulae phycologicae. Volume 7, 40 pages, 100 plates, Nordhausen.

Lamouroux, J.V.

1809a. Mémoires sur les Caulerpes, nouveau genre de la famille des algues marines. Journal de Botanique, 2:136–146.

1809b. Observation sur la physiologie des algues marines, et description de cincq nouveau genres de cette famille. Nouveau Bulletin des Sciences de la Societe Philomatique de Paris, 1:330–333.

Meñez, E.G.

1961. The Marine Algae of the Hundred Islands, Philippines. The Philippine Journal of Science, 90(1):37–86, 12 plates.

Montagne, J.F.C.

1837. Centurie de plantes cellulaires exotiques nouvelles. Annates des Sciences Naturelles (Botanique), series 2, 8:345–370.

1838. De l’organisation et du mode de reproduction des Caulèrpees, et an particulier du Caulerpa Webbiana espèce nouvelle des lies Canaries. Annales des Sciences Naturelles, Botanique, 9:129–150.

1845. Voyage au Pole Sud et dans l’Oceanie, exécuté par les corvettes l’Astrolabe et la Zélée. 168 pages, 20 plates. Paris.

Reyes, A.Y.

1972. A Survey of the Littoral Benthic Algae of the Coastal Areas of Dumaguete City. The Philippine Journal of Science, 99(3–4): 131–163.

1976. The Littoral Benthic Algae of Siquijor Province, I: Cyanophyta and Chlorophyta. The Philippine Journal of Science, 105(3):133–191, 11 plates.

Taylor, W.R.

1928. The Marine Algae of Florida, with Special Reference to the Dry Tortugas. In Papers from the Tortugas Laboratory, 25. Carnegie Institution of Washington Publication, 379: 219 pages, plates 1–37.

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1977. Notes on Plants of the Genus Caulerpa in the Herbarium of Maxwell S. Doty at the University of Hawaii. Atoll Research Bulletin, 208:1–7.

Trono, G.C., Jr.

1972. Annotated Checklist of Some Marine Benthic Algae from Tawi-Tawi, Sulu Archipelago. University of the Philippines Natural and Applied Science Bulletin, 24(3):85–112.

1973. The Marine Algae of Siasi Island and Vicinity, I: Introduction and Chlorophyta. Kalikasan, Philippine Journal of Biology, 1:207–228.

1975. The Marine Algae of Bulusan and Vicinity, province of Sorsogon, I: Introduction and Chlorophyta. Kalikasan, Philippine Journal of Biology, 4:23–41.

1978. Notes on Some Marine Benthic Algae of Sta. Cruz, Marinduque, Philippines. University of the Philippines Natural Science Research Center Technical Report, 54: 24 pages.

Trono, G.C., Jr., and A. Tuazon

1978. Notes on Some Marine Benthic Algae from Bakawan and Sula Islands, Province of Catanduanes, Philippines. University of the Philippines Natural Science Research Center Technical Report, 53: 19 pages.

Turner, D.

1808. Fuci sive plantarum fucorum genen a botamcis asenptarum icones, descriptiones et historia. Volume 1, 165 pages, 71 plates. London.

1819. Fuci sive plantarum fucorum genen a botamcis ascriptarum icones, descriptiones et historia. Volume 4, 153 pages, plates 197–258, index 1–7. London.

Vahl, M.

1802. Endeel Kryptogamiske planter (fuci) fra St. Croix. Skrifter af Naturhistone Selskabet, 5(3):29–47.

Velasquez, G.T.

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1898. Monographic des Caulerpes. Annales du jardin Botanique de Buitenzorg, 15:243–401, plates 20–34.

Yamada, Y., and T. Tanaka

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PLATES

PLATE 1

(illustrations reduced to 53%)

FIGURE A.—Caulerpa fastigiata, habit, X 2.

FIGURE B.—Caulerpa cupressoides, habit, X 2.

FIGURE C.—Same, pinnules, showing 3-ranked branching, X 4.

FIGURES D., E.—Caulerpa crassifolia, habit, X 1.

FIGURE F.—Caulerpa lentillifera, ramelli, showing imbricate branching and constricted bases, X 13.

FIGURE G.—Same, habit, X 2.6.

FIGURE H.—Caulerpa arenicola, habit, X 3.

FIGURE I.—Caulerpa brachypus, habit X 1.3.

PLATE 2

(illustrations reduced to 53%)

FIGURE A.—Caulerpa racemosa var. clavifera, habit, X 2.5.

FIGURE B.—Caulerpa racemosa var. occidentalis, habit, X 1.

FIGURE C.—Caulerpa racemosa var. macrophysa, habit, X 1.

FIGURE D.—Caulerpa racemosavar. uvifera habit, X 1.

FIGURE E.—Caulerpa serrulata, typical, habit, X 1.

FIGURE F.—Caulerpa serrulata var. boryana f. occidentalis, habit, X 1.

FIGURE G.—Caulerpa webbiana, habit X 2.

FIGURE H.—Same, erect branch with whorled branchlets, X 12.

FIGURE I.—Same, erect branch with distichous branchlets, X 16.

FIGURE J.—Same, stolon beset with determinate branchlets, X 16.

FIGURE K.—Caulerpa racemosa var. peltata, habit, X 2.

FIGURE L.—Caulerpa microphysa, habit, X 2.

PLATE 3

(illustrations reduced to 55%)

FIGURE A.—Caulerpa verticillata, habit, X 3.3.

FIGURE B.—Same branchlets with toothed apices, X 31.

FIGURE C.—Same, portion of erect branching showing whorled branchlets, X 7.

FIGURE D.—Caulerpa urvilliana, habit, X 1.

FIGURE E.—Same, erect branch with mamillate protuberances, X 6.

FIGURE F.—Caulerpa sertularioides, habit, X 2.

FIGURE G.—Caulerpa reyesii, new species, habit, X 1.

FIGURE H.—Same, erect branch with multi-ranked branchlets, X 3.5.

FIGURE I.—Same, erect branch with distichous branchlets, X 3.

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