The Genus Caulerpa

from Central Visayas,
Philippines

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~~ “HILCONIDA P- CALUMPONG sen al sa

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(OTHE MARINE SCIENCES * NUMBER 1

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SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES © NUMBER 17

The Genus Caulerpa
from Central Visayas,

Philippines

Ernani G. Menez
and FHilconida P. Calumpong

ISSUED
OCT 5 1982

SMITHSONIAN PUBLICATIONS

SMITHSONIAN INSTITUTION PRESS
City of Washington
1982

ABSTRACT

Menez, 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 reyesi1, 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 589.477 82-5976 AACR2

Contents

EERO CUCHON EE Aee Tere me re Ms es ake dl ee eens
FNC KMOWLCASINCIATS eee nr ME nA Went te) Maeno nay vlan wes
Wesemptroniols Collecting Sites e 5 a2. no ns ence ee bee ee lee oe eee
lassi C:HUOROPHY CHAP Mere y ieaciys ry cps none a lode encase debe a.
aImMily ga CAUTEERPACHAE pea ean yoi etic csi 5 ss sf alctne tiles epistles ees
Genus CAMGADE cps ding 60.0 F038 6 OO ATO > RO ee an gree
Key to Species of Caulerpa from Central Visayas ...............
CORWGHDE, CHGOGHA, VWASTIOSE, 7 oho ie oa ee ees erie ae
Caulerpakurachy pusmllaive yam yates oe oe esse ec
Caulerparcrassijolia (@ Agardh) J, Agardh ~.........-......--:-:
Caulerpa cupressoides (West in Vahl) C. Agardh .................
CaulerpanasiveratagNontagne © seen aon sd oe
Caulerpanlentilujcram|peNeacd ieee oe es eine eee eta
Caulerpa microphysa (Weber-van Bosse) Feldmann ..............
Caulerpaacemosas(Korsskal) J. Agardh ..................-......
Caulerpa racemosa var. clavifera (Turner) Weber-van Bosse .......
Caulerpa racemosa var. macrophysa (Kutzing) Taylor .............
Caulerpa racemosa var. occidentalis (J. Agardh) Bgrgesen .........
Caulerpa racemosa var. peltata (Lamouroux) Eubank .............
Caulerpa racemosa var. uvifera (Turner) J. Agardh ...............
Caulerparsernulatas(Horsskal) ee Agardh 0 .......2.....-....--2-
Caulerpa serrulata var. boryana f. occidentalis (Weber-van Bosse)
NWamadasancdalianakaurear sien ite vc cine ee ase lee
Caulerparsertulariovdesi(Gmelin)iHowe ..-.......5..........2..-.
CanlerparuruilianasNMiontaeme wns pee 12s oee ene
CaulerparwebvianaiNMontasne sss] -eeee ee eae eee ee
Gamlerpaucnticilatam)| py NG acai re hoy en ee ee be
COUCH PTAC CSUICWESPECLESM i eee ae ook ei
i Lareerraitune (Caeel seh eee oo sla ath ae hee A gece oe Soa
[PILES 5 c's w.dtrip'e p88) Bleiots Wie cLeeS oo 8 ote Se Remmi ae ar ee a

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OMOWmWAWMWOnNI NYITNNAMAAAVVH SL HK W

The Genus Caulerpa
from Central Visayas,

Philippines

Ernani G. Menez
and Flilconida P. Calumpong

Introduction

During the summers of 1978 and 1979, Smith-
sonian Institution conducted biological collecting
expeditions in Central Visayas, Philippines, in
order to collect marine plant and animal orga-
nisms for purposes of systematic, ecological, and
distributional studies. Included in the plant col-
lection 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 previ-
ously 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 Archi-
pelago (Figure 1). The area consists of the islands
of Negros, Cebu, Bohol, Siquijor, and their sat-
ellite islets. Except for the southwestern portion

Emani G. Menez, Smithsonian Oceanographic Sorting Center, Smith-
sonian Institution, Washington, D.C. 20560. Hilconida P. Calum-
pong, Department of Biology, Silliman University, Dumaguete City
6501, Philippines.

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 phy-
togeographic 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 re-
corded 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, includ-

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

VISAYAN
SEA

CAMOTES
SEA

20,

1 SUMILON
@ . 1.

ae

SULU SEA

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

NUMBER 17

ing 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 in-

cluded. Collector is designated as EM for E.G. |

Menez. The numbers preceding the collector cor-
respond 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. Na-
tional Herbarium, Smithsonian Institution, the
Siliman University Herbarium in Dumaguete
City, Philippines, and the Philippine National
Herbarium in Manila, Philippines.

ACKNOWLEDGMENTS.—Ernani G. Menez ac-

knowledges the financial support of the Smith-
sonian 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 Silli-
man University Faculty and Staff Development
Program through Dr. Angel C. Alcala. Ms. Cal-
umpong’s research in the USA was made pos-
sible through a fellowship award from the Smith-
sonian Institution Educational Outreach Pro-
gram.
We are indebted to Dr. William J. Gilbert,
Department of Biology, Albion College, to Dr.
Robert T. Wilce, Department of Botany, Univer-
sity 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
reyestt, 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.

Y

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 dur-
ing 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, Siquyor 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 Ori-
ental (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 col-
lected 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

4 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

(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). Al-
gae 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°-
1600” E). Rocks and stony corals provide excellent sub-
strate 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 Ori-
ental (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 sub-
strate, 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 col-
lected 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 Siqui-
jor 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 col-
lected 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, usu-
ally bearing terete, flat or compressed branchlets.
Thallus internally traversed by extensions (tra-
beculae) 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 sub-
tropical seas.

Key to Species of Caulerpa from Central Visayas

1. Erect branches angular, with thorn-like or dentate projections, stolons

naked

‘Ans dibchie reser ae C. urvilliana

NUMBER 17 5
2 Stolonsrand branches similanim forme). oy) 4 3
Stolons and branches dissimilar in form ...........................-. 4
3. Erect branches bearing verticillate branchlets ........... C. verticillata
Erect branches without verticillate branchlets ............ C. fastigiata
Ane recthoranches tation Compressed.) cst yas ea ne 5
Rrectibranchesmottlat or compressed): 7.0.) 94 ea 10
Di, IRBVATIETUN fOpRES Tea ES sie eee ers a eee Reape ae = Laat As8 ter Rd 6
Reatnielligalleseinitigeen etter er nar, Nm a eat gral nn Atel NE ate ay 9
Ommeanmellineyiadiically icp iqu te ae ee ih eh oe ue UL RE UE 7
ENamelligilatconcompressedi ry... ake e les ei ce ek Se 8
7. Plants small, to 15 mm tall, branches substipitate, bearing pinnules to
Iho) HOTT, IKOVEYeA. Co tale eect at Ce mene Sanne On Pe tT C. arenicola
Plants moderately large, to 60 mm tall, branches long-stalked, bearing
pinmulesicorsimamulong. ee. 6 es We en C. sertularioides
8. Branchlets oppositely-pinnate; stolons 200 wm in diameter ............
ey cir Ames, fu is bh C. crassifolia
Branchlets tetrastichous, distichous; stolons 1-2 mm in diameter ......
5 ode a Bt SS Geh ESSAY aaa ese ALE or UE et C. reyesii, new species
9. Branches dichotomous, margins regularly serrate ......... C. serrulata
Branches proliferous, margins irregularly serrate ......... C. brachypus
NOME Stolommtomientose ee yd chaetags velo Lye C. webbiana
StOlOiMpiMacemene ee a nee ee 5 SONIA Ca nd no 22k Dee en eat 11
11. Ramelli seminaviculate, in several ranks .............. C. cupressoides
Ramelli clavate or abruptly ending in swollen tips ................. 12

12. Ramelli thick-walled, not translucent
Ramelli thin-walled, translucent

3

13. Ramelli many, imbricate or in 4-rows; branches long and near each

other

* 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 pm in diameter,
bearing oppositely-pinnate, cylindrical, filiform
pinnules, 250-300 wm in diameter, to 1.5 mm
long, slightly upcurved to straight, distal end
conical, apiculate with basal end somewhat nar-

ree Deen eenre EN a Ul C. lentillifera

rower; 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).
PHrtippInE DistriBuTION.— Visayas (Siquijor,
Negros Oriental).

Caulerpa brachypus Harvey
Puate 11

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, form-

6 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

ing 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 substip-
itate, erect foliar branches proliferous occasion-
ally, ligulate, 5-40 mm long, 2-4 mm wide, with
serrate margins or almost entire in a few speci-
mens, 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 (Pangasi-
nan); Visayas (Siquiyor, 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.—Menez, 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 um 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 por-
tions; 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 col-
lected by H.P. Calumpong).

PHILipPINE DistripuTion.—Luzon
nan); Visayas (Negros Oriental).

REMARKS.—Specimens were collected from
sandy bottom, 10 m deep, from Ajong, Sibulan,
Negros Oriental on 12 November 1979. Plants

(Pangasi-

examined are smaller compared to those de-
scribed 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, form-
ing 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, fili-
form 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 occasion-
ally pinnate, upcurved, compressed, seminavicu-
late, 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 DiIsTRIBUTION.— Visayas
Oriental, Siquijor).

(Negros

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 um in diameter, with laxly
branched, filiform rhizoids borne at the ends of
descending filaments; the ascending filaments
morphologically similar to stolons, with subop-
posite, alternate, dichotomous or occasionally
subverticillate branches, to 1 mm long, 150-225
jum in diameter, with obtuse or rounded apices.

SPECIMENS STUuDIED.—78 EM-5(215-225); 78

NUMBER 17

EM-27(29A); 78 EM-31(25); 78 EM-34(643); 78
EM-37(16).

PHILIPPINE DistRipsuTiIoN.—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 di-
ameter, 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).

PuiuipPiInE DistripsuTion.—Luzon (Oriental
Mindoro, Pangasinan, Sorsogon, Catanduanes,
Marinduque); Visayas (Negros Oriental, Siqui-
jor); 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 | mm in
diameter, bearing descending branches with
branched, filiform rhizoids at the ends; the as-
cending branches on the upper side of the stolon

shortly stipitate, simple or occasionally branched,
with terete rachis bearing crowded, spherical ra-
melli, 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 (Forsskal) J. Agardh

Fucus racemosus Forsskal, 1775:191.

Caulerpa racemosa (Forsskal) J. Agardh, 1873:35.—Gilbert,
1946:78.— Velasquez, 1953: 100.—Menez, 1961:51.—Tay-
lor, 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.—Velas-
quez, Trono, and Doty, 1975:131.—Trono and Tuazon,
1978:3.—Cordero, 1978:280.

Description.—Plants highly variable, forming
loose to dense, sometimes entangled colonies, con-
sisting of few to many erect branches, simple or
branched, bearing clavate, turbinate, spherical,
hemispherical or discoid ramelli, which are stipi-
tate or substipitate, having obtuse or flattened
tips; ramelli distichously, multiseriately or imbri-
cately 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 oc-

8 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

casionally 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.—/8 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, Panga-
sinan, Bataan, Batangas, Quezon, Oriental Min-
doro); Visayas (Panay, Negros Oriental, Apo);
Mindanao (Zamboanga, Misamis Oriental, Bas-
ilan, Sulu).

Caulerpa racemosa var. macrophysa (Kutzing)
Taylor

PLATE 2c

Chauvinia macrophysa Kutzing, 1857:6.

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

DescriptTion.—Plants large, to 100 mm tall;
erect branches simple or occasionally branched,
few and distantly spaced on stout, creeping sto-
lon; 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); 799 EM-22(64).

PHILipPINE Distripution.—Luzon (Pangasi-
nan, Oriental Mindoro); Visayas (Cebu, Negros
Oriental, Siquijor); Mindanao (Basilan, Sulu).

** Caulerpa racemosa var. occidentalis

(J. Agardh) Borgesen
PLATE 2B

Caulerpa chemnitzia var. occidentalis J. Agardh, 1873:37.
Caulerpa racemosa var. occidentalis (J. Agardh) Bgrgesen,
1907:379.—Domantay, 1962:278.—Taylor, 1966:351.

Description.—Plants up to 50 mm tall; creep-
ing, terete stolon giving off erect branches, simple
or rarely branched, bearing ramelli abruptly ex-

panded 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).

PuivipPINnE Distripution.—Luzon (Oriental
Mindoro, Pangasinan, Sorsogon, Catanduanes,
Marinduque); Visayas (Negros Oriental, Siqui-
jor); 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 con-
sistently 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 sev-
eral 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 wm in
diameter, bearing single, stipitate, peltate disks,
which are less than 2 mm in diameter; disks
sometimes several on erect branches; from mar-
gins of disks, occasionally other branches are pro-
duced. The stolon bears short, descending
branches, with branched filiform rhizoids at the
ends.

NUMBER 17

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 DistRipuTION.—Luzon (Oriental
Mindoro); Visayas (Negros Oriental, Siquijor,
Apo, Sumilon, Pescador); Mindanao (Zam-
boanga).

** 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.

DescriptTion.—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 (Forsskal) J. Agardh
PLATE 2E

Fucus serrulatus Forsskal, 1775:189.

Caulerpa serrulata (Forsskal) J. Agardh, 1837:174.—Gilbert,
1942:14; 1946:78; 1961:440.—Domantay, 1962:278.—
Menez, 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, wide-
spread, in dense colonies on rocks and coral rub-
bles, 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 mar-

gins, 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 DistRipuTION.—Luzon (Ilocos Sur,
Pangasinan, Bataan, Batangas, Catanduanes,
Sorsogon, Marinduque, Oriental Mindoro); Vi-
sayas (Leyte, Siquijor, Sumilon, Apo, Negros Ori-
ental); 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 1938:62.—Gilbert, 1942:15;

1961:441.

and ‘Tanaka,

Description.—Erect branches to 30 mm tall,
1-3 times dichotomously branched, flat or com-
pressed, supported by a long, terete stalk, to 7
mm long; branches not twisted.

SPECIMEN STUDIED.—/8 EM-24(1).

PuivipPinE Distripution.—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.—Menez, 1961:52.—Taylor, 1966:351.—Reyes,
1972:143.—Velasquez, Cornejo, Santiago, and Baens-Ar-
cega, 1973:14, pl. 4: fig. 19.—Trono, 1972:96.—Cordero,
1978:280.

Description.—Plants moderately large, wide-
spread, 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

10 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

erect, simple or occasionally branched, feather-
like branches with long stalks extending into a
terete rachis, to | mm in diameter, bearing cylin-
drical, pinnately arranged pinnules, commonly 8
mm long, 200 um in diameter, upcurved with
mucronate tips.

SPECIMENS StupIED.—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).

PuiuippInE DistripuTion.—Luzon (Babuyan
Island, Pangasinan, Bataan, Batangas, Manila
Bay, Oriental Mindoro, Sorsogon); Visayas (Si-
quior, Aklan, Negros Oriental, Negros Occiden-
tal); 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 form-
ing 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
times dichotomously or alternately
branched, with mamillate, multiseriately dis-
posed protuberances and mucronate tips; these
erect branches, 20-140 mm tall, supported by
terete stalk, 10-60 mm long.

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

PHILIPPINE DistTRIBUTION.—Luzon (Sorsogon);
Palawan; Visayas (Siquijor, Negros Oriental, Ne-
gros Occidental); Mindanao.

several

* Caulerpa webbiana Montagne
PLATE 2c-}
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; as-
cending 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 branch-
lets.

SPECIMENS STUDIED.—/8 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, descend-
ing branches, terminated by laxly branched, fili-
form rhizoids; the terete, erect branches, simple
or irregularly branched, bearing abundant tufts
of whorled, determinate branchlets, to 180 wm 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 DistrrpuTion.—Palawan; Visayas
(Siquijor, Apo); Mindanao (Sulu, Basilan).

* Caulerpa reyesii, new species
b )

Figure 2; PLATE 3G-1

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 sim-
plices vel interdum frondiformes; frondes basi vel
ad medio ramosae basi breviter terete stipitatae,
stipitis in rachidis compressis | mm latis contin-
uis, ramulis subsessilibus complanatis obovatis

NUMBER

17

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

11

IW SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

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.

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 com-
pressed rachis; erect branches supported by short,
cylindrical stalks that extend into a compressed
rachis, | mm broad, bearing subsessile, flat, over-
lapping, obovate, clavate ramelli, which are adax-
ially incurved, tetrastichous, distichous, to 7 mm
long and 2 mm at the widest portion, mostly

mucronate or cuspidate at the tips.

Hototypee.—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 Insti-
tution, Washington, D.C., U.S.A. Isotypes depos-
ited in the Philippine National Herbarium, Ma-
nila, Philippines and Silliman University Her-
barium, Dumaguete City, Philippines.

PARATYPES STUDIED.—78 EM-5(24-28).

RemaARKS.—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 re-
corded and described this species as Caulerpa spe-
cies. His specimens were collected from “dead
coral-sandy bottom at the lower intertidal zone”
at Siquijor Island, Philippines.

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~ PLATES

16

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

PLATE 1

(illustrations reduced to 53%)

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

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

FicurE c.—Same, pinnules, showing 3-ranked branching, X 4.

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

Ficure F.—Caulerpa lentillifera, ramelli, showing imbricate branching and constricted bases,
xX 13.

FicureE G.—Same, habit, X 2.6.

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

Ficure 1.—Caulerpa brachypus, habit X 1.3.

NUMBER 17

18

FIGURE
FIGURE
FIGURE
FIGURE
FIGURE
FIGURE
FIGURE
FIGURE
FIGURE

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

PLATE 2

(illustrations reduced to 53%)

A.—Caulerpa racemosa var. clavifera, habit, X 2.5.
B.—Caulerpa racemosa var. occidentalis, habit, X 1.
c.—Caulerpa racemosa var. macrophysa, habit, X 1.
pD.—Caulerpa racemosa var. uvifera, habit, X 1.

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

F.—Caulerpa serrulata var. boryana f. occidentalis, habit, X 1.
G.—Caulerpa webbiana, habit X 2.

H.—Same, erect branch with whorled branchlets, * 12.
1.—Same, erect branch with distichous branchlets, < 16.

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

FIGURE
FIGURE

K.—Caulerpa racemosa var. peltata, habit, X 2.
L.—Caulerpa microphysa, habit, X 2.

19

NUMBER 17

20

Ficu
Ficu
Ficu
Ficu
Ficu

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

PLATE 3

(illustrations reduced to 55%)

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

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

RE C.—Same, portion of erect branching showing whorled branchlets, X 7.
RE D.—Caulerpa urvilliana, habit, X 1.

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

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

Ficu

Fict

RE G.—Caulerpa reyesit, new species, habit, X 1.
RE H.—Same, erect branch with multi-ranked branchlets, X 3.5.

FiGure 1.—Same, erect branch with distichous branchlets, X 3.

21

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