».._ ERNANI G.-MENEZ, RONALD C. PHILLIPS, |
_. -...,and-HILCONIDA P. CALUMPONG .

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THE MARINE SCIENCES
een

- NUMBER2I
oe enon. ere

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SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES NUMBER 21

Seagrasses from the Philippines

Ernani G. Menez, Ronald C. Phillips,
and Filconda P. Calumpong

ISSUED
DEC 1 1083

SMITHSONIAN PUBLICATICNS

SMITHSONIAN INSTITUTION PRESS
City of Washington
1983

ABSTRACT

Menez, Ernani G., Ronald C. Phillips, and Hilconida P. Calumpong. Sea-
grasses from the Philippines. Smithsonian Contributions to the Marine Sciences,
number 21, 40 pages, 26 figures, 1983.—Seagrasses were collected from various
islands in the Philippines during 1978-1982. A total of 12 species in seven
genera are recorded. Generic and specific keys, based on vegetative characters,
are provided for easier differentiation of the seagrasses. General discussions of
seagrass biology, ecology, collection and preservation are presented. Local and
world distribution of Philippine seagrasses are also included.

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.

Seagrasses from the Philippines.

(Smithsonian contributions to the marine sciences ; no. 21)

Bibliography: p.

Supt. of Docs. no.: SI 1.41:21

1. Seagrasses—Philippines. I. Phillipps, Ronald C. II. Calumpong, Hilconida P. III. Ti-
tle. IV. Series.

QK495.A14M46 1983 584.73 83-600168

Contents

Page

MAGROCUC TOME tree trary U ame ME sud Ea ark ia ele 1
INCKMOWICAM AINE Mts Mae wees CHEF ee eames win tae anne teas 3
IMatenialstandeWicthodsiiar, ire ie Os aici aa. edn noes 3
Collectineyandebresernvinls, SCACrasses 2) ee ah aa 4
General Features of Seagrass Biology and Ecology ....:............... 6
INevatonchey Philippine: SEASTASSES® aysiu ak eee eine sete es Z
DIV AST OMAN DELO PELVA AW Pp eras ee ne ti cs Pine a nn a Mle eae ele Ug ee haces 8
ClassyMONOCOMUEDONEAB in: 2) 045. Glows Vos she. hace eee nee ees 8
@denEVORIAR meri at cyclen nei dla ie Ua Eee eo
HamilyebOLAMOGERONACEAE: «). 0/048 tee Gaver a sioee ds eaentaan 8
Cymodocea rotundata Ehrenberg and Hemprich, ex Ascherson . 8
Cymodocea serrulata (R. Brown) Ascherson and Magnus ...... 8
Haloaulepinifolia’ (Muka) den \Hartog (2.2). 225... 202. 22s 13

Halodule uninervis (Forskal) Ascherson ....................-.-. 13
Syringodium isoetifolium (Ascherson) Dandy .................-. 18
Thalassodendron ciliatum (Forskal) den Hartog ................ 18
RannilyallyDROCHARITACEAB (04 br. Ge ah aneG we scradie «eens ste al 23
npalusiacorordesa (legis) pINOV IC Haney fanaa ea Sabie oc tee. 23

[MG OUT DAGGER, ENSONSTON B54 oy sono dhe spud sccugsaes oda phos 24
ialophulami«or (Zollinger) dent Hartogy. 34.4.4. -2. 224.35... 26
Halopiniiaovaliss (Re Brown) Elookerif, ).-.4...--....2..2+4--: 30
Halophila spinulosa (R. Brown) Ascherson .................... 33

Thalassia hemprichu (Ehrenberg) Ascherson .................. 33
iterates Clte Cmte er to ee GCN ay ee eM owa de 39

ll

Soa oft bee Aes ey

: |
5

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|

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;

Seagrasses from the Philippines

Ernani G. Menez, Ronald C. Phillips
and Filconida P. Calumpong

Introduction

The seagrass flora of the western tropical Pa-
cific is unusually diverse (den Hartog, 1970).
Tsuda, Fosberg, and Sachet (1977) reported ten
species for Micronesia; Johnstone (1979) listed 13
taxa for Papua New Guinea; and den Hartog
(1970) reported 11 species for the Phillipines.
Despite the interesting questions associated with
this diversity and the widespread occurrence of
seagrasses in the Philippines, only a few papers
have been published which treat Philippine sea-
grasses. Earlier records include Blanco’s (1837,
1845, 1879) reports of Vallisneria sphaerocarpa (=
Enhalus acoroides) from Zambales. Another report
of Vallisneria sphaerocarpa from Palawan was by
Merrill (1918). Ostenfeld (1909) recorded Halo-
phila ovata (= Halophila minor) from the Philip-
pines, based on Loher’s specimen from Luzon
and later, Merrill’s collection from Manila Bay.
A critical morphological study of Thalassia hem-
prichtt was published by Pascasio and Santos
(1930). Domantay (1962) listed eight species of
seagrasses in his study of the marine vegetation
of the Hundred Islands in Pangasinan. Merrill
(1912, 1915, 1918, 1925), Mendoza and del Ro-
sario (1967) included seagrasses in vascular plant
floras. Calumpong (1979) reported three sea-

Ernant G. Menez, Smithsonian Oceanographic Sorting Center, Na-
tional Museum of Natural History, Smithsonian Institution, Wash-
ington, D.C. 20560. Ronald C. Phillips, School of Natural and
Mathematical Sciences, Seattle Pacific University, Seattle, Washing-
ton, 98119. Hilconida P. Calumpong, Department of Biology, Silli-
man University, Dumaguete City, Negros Oriental 6501, Philippines.

grasses in an ecological study of the sea hare,

Dolabella auriculana (Lightfoot), in Central Vi-

sayas region. Cordero (1981) illustrated and de-

scribed the morphology and distribution of three
species of seagrasses.

The Philippines is an archipelago of about
7,100 islands. It lies between latitudes 4°40’—
21°50’ N and longitudes 116°50’-126°35’ E, sur-
rounded by the South China Sea in the west,
Sulu Sea and Celebes Sea in the south, the Phil-
ippine Sea, which opens into the Pacific Ocean,
in the east, and the Luzon Strait in the north.
The country is naturally divided into four geo-
graphical regions (Figure 1): the Luzon Region,
including the islands of Luzon, Babuyan, Catan-
duanes, Mindoro, Masbate, Romblon, and Mar-
induque; the Mindanao region, including Min-
danao, Basilan, and the Sulu Group; the Visayas
region, including Samar, Leyte, Bohol, Cebu,
Negros, and Panay; and the Palawan region,
including Palawan, Balabac, Culion, and the
Cuyo Group.

Seagrasses are important, but their role has
often been overlooked due largely to their sub-
merged state. Thayer, Wolfe, and Williams
(1975) gave an overall summary of the impor-
tance of seagrasses:

1. Seagrass has a high growth rate, producing an
average of about 300-600 g dry weight/m*/
year, not including root production. Com-
pared to world averages for cultivated corn
(412 g C/m”’) or rice (497 g C/m’), seagrass
beds are more productive.

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

2 2G) LUZON
ee re REGION

PALAWAN
REGION

MINDANAO {ne XQ Mindanao
REGION aiogr sr ee ee
x Ss asilan

Seas

ae

<i °
ee
Sy g

awi-Tawi

Ficure 1.—General map of the Philippines.

NUMBER 21

2. The leaves support a large number of epiphy-
tic organisms, with a total biomass perhaps
approaching that of the seagrass itself.

3. Although a few organisms may feed directly
on the seagrass and several may graze on the
epiphytes, the major food chains are based on
seagrass detritus and its resident microbes.

4. The organic matter in the detritus and in
decaying roots initiates sulfate reduction and
maintains an active sulfur cycle.

5. Seagrass roots bind the sediment together and,
with the protection afforded by the leaves,
surface erosion is reduced, thereby preserving
the microbial flora of the sediment and the
sediment-water interface.

6. Seagrass leaves retard the currents and in-
crease sedimentation of organic and inorganic
materials around the plants.

7. Seagrass absorbs phosphorus through the roots
and the leaves; it may be that the phosphorus
absorbed through the roots is released through
the leaves, thereby returning phosphate from
sediments to the water column. Nitrogen also
is taken up by the roots and transferred to the
leaves and into the medium.

It is now imperative that we properly recognize
how they contribute to the oceanic realm of the
Philippines. All over the world humans and sea-
grasses are coming into conflict for space along
the coast in areas that are in demand for human
activities, from the creation of recreational places,
such as marinas, to industrial ones, such as power
plants, mining sites, freighter and tanker termi-
nals. Unfortunately, discharges of warm water
from power plants and silt and nutrients from
mining, sewage treatment plants, and tanker ter-
minals directly harm seagrasses. Before we de-
stroy or even degrade any more seagrass habitats,
we must know where the seagrasses are and the
true value of this coastal resource.

The resolution of this conflict need not be an
inevitable loss of seagrasses. It is our hope that
this study not only will alert people to the sea-
grasses by allowing them to identify the various
species, but also by making them aware of where
to find them and to appreciate some of the plant
and animal associations that occur within the

2)

seagrass ecosystems. Most importantly, seagrass
communities in the Philippines serve as habitats
and breeding grounds for various marine orga-
nisms that are economically important to the
local populace. Appropriate planning and reason-
able environmental management can ensure the
perpetuation of these ecosystems.

ACKNOWLEDGMENTS.—We wish to thank Dr.
Angel Alcala, Acting President, Silliman Univer-
sity, for allowing us to use the SU Marine Labo-
ratory for our work and for providing accommo-
dations. We appreciate the local transportation
and accommodations provided by the University
of San Carlos, through Prof. Cristobal Plateros,
Chairman, Department of Biology.

Hilconida Calumpong acknowledges the assist-
ance of Mr. Willy Rasay and Ms. Janet Estacion
in collecting and preparation of herbarium spec-
imens of seagrasses. Ronald Phillips appreciates
the financial aid, given by the Seattle Pacific
University Institute for Research, for travel in
March 1982. A portion of this research was sup-
ported by the National Science Foundation,
International Decade of Ocean Exploration, Liv-
ing Resources Program, Seagrass Ecosystem
Study (OCE77-25559). Ernani Menez appreci-
ates the travel and research support in the Phil-
ippines from the Smithsonian Institution Fluid
Research Fund.

For their critical review of this paper, we are
indebted to Dr. Raymond Fosberg, Botanist
Emeritus, Smithsonian Institution, to Dr. C. den
Hartog, Catholic University, Nijmegen, The
Netherlands, and to Dr. Calvin McMillan, Pro-
fessor of Botany, University of Texas at Austin.

The loan of Halophila beccarit and H. spinulosa
specimens from Rijksherbarium, Leiden, is grate- |
fully acknowledged.

MATERIALS AND METHOops.— The seagrasses col-
lected during the Smithsonian Institution Biolog-
ical Expeditions of 1978 and 1979 to Central
Visayas, Philippines, and the authors’ personal
collections during 1980-1982 from the Visayas
region, Palawan, La Union, and Ilocos Norte,
Philippines, were used in this study. Replicates of
material are being deposited in the U.S. National
Herbarium (Smithsonian Institution), the Seattle

4 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

Pacific University Herbarium, the Silliman Uni-
versity Herbarium, the National Herbarium of
the Philippines, and the Rijksherbarium.

In the systematic section of this paper, the
author, date, and page of the original publication
of each taxon, plus basionyms and published
records from the Philippines, are given. The char-
acteristics, natural history, local and world distri-
bution of each species are also given. A dichoto-
mous key and line drawings are included in order
to facilitate easier identification of the twelve
species of seagrasses from the Philippines. Most
of the natural history and distribution data of
Philippine seagrasses in this study are from den
Hartog’s (1964, 1970) publications. Additional
information has been taken from Isaac (1968),
Kay (1971), Kirkman (1975), McMillan (1980a,
b), McMillan, Bridges, Kock, and Falanruw
(1982) and Johnstone (1982).

COLLECTING AND PRESERVING SEAGRASSES.—
Seagrasses may be collected as drift plants cast
ashore or as freshly dug specimens. The major
problems in the use of drift plants is that they
may be broken and their original attachment site
is uncertain. It is known that seagrass plants are
buoyant and can float for weeks if detached from
the bottom.

The recommended procedure for collecting
and preserving seagrass specimens for herbarium
storage and study is as follows. A collector should
get the entire plant including the root and rhi-
zome system. One should use a shovel or dig by
hand deeply enough into the substrate to get
below the rhizome mat. The plant mass can then
be torn loose from the bottom. All sediment and
animals should be washed from the rhizome and
roots. Specimens covering all development phases
should be collected from the same area. It is best
to include plants with flowers, fruits, and seeds,
as well as vegetative plants with intact leaf tips
and root systems.

The plants should be returned to the laboratory
in a plastic bag, preferably in an ice chest. The
plants should be kept moist and should not be
exposed to direct sunlight.

The plants can be preserved by dry-mounting
on herbarium paper or by soaking first in a

solution of seawater formalin. Soaking the plants
first in formalin keeps down the growth of fungi
(mildew) and destruction by insects, but renders
the plants useless for chemical analysis. If one
wishes to kill the plants before mounting on
herbarium paper, the plants should be soaked for
one hour in a 5% solution of seawater-formalin
(8 parts seawater to 1 part formaldehyde). Then
the plants should be drained dry and mounted
on herbarium sheets or other stiff, high-quality
paper.

Seagrasses, such as the small and fragile Halo-
phila species, are most easily mounted on herbar-
ium sheets if they are floated first in a tray of
seawater. A sheet of herbarium paper is placed
under the plant in the water, the plant arranged
on the sheet, the sheet is lifted out and the excess
water drained off. The sheet is then ready for the
plant press (Figure 2).

The plant press can be made from two pieces
of 3/8-inch plywood, each 12 X 17 inches. Inside
the press there should be a complement of cor-
rugated cardboard ventilators, heavy blotters,
and waxed paper, in addition to the herbarium
paper with the specimens. Each of these sheets is
the same size as the plant press. Two or more
straps (belts or pieces of clothesline) are needed
to hold the press together.

The plant press is assembled by placing a
herbarium sheet with its plant specimen on a
blotter. A sheet of waxed paper is placed on top
of the specimen. Another blotter is placed on top
of the waxed paper. Cardboard ventilators are
used to separate blotters. When all of the plants
have been put into the press, the straps are used
to bind the press tightly together. The press may
be placed over a source of mild heat to allow a
current of warm dry air to rise through the cor-
rugations of the ventilators. Until the plants are
dry, the press should be opened each day and dry
ventilators and blotters exchanged for the wet
ones. When the sheets are dry, the dried plants
should be fixed to the herbarium sheets using
clear Duco cement.

The following information should be typed on
a specimen-label form: scientific name (with au-
thor), habitat information (depth of water, type

NUMBER 21

Full Plant Press

Genus/Species:
Habitat Data:
Location:
Date:
Collector:

Herbarium Sheet with Data

SS a | Plywood

( CPP LEP LP EP LLLP EP LAY OP AOL ED PEP ET) Cardboard Ventilator
MME EGET cr asa Blotter
SE EESOOEOr— Waxed Paper
Gest ean Herbarium Sheet with Specimen
————SSSeSeSeSeSe===——E Blotter
GSS SSSSSSSSSSSSSSSSS Cardboard Venti lator

LE Se Plywood

Plant Press Arrangement

Ficure 2.—Illustration showing proper plant press arrangement for drying seagrasses.

6 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

of substrate, etc.), location (specific but also gen-
eral area), date of collection, collector’s name.
The label should be pasted in the lower right
corner of the sheet.

General Features of Seagrass Biology and
Ecology

The narrow fringe of the edge of the sea is both
the most productive and the most threatened
portion of the ocean. In this area of the shallow
water light can penetrate to the bottom and
nutrients arrive from both land and sea. It is here
that marine plant ecosystems flourish. These ma-
rine systems are the foundations of inshore pro-
ductivity and form the basis for animal life.

The value and significance of the shallow-water
habitats have been overlooked in the past, in part
because the connections from the plants to ani-
mals are complex and often indirect. Paradoxi-
cally, the most important contribution of all sea-
grass ecosystems to the chains of life is death. The
bacteria that devour the dead leaves of the plant
not only nourish and enrich the area in which the
plants grow, but also affect waters for miles
around. Seagrass and mangrove leaves have been
found in deep-sea trenches far from shore and
thousands of feet deep. Even in these most remote
regions of the sea, the leaves provide food and
shelter for resident animals.

Seagrasses are not true grasses, but they do
resemble them and have many similarities with
them in the biology of their life cycle. Seagrasses
usually form dense meadows that resemble more
familiar prairies or grasslands. Seagrasses are vas-
cular plants. The vascular system in the seagrasses
consists of an internal network of tubes that is
continuous from the roots to the leaf tips, trans-
porting water, nutrients, and gases for tissue
growth. Because of this system, seagrass growing
in sand or mud can use the highly concentrated
nutrients of the bottom, nutrients that may be
hundreds or thousands of times more concen-
trated than those in the overlying water column.
This source, the bottom sediment, is unavailable
to all other marine plants; these others must
depend for growth only on nutrients dissolved in
the water. Seagrasses can absorb nutrients from

water through their leaves like algae, but are not
limited to that source. Seagrasses, then, are a
mechanism for returning nutrients to other ma-
rine life that would otherwise be lost in the sedi-
ments. They are the only plants in the sea that
can do this. The significance of this dual nutrient
source is reflected in growth rates that are high
or higher than those of any other plant commu-
nity in the sea. The roots and rhizomes not only
absorb nutrients from the sediment, but also pre-
vent the movement of bottom sediments. In dense
meadows of Thalassia hemprichiu (turtle grass) and
Enhalus acoroides, the roots and rhizomes form a
dense mat that prevents erosion of the coast that
would otherwise occur from strong currents and
severe typhoons. Since they occur inshore of the
coral fore-reef, they act in concert with the reef in
coastal protection.

Part of the importance of seagrasses is their
high productivity. Seagrass beds have been shown
to be among the most productive areas of the
earth, including those under intensive agricul-
ture. Seagrasses have a complement of attached
algae, called epiphytes, which contribute to the
high productivity of the seagrass ecosystem. This
primary productivity is the first link in the food
chains of these ecosystems, and generally supports
a high productivity in the associated animal com-
munity.

Seagrasses constitute a haven and refuge for
animals in the sea. They are especially important
as nurseries for fish, shrimps, lobsters, scallops,
and other valuable food animals. The protection
that these plant systems afford extends beyond
their associated animal life to the coast itself.
Although many of the animal associations with
seagrasses are indirect and inconspicuous, there
are notable exceptions: dugongs, sea turtles, some
fish, and sea urchins and other invertebrates feed
directly on these plants.

A seagrass meadow or bed may consist of a few
hundred to a few thousand leafy shoots per square
meter and can cover bottom areas from a few
acres up to many square miles. These meadows
are crucial habitats for a variety of marine life.
To some animals they are a refuge; to others they
are the local swim-in restaurant where a free
lunch is always available. One of the reasons for

NUMBER 21

this concentration of marine life in seagrass
meadows is the great habitat complexity resulting
from the dense canopy of leaves; the leaves pro-
vide a surface area that can be 20 or more times
greater than that of the bottom they occupy — a
veritable jungle. A myriad of small animals and
plants, some epiphytes and some merely clinging,
can be found on seagrass leaves. These form an
essential part of the complex food webs that exist
in a seagrass meadow.

An essential feature of any seagrass meadow is
the fate of the leaves as they age and become
detached from the leaf stalk, an event similar to
the autumn drop of leaves in a forest. A large
proportion of these leaves sink to the bottom
where they become the basis of yet another com-
plex food web. Through wave action and brows-
ing activities of invertebrates, the leaves break
down and decompose into particulate matter that
remains suspended in the water column. In so
doing, they provide inorganic nutrients that are
essential to the continued growth of the living
seagrasses, and nourishment to a variety of bac-
teria. Some of the soluble plant chemicals leach
out into the surrounding water and help to feed
the same bacteria. These bacteria are the food of

al

diverse species of small animals that are them-
selves the food for slightly larger species of ani-
mals and so it goes. This decomposer food chain
begins with bacteria in the upper levels of bottom
sediments and around the surface of the particu-
late plant matter, an environment devoid of ox-
ygen, but eventually it leads to the more fa-
miliar larger food animals, worms, clams, shrimps
and consequently to fishes. The steps in this food
chain are still more obscure than the one based
on direct grazers and there is still doubt about its
significance in the economy of the sea. The pro-
duction of detritus from the constant sloughing
of leaves in the seagrass meadow results in a
relatively constant source of food for the bacteria
and animals of the decomposer food chain
throughout the year.

Many studies in the United States have proven
that seagrass presence and growth is absolutely
dependent on relatively clear water. Seagrasses
stabilize bottom sediments and baffle water mo-
tion inside the meadow, but additions of dirty
water from dredging, mining, sewage, and boat-
ing activities can sharply curtail or eliminate the
productivity or even presence of seagrass in an
area.

Key to the Philippine Seagrasses

Wee eavesmtenete: puke cis ee

1 LPSaKES TAN oo os Se aca, 5 5 6 be Ros aloe eee nee: See ee a 2

2. Leaves elliptic, ovate, lanceolate or linear-oblong; 4—30 mm long
doa 6 diggs 9 66g oR Aerees es oie AeA Snene nS eet nnn Ree ere Halophila
Pemleaves lanceolate wiAthOUnlelOSS-VEINSE 9. . 1.5456 fe. oe ee H. beccarii
eayes@tnenwiSe)WitheCrOSS-VEINS si 445.57 F ej dae ochales bec ne ae. B

BS eaves with 2-22 pairs of cross-velms 2... 2.) 25.5. 522808 te H. ovalis
CAVES RW AUN = APAITS Ol! CLOSS= VEINS geiko alge ys Seid ene Ne oie Aesstaoe C

C. Leaves up to 22 pairs, distichous on long shoot .......... H. spinulosa
eavesinepairs omevery short Shoot. . yi eos ic. spe ek ei H. minor
eavesviimearmsnally5—1OO"emmMlong 23008 shee ge tee Nene SE 3
IRC AV ESM GU ALC MMR ec P ee Er. hes ee eae aS eA OED oe os ate RS 4
ecyy CSB MNO UN Ate rt orc Cl) ae Dr ye Se A 8 ok iS 6

Ae Nerves monmorewthane ae gic isco Bis. Ml ihe te ekg SMe Bees + Halodule

A. Leaf tip with irregular serrations; tip of midrib often furcate .........

Leaf tip tridentate; tip of midrib not furcate
INEnVesmmOrennalyo et oa) a. ae.

BAS fre one a Te es are ee H. pinifolia

Be has een H. uninervis

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

5. Unbranched or once-branched shoots produced at every fourth internode
on rhizome; leaf tips with bi- trifurcate teeth; roots borne on internode
with numerous tough, wiry laterals; internodes 3-10 mm _ long

; SS ee Thalassodendron ciliatum
Beare Pee em 9h. Cymodocea

A. Leaf tip smooth or slightly serrulate; leaf blades linear . C. rotundata

Leaf tip serrate-dentate; leaf blades linear-falcate

Dane C. serrulata

6. Rhizome 2-4 mm in diameter; roots densely beset with filiform laterals;

leaves linear-falcate; without basal black fibers

.. Thalassia hemprichii

Rhizome more than | cm in diameter; roots naked; leaves linear; with

persistent basal black fibers ......

Division ANTHOPHYTA
Class MONOCOTYLEDONEAE
Order HELOBIAE
Family POTAMOGETONACEAE

Cymodocea rotundata Ehrenberg and
Hemprich, ex Ascherson

FiGuRES 3A,B, 4

Cymodocea rotundata Ehrenberg and Hemprich, ex Ascherson,
1871a:83.—Merrill, 1925:24.—Mendoza and del Rosario,
1967:12.—den Hartog, 1970:166, fig. 47.

CHARACTERISTICS.—Plants of moderate size,
with moderately robust rhizomes, reaching 2 mm
in diameter; internodes 0.5-3 cm long; laxly
branched roots usually 1-3 per node. Erect shoots
short, generally with 2-4 leaves. Persistent leaf
sheaths up to 4 cm long, usually forming scarious
mass at maturity; when shed, conspicuous scars
are left on stem. Leaf blades linear, 2-4 mm wide
and up to 10 cm long, with round apices or
slightly serrulate, nerves 8-12.

NaTuRAL History.—Cymodocea rotundata occurs
in shallow water, on sand-mud or sand substrates
in sheltered coves or bays, lagoons, mouth of
rivers and coral reefs. The plant may be found
growing with Halophila ovalis, Halodule uninervis,
Enhalus acoroides, Thalassia hemprichii, Syringodium
isoetifolium, or Cymodocea serrulata. Epiphytic algae
found on leaves of the plant include melobesioids
and Hypnea. In New Caledonia and Thursday

| PRISE et Soa are Enhalus acoroides

Island, staminate flowers were observed during
April and November, respectively. Pistillate flow-
ers have been reported during November at
Thursday Island. At Bristow Island, both stami-
nate and pistillate flowers were found in March.
Plants with fruits were recorded from Yap,
Tomil-Gagil, and Map islands in Micronesia dur-
ing March and June-July. In New Guinea and
the island of Bali, fruits were observed in June.

PuILipPINE DistriBuTION.—Luzon (Lingayen
Gulf, Catanduanes, Mindoro); Palawan (Cuyo
Islands); Visayas (Samar, Negros, Siquijor); Min-
danao (Pearl Bank, Tawi-Tawi Group).

RanceE.—C. rotundata is widely distributed in
the Indian and western Pacific oceans. It occurs
in Egypt, Sudan, Kenya, Tanzania, Mozam-
bique, Madagascar, Yemen, Thailand, Andaman
Islands, Ryukyu Islands, Malaysia, Indonesia,
Papua New Guinea, Caroline Islands, Queens-
land, and New Caledonia.

Cymodocea serrulata (R. Brown) Ascherson
and Magnus

FicurREs 5A-c, 6

Caulinia serrulata R. Brown, 1810:339.

Cymodocea serrulata (R. Brown) Ascherson and Magnus, in
Ascherson, 1871a:84.—Meerrill, 1925:24.—Mendoza and
del Rosario, 1967:13.—den Hartog, 1970:171, fig. 48.

CHARACTERISTICS.—Plants of moderate size,
rhizomes 2-3 mm in diameter; internodes 1-4 cm
long; usually 2 to several sparsely branched roots
per node. Erect shoots short, bearing up to 4

NUMBER 21

Figure 3.—Cymodocea rotundata: a, habit of plant, X 1.4; B, upper portion of a leaf showing
perpendicular veins, X 4.4.

10

SMITHSONIAN CONTFIBUTIONS TO THE MARINE SCIENCES

Figure 4.—Cymodocea rotundata: top, world distribution; bottom, Philippine distribu-
tion.

NUMBER 21

Ficure 5.—Cymodocea serrulata: A, habit of plant, X 1.2; B, upper portion of a leaf showing
serrate tip and perpendicular veins, X 2.2; c, fruit, X 1.8

2

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

Figure 6.—Cymodocea serrulata: top, world distribution; bottom, Philippine distribu-
tion.

NUMBER 21

leaves. Leaf sheaths 1-3 cm long, tapered towards
the base; when shed, leaving conspicuous scars
around the stem. Leaf blades linear-falcate, 3-7
mm wide, up to 9 cm long, narrowed at the base;
with round to obtuse, serrate leaf tips; margins
mostly entire, spinulose only near the tip; having
8-16 nerves connected by cross-veins. Fruits
nearly elliptic, compressed, sessile and with dorsal
ridges.

Naturat History.—Cymodocea serrulata occurs
in the lower intertidal zone, in sheltered localities,
on coarse coral-sand, sand-mud, or mud with
coral rubble substrates. The plant is reported
growing together with Thalassia hemprichit, Enhalus
acoroides, Halophila ovalis, Halodule uninervis, and
Syringodium isoetifolium. Pistillate flowers have been
reported during April in New Caledonia; in
Kenya, they have been observed in January and
August. Plants with staminate flowers have been
recorded during February and March from Mor-
eton Bay, Queensland. In Kenya, they occurred
in September. Contrary to den Hartog’s (1970)
comment that Cymodocea serrulata is absent in
places where there is freshwater influence, in
Albay Gulf, southeastern Luzon, Philippines, the
plant was collected from boulders near the mouth
of a stream, growing together with Halodule uni-
nervis. Epiphytic algae found on the leaves of
Cymodocea serrulata were melobesioids, Ceramium
and Acrochaetium. In Negros, Philippines, fruits
were found on Cymodocea serrulata in August.

PuivipPpINE DistripuTiIoN.—Luzon (Pangasi-
nan, Albay, Catanduanes, Mindoro); Palawan
(Bajallanura Island, Double Island); Visayas (Ne-
gros, Leyte); Mindanao (Zamboanga).

RanceE.—C. serrulata is common in tropical and
subtropical regions. It occurs in Egypt, Sudan,
Kenya, Tanzania, Mozambique, Madagascar,
Comoro Islands, Seychelles, Yemen, Sri Lanka,
Malaysia, Indonesia, Western Australia, Queens-
land, Papua New Guinea, Caroline Islands, and
New Caledonia.

Halodule pinifolia (Miki) den Hartog
FIGURES 7A,B, 8

Diplanthera pinifolia Miki, 1932:787.
Halodule pinifolia (Miki) den Hartog, 1964:309; 1970:158.

13

CHARACTERISTICS.—Plants small, with slender
rhizomes, up to 1.5 mm in diameter; internodes
usually 0.5-2.0 cm long; simple or laxly branched
roots, 2 to several per node. Short, erect shoots
consisting of 2-4 leaves borne at each node. Leaf
blades linear, 4-15 cm long, not more than 1.5
mm wide. Leaf tips obtuse with few irregular
serrations. Leaf margins entire; conspicuous mid-
rib furcate at the tip. Basal portion of leaves
enclosed by sheaths, 3 cm long. Ovate scales on
each node not persistent at maturity.

NaTuRAL History.—Halodule pinifolia occurs in
sheltered and semi-exposed bays, coral reef plat-
forms, in pools, also in wave-beaten sites. The
plant has only been collected from two sites along
the coast of Negros and Samar islands of the
Visayas region of the Philippines. Those collected
from Negros were from the upper sublittoral zone
and were sterile. The rarity of this species in our
collections suggests that it is less tolerant of pre-
vailing hydrographic parameters compared to
other seagrasses. In the literature (den Hartog,
1970), its companion species on compact mud is
Cymodocea rotundata and on soft mud, Halophila
ovalis. At Cairns, Queensland, flowering and fruit-
ing plants of Halodule pinifolia were found during
November. Species of crustose melobesioids Acro-
chaetium, Ceramium, Hypnea, and Polysiphonia, were
found growing on leaves of Halodule pinifolia.

PHILIPPINE DisTrRiBUTION.—Luzon (Mindoro);
Visayas (Negros, Samar, Bohol); Mindano (Cav-
illi Island, Pearl Bank).

Rance.—H. pinifolia is most commonly distrib-
uted in tropical and subtropical localities. It oc-
curs in Ryukyu Islands, Vietnam, Indonesia,
Tonga Islands, Papua New Guinea, Fiji Islands,
Queensland and New Caledonia.

Halodule uninervis (Forskal) Ascherson
Ficures 9a,B, 10

Zostera uninervis Forskal, 1775:157.

Halodule uninervis (Forskal) Ascherson, in Boissier, 1882:24.—
den Hartog, 1970:147.

Diplanthera uninervis Ascherson, in Engler and Prantl,
1897:37.—Domantay, 1962:275.—Mendoza and del Ro-
sario, 1967:13.

CHARACTERISTICS.—Plants of moderate size,

14

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

Ficure 7.—Halodule pinifolia: a, habit of plant, X 2.1; B, upper portion of leaf showing furcate
midrib and irregular teeth at tip, X 28.

| NUMBER 21

FicureE 8.—Halodule pinifolia: top, world distribution; bottom, Philippine distribution.

15

16 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

B IBS

Ficure 9.—Halodule uninervis: a, habit of plant, X 0.95; B, leaf tip showing two acuminate teeth
and median nerve, X 10.4.

with slender rhizomes barely reaching 2 mm in __ below by a 1-2 cm long leaf sheath; when shed,
diameter; internodes 2-3 cm long; unbranched to _ leaf sheaths leave conspicuous annular scars on
sparsely branched roots not more than 5 pernode. — stem. Leaf blades linear and narrow at the base,
Erect shoots short, with 2-4 leaves enveloped 2-5 mm wide, reaching 15 cm in length. Blades

NUMBER 21

Ficure 10.—Halodule uninervis: top, world distribution; bottom, Philippine distribu-
tion.

17

18 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

with 3 nerves; the middle one very distinct and
ending in the median apical obtuse tooth, while
the lateral nerves ending in 2 abruptly acuminate
teeth. Leaf tip tridentate; leaf margin entire.

Natura History.—Halodule uninervis occurs in
sheltered or exposed localities in the shallow in-
tertidal to the upper subtidal zone, on sand-mud,
coarse coral-sand, or sand substrates. In the Phil-
ippines, occasional patches of the plant may be
mixed with Thalassia hemprichu, Halophila ovalis,
Syringodium isoetifollum, Enhalus acoroides, and Cy-
modocea serrulata. On the leaves and particularly
the basal parts of Halodule univervis, epiphytic
algae, such as Colpomenia sinuosa,, Dictyopterts deli-
catula, Jama adherens, Mastophora rosea, and species
of Ceramium, were found. Flowering and fruiting
plants of Halodule uninervis have not been observed
to date in the Philippines. At Queensland, flow-
ering plants were found during October and No-
vember.

PHILIPPINE DistripuTion.—Luzon (Manila
Bay, Pangasinan, Albay, Catanduanes, Mindoro,
La Union, Ilocos Norte); Palawan (Cuyo Islands);
Visayas (Negros, Bohol, Leyte, Cebu); Mindano
(Zamboanga).

Rance.—Halodule uninervis is widely distributed
along the tropical coast of the Indian and western
Pacific oceans. It occurs in Egypt, Eritrea, So-
malia, Tanzania, Madagascar, Mozambique,
Mauritius, Seychelles, Saudi Arabia, Kuwait,
Bahrain Islands, Yemen, Sri Lanka, Andaman
Islands, Thailand, Vietnam, Malaysia, Indonesia,
Papua New Guinea, Western Australia, Queens-
land, New Caledonia, Tonga Islands, Caroline
Islands, Mariana Islands, and Japan.

Syringodium isoetifolium (Ascherson) Dandy
Ficures 11aA—p, 12

Cymodocea isoetifolia Ascherson, 1868:163.
Syringodium isoetifolum (Ascherson) Dandy, in Dandy and
Tandy, 1939:116.—den Hartog, 1970:177, figs. 50, 51a—d.

CuaraActeERIstics.—Plants of moderate size,
with slender rhizomes not more than 1.5 mm in
diameter; internodes 3-15 mm long; one to sev-
eral laxly branched roots below the erect shoots.

Erect shoot consisting of 2-3 terete leaf blades per
node. Leaf blades reaching 25 cm long, 1.0-2.5
mm in diameter, narrow towards the base; central
vascular bundle encircled by 6-11 air channels;
pericentral vascular bundles 7-9. Leaf sheaths up
to 5 cm long. Inflorescence cymose; sheaths of
reduced leaves in the inflorescence mostly 4-7
mm long. Staminate flowers with 2 ovate anthers,
4 mm long and connected by the middle to the
apex of a 2 mm, terete stalk.

Natura Hisrory.—Springodium isoetifolium oc-
curs in sheltered and semi-exposed habitats, on
coarse coral-sand substrate. The plants are found
in the upper subtidal zone, forming thick carpets
with Thalassia hemprichiu, Cymodocea rotundata, C.
serrulata, and Halodule uninervis. In the Philippines,
pistillate and staminate flowers were found dur-
ing May. Crustose melobesioids sometimes heavy
on leaf blades; Chaetomorpha crassa, Tolypiocladia
glomerulata, Hypnea valentiae, Dictyota species, and
Gracilaria blodgettu attached at the base of leaves
and on leaf sheaths of S. zsoetzfolium.

PuivipPpInE Distripution.—Luzon (Pangasi-
nan, Albay, Catanduanes, Mindoro, La Union);
Visayas (Negros, Siquijor, Bohol); Mindanao
(Zamboanga).

RANGE.—Syringodium isoetifolium is widely dis-
tributed in the tropics. It occurs in Egypt, Kenya,
Tanzania, Mozambique, Madagascar, Sey-
chelles, Mauritius, Yemen, Bahrain Islands, Sri
Lanka, Nicobar Islands, Vietnam, Malaysia,
Papua New Guinea, Tonga Islands, Caroline Is-
lands, Ryukyu Islands, Western Australia,
Queensland, and New Caledonia.

Thalassodendron ciliatum (Forskal) den
Hartog

Ficures 13a-c, 14

Zostera ciliata Forskal, 1775:157.
Thalassodendron ciliatum (Forskal) den Hartog, 1970:186, figs.
52a—j.—Menez and Calumpong, 1983:103-111.

CHARACTERISTICS.—Plants moderately large,
occasionally branched once, reaching a height of
45 cm. Rhizome thick, up to 5 mm in diameter;
internodes 3-10 mm long; roots 1-6 borne on

NUMBER 21

ae Seager s
> a

Ficure 11.—Syringodium isoetifolium: a, habit of plant, X 0.3; B, staminate flower with
2 anthers, X 4.1; c, portion of male plant, X 3.1; D, section of leaf-blade showing
central vascular bundle and air channels, X 21.8.

19

20

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

Figure 12.—Syringodium isoetifolium: top, world distribution; bottom, Philippine distri-
bution.

22

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

Ficure 14.—Thalassodendron ciliatum: top, world distribution; bottom, Philippine dis-
tribution.

i
1)

NUMBER 21

internode, with tough, wiry laterals, no more than
1.5 mm in diameter. Two stems produced at
every fourth internode on rhizome; one fully de-
veloped, long and erect with numerous leaf scars
and another, usually an undeveloped stem-bud.
Upper part of erect stem compressed, but the
basal portion terete. Leaf blades usually 6, linear-
falcate, narrowed towards the base, up to 10 mm
wide and 13 cm long; leaf tips rounded, dentic-
ulate; apical teeth bi- trifurcate; upper half of
leaf margin with irregular serrations, becoming
distant and few towards the base; lower half of
leaf margin almost entire. Leaf nerves 12-18,
connected by a few oblique cross-veins. Leaf
sheaths obtuse, up to 12 mm wide and 20 mm
long, with obtuse auriculae and ligules.

NaTuRAL History.— Thalassodendron ciliatum oc-
curs in sheltered and semi-exposed localities, in
the upper sublittoral zone. The plants from the
Philippines were sterile and found in small
patches among dead and living corals in clear
water. In Kenya and Mozambique, staminate
and pistillate flowers were observed during Au-
gust and January, respectively. In Tanzania,
staminate flowers have been recorded during Au-
gust. Algal epiphytes found growing on Philip-
pine specimens of T° cilzatum include Hypnea val-
entiae, Jania adherens, Amphiroa rigida, Codium arabi-
cum, Giffordia rallstae, Sphacelaria furcigera, Polysi-
phonia mollis, species of Padina and Dictyota, Cer-
amium mazatlanense, and Champia parvula. The latter
two algal taxa were present on both leaves and
stems, while the rest were only on the stem.

The authors wish to point out that Thalassoden-
dron ciliatum may often be confused with the “‘long-
stemmed” Cymodocea serrulata. Johnstone (1982)
has found two growth forms of Cymodocea serrulata
in New Guinea and reported that the “normal”
form to be the most common. McMillan (1983)
has found evidence that the “long-stemmed”
plants are controlled by light intensity. 7. czliatum
and C. serrulata are easily distinguished by a care-
ful morphological examination.

PHILIPPINE DistRIBUTION.— Palawan (Bajallan-
ura Island, Tidepole Island, Double Island, west-
ern side of Tabon Cave at Quezon).

Rance.—T. ciliatum occurs in Egypt, Sudan,

73)

Somalia, Kenya, Tanzania, Mozambique, South
Africa, Madagascar, Chagos Archipelago, Sey-
chelles, Amirante Islands, Aldabra Islands, Com-
oro Island, Mauritius, Saudi Arabia, Iran, Indo-
nesia, Papua New Guinea, Solomon Islands, Car-
oline Islands, and Queensland.

Family HyDROCHARITACEAE

Enhalus acoroides (L. f.) Royle
FicureE 15, 16

Stratiotes acoroides L. f., 1781:268.

Vallisneria sphaerocarpa Blanco, 1837:780; 1845:538; 1879:
186.—Merrill, 1918:59.

Enhatus acoroides (L. f.) Royle, 1840:453.—Merrill, 1925:27.—
Domantay, 1962:275.—den Hartog, 1970:215, fig. 60.—
Cordero, 1981:321, fig. 1.

CHarRACTERISTICS.—This is the largest of the
seagrass species from the Philippines, reaching a
height of more than a meter. Rhizome thick, up
to 1.5 cm in diameter, deeply embedded in sub-
strate, bearing many roots, 2-5 mm in diameter
and up to 15 cm long. Persistent leaf fibers very
dense and masking rhizome. Leaves 3 or 4, pro-
duced directly from rhizome; leaf blades linear,
to 1.5 cm wide and reaching | m in length; leaf
tips rounded, beset occasionally with minute ser-
rate projections when young; leaf margins entire,
slightly rolled. Characteristic tough, black vas-
cular bundles persistent after death and decay of
other leaf tissues. Nerves 10-40, parallel to about
as many air-channels. Fruits ovoid, 4-7 cm long,
with acuminate tips and densely covered with
bifid projections. Fruit stalk long and coiled.

Natura History.—Enhalus acoroides occurs in
sheltered localities in the lower intertidal and
upper subtidal zones, on sand-mud, coarse coral-
sand or mud substrates. In the Philippines, the
plant was observed in pure stands next to a
mangrove where silt was thick and water was
murky, or in occasional patches in coarse coral-
sand growing together with Thalassia hemprichiz,
or mixed with Halodule uninervis, Cymodocea serru-
lata, Syringodium isoetifolium, and Halophila ovalis.
Fruiting plants of Enhalus acoroides were found in
May-October in Central Visayas and during

24

|

=
Se

<s
=

SS

Ficure 15.—Enhalus acoroides: habit of plant, X 0.4.

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

April staminate flowers were collected from
southern Luzon in the Philippines. According to
den Hartog (1970), this plant probably flowers
and fruits the whole year round. Crustose melo-
besioids are sometimes very thick on leaves and
on black “hairs.” Hypnea and Cladophora were
found attached to the melobesioids. Few thalli of
Ceramium were epiphytic on Enhalus acoroides
leaves.

PuiLipPINE DistriBuTION.—Luzon (Zambales,
Catanduanes, Mindoro); Palawan (Tay-tay Bay,
Quezon, Cuyo Islands); Visayas (Cebu, Samar,
Leyte, Negros, Bohol); Mindanao (Surigao, Zam-
boanga, Basilan).

Rance.—E. acoroides is widely distributed in
the tropics. It occurs in Kenya, Seychelles, Mad-
agascar, Saudi Arabia, Yemen, India, Sri Lanka,
Andaman Islands, Malaysia, Kampuchea, Viet-
nam, Ryukyu Islands, Indonesia, Papua New
Guinea, Mariana Islands, Caroline Islands,
Queensland, and New Caledonia.

Halophila beccarii Ascherson
Ficures 174,B, 18

Halophila beccarii Ascherson, 1871b:302.—Merrill, 1912:70;
1925:25.—Mendoza and del Rosario, 1967:21.—den Har-
tog, 1970:261.

CHARACTERISTICS.—Plants small, with slender
rhizomes not more than | mm in diameter; inter-
nodes 1-3 cm long; with one root at each node.
Uneven pair of scales enveloping the erect shoot
that consists of up to 10 closely set, petiolated
leaves. Leaf blades lanceolate, with acute apices
and cuneate or attenuate bases, 8-15 mm long
and 1-2 mm wide; margins entire, rarely mi-
nutely spinulose. Conspicuous midrib extending
to leaf apex, with one intramarginal vein on each
side that is connected only at the base of the
midrib and joining it again just below the tip.
Cross-veins absent.

Natura History.—Halophila beccartt occurs in
the shallow, intertidal zone in sheltered localities,
such as mangroves, mouths of rivers, and brackish
ponds. The plant seems to favor sand-mud, par-
ticularly muddy substrates. In Singapore, samples
of the species were found abundant on mud and

NUMBER 21

Ficure 16.—Enhalus acoroides: top, world distribution; bottom, Philippine distribution.

25

26 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

Rex i,
ATR &>

Ficure 17.—Halophila beccari: a, habit of plant, X 2.9; B, leaves showing entire and spinulose
margins, including conspicuous midrib and intramarginal veins, X 7.3.

on sand in shaded stream beds, and in brackish
water in mangroves. Flowering of Halophila bec-
carut have been observed during February, June,
and August. The plant may be found growing in
pure stand or together with Halophila ovalis, H.
minor, or Halodule uninervis. A few patches of crus-
tose melobesioids were observed growing on
leaves of Halophila beccaru collected from the Phil-
ippines.

PHILIPPINE DistrrpuTion.—Luzon (Manila
Bay, Paranaque).

Rance.—H. beccarit occurs in Chilka Lake, a
salt-water lake along the eastern coast of India. It
has also been recorded in Sri Lanka, Burma,
Vietnam, and Malaysia.

Halophila minor (Zollinger) den Hartog
Ficures 19a—c, 20

Lemnopsis minor Zollinger, 1854:75.
Halophila minor (Zollinger) den Hartog, 1957:410.—Mendoza
and del Rosario, 1967:21.

NUMBER 21

Ficure 18.—Halophila beccarit: top, world distribution; bottom, Philippine distribution.

27

28

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

Ficure 19.—Halophila minor:a, habit of plant, X 4.9; B, leaf showing cross veins, X 9.7; c, scale,
x 29.1.

\
NUMBER 21

Ficure 20.—Halophila minor: top, world distribution; bottom, Philippine distribution.

99

30 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

Halophila ovata sensu Ostenfeld, 1909:68.—sensu Merrill,
1912:70; 1925:26.—sensu den Hartog, 1970:251. [Non
Gaudichaud, 1826 [1829]:430 (= H. ovalis (R. Brown)
Hooker f.).]

CHARACTERISTICS.— This is probably the small-
est of the seagrasses found in the Philippines, with
rhizomes barely reaching 1 mm in diameter;
internodes 7-20 mm long; usually one root below
each erect shoot. Erect shoot short, consisting of
a pair of leaves on each node. Leaves with pe-
tioles, 2-8 mm long, each enveloped by a pair of
transparent, orbicular scales with rounded apices.
Leaf blades elliptic or obovate, width 2-3 mm,
length 4—7 mm, apices rounded, bases attenuate;
margins entire; with 4-7 pairs of cross-veins.
Staminate flowers pedicellate, with obtuse, con-
vex tepals; anthers approximately 2 mm long.
Pistillate flowers with 3 styles, each up to 15 mm
long; ovary ovoid, reaching a length of 3 mm.
Ellipsoid, ovoid or globose fruits up to 4 mm long,
bearing 2-6 mm long beaks; seeds subglobose,
about 0.5 mm long.

The authors follow the remarks of Sachet and
Fosberg (1973:441) that the correct name for this
species is Halophila minor, since Halophila ovata is a
superfluous name, hence illegitimate.

NaturRAL History.—Halophila minor occurs in
sheltered coasts on mud, sand, or coral rubble
substrates. The plant seems to prefer muddy bot-
toms and tolerates heavy sedimentation, as gen-
erally found in deep bay localities that are well
protected from wave action. In the Philippines,
H. minor plants were collected from sand-mud or
coral rubble substrates during January—May.
They were abundant in March—May and bearing
flowers and fruits. Flowering plants have been
observed in January and in April, including
fruits, at New Caledonia. Plants with fruits and
flowers were collected from Kenya in July. Den
Hartog (1970) suggested that in areas where the
plants are reproductive, they are annuals, and in
places where they do not flower, they may be
perennials. Halophila minor have been found grow-
ing together with Halophila ovalis, Enhalus acoroides,
Thalassia hemprichu, and Halodule uninervis. Exam-
ination of epiphytes revealed few patches of crus-

tose red algae on leaves of H. minor from the
Philippines.

PHILIPPINE Distripution.—Luzon (Manila
Bay, Albay, Mindoro); Visayas (Cebu, Negros).

RanceEs.—Halophila minor occurs in Kenya, In-
dia, Singapore, Indonesia, Papua New Guinea,
Western Australia, Queensland, New Caledonia,
Samoa, Fiji, Wallis Islands, Caroline Islands,
Mariana Islands, Hawaii.

Halophila ovalis (R. Brown) Hooker f.

Ficures 21a-—c, 22

Caulina ovalis R. Brown, 1810:339.

Halophila ovalis (R. Brown) Hooker, 1858:45:—Merrill,
1925:26.—Domantay, 1962:275.—Mendoza and del Ro-
sario, 1967:22.—den Hartog, 1970:240.—Cordero, 1981:
322;

Halophila ovata Gaudichaud, 1826 [1829]:430.

CHARACTERISTICS.— Plants small, with long and
narrow rhizomes, not more than 2 mm in diam-
eter; internodes 5-40 mm long; usually one root
present below each erect shoot. Erect shoot short,
consisting of a pair of leaves borne on each node.
Leaves petiolated; petioles 5-50 mm long, each
enveloped by a pair of transparent scales; apex of
scale emarginate, the base auriculate. Leaf blades
oblong, oval, elliptic or spathulate, occasionally
lanceolate or linear, width 4-12 mm, length 8-
25 mm, apices rounded, bases cuneate, attenuate
or truncate; margins entire; with 12-22 pairs of
cross-veins which are often forked. Midrib con-
nected to the intramarginal nerve at the top.
Staminate flowers consist of small, elliptic, convex
tepals generally obtuse or slightly apiculate; an-
thers oblong. Pistillate flowers with small ovoid
ovaries; 3 long styles. Fruits globose with beaks
up to 6 mm long.

NatuRAL History.—Halophila ovalis occurs in
open or sheltered coasts, from midtide to 10 m
deep in Central Philippines. The plant grows on
a wide variety of substrate, such as mud, living
corals, or coral rubble, and in sand where the
plant is occasionally almost completely buried.
According to the literature, H. ovalis is a euryha-
line species that can grow in waters with de-

1
]
|
}

NUMBER 21

Ficure 21.—Halophila ovalis: a, habit of plant, X 2.2; B, leaf showing cross veins, X 5.6; c, scale,
X< 10.4.

3]

32

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

Ficure 22.—Halophila ovalis: top, world distribution; bottom, Philippine distribution.

NUMBER 21

creased salinities of 10%o, as in estuaries and
brackish lagoons. It is also eurythermic, but can-
not survive in waters with temperatures below
10° C. There is a great variability in its mor-
phology (i.e., leaf size and shape, plant size, ner-
vation) which is probably influenced primarily
by its environment. In den Hartog’s (1970) view,
H. ovalis represents a “collective species” and only
H. ovata (= H. minor) had been separated from it
(Ostenfeld, 1909).

Halophila ovalis is an abundant seagrass in trop-
ical and warm temperate areas, occasionally
found in pure stands or mixed with Thalassia
hemprichi, Halodule uninervis, H. pinifolia, Cymodocea
rotundata, or Enhalus acoroides. On a sand-bar on
the northwestern end of Sumilon Island, Philip-
pines, it was observed that Halophila ovalis ap-
peared as a pioneer species on new sand-coral
rubble substrate. The plant flowers and fruits
during the warmest months. Algal epiphytes
found on the leaves of the plant include crustose
melobesioids, a few thalli of species of Acrochae-
tum, Ceramium, Hypnea, Janta, and Polysiphonia.

PHILIPPINE DisTRiBUTION.—Luzon (La Union,
Catanduanes, Pangasinan, Albay, Mindoro,
Masbate); Palawan (Cuyo Islands, Quezon); Vi-
sayas (Leyte, Negros, Cebu, Panay, Sumilon, Si-
quijor, Bohol, Samar); Mindanao (Davao, Zam-
boanga, Bancoran Island).

Rance.—Halophila ovalis is widely distributed
in the Indo-West Pacific. It occurs in Egypt,
Sudan, Kenya, Tanzania, Mozambique, South
Africa, Madagascar, Seychelles, Mauritius, Israel,
Saudi Arabia, Yemen, Kuwait, Iran, Bahrain
Islands, Sri Lanka, Burma, Andaman Islands,
Thailand, Vietnam, Hong Kong, China, Ryukyu
Islands, Malaysia, Indonesia, Papua New Guinea,
Western Australia, South Australia, Tasmania,
New South Wales, Lord Howe Island, Queens-
land, Northern Territory, Caroline Islands, Ha-
waiian Islands, Samoa Islands, Tonga Islands,
Fiji Islands, New Caledonia, and India.

Halophila spinulosa (R. Brown) Ascherson
Figures 234,B, 24

Caulina spinulosa R. Brown, 1810:339.

33

Halophila spinulosa (R. Brown) Ascherson, 1875:368.—Mer-
rill, 1915:2; 1925:26.—Pascasio and Santos, 1930:4.—Do-
mantay, 1962:275.—Mendoza and del Rosario, 1967:
22.—den Hartog, 1970:263.

CHaARACTERISTICS.—Plants small, with long and
narrow rhizomes, not more than | mm in diam-
eter; internodes 13-35 mm long; usually one root
per node; roots 5-10 cm long, beset with filiform
hairs. Erect shoots pedicellate with 2 basal, ovate
scales with serrate apex. Sessile leaf blades up to
22 pairs, distichously arranged on erect shoot;
blades oblong-linear, semi-amplexicaulous, 20
mm long, 3.5 mm wide; margins serrate; basal
portion of one side of blade folded; a few incon-
spicuous cross-veins present and connected to the
intramarginal nerve; tip of conspicuous midrib
joining intramarginal nerve.

Natura History.—Halophila spinulosa occurs
on sand, mud, or coral rubble in the sublittoral
zone. The plant may grow together with Thalassia
hemprichu, Enhalus acoroides, Halophila ovalis, and
Halodule pinifolia. Flowering H. spinulosa has been
observed during March, October, November, and
December. Fruits were observed in October. The
plant can tolerate brackish water conditions (den
Hartog, 1970). The few specimens of H. spznulosa
examined did not reveal any epiphytes.

PHILIPPINE DistriBuTion.—Luzon (Manila
Bay, Mindoro, Camarines); Mindanao (Davao).

Rance.— Halophila spinulosa occurs in Malaysia,
Singapore, Indonesia, Western Australia, Papua
New Guinea, and Queensland.

Thalassia hemprichii (Ehrenberg) Ascherson
Ficures 25a,B, 26

Schizotheca hemprichu Ehrenberg, 1836:429.

Thalassia hemprichu (Ehrenberg) Ascherson, 1871¢:242.—
Merrill, 1912:71; 1925:27.—Pascasio and Santos, 1930:1,
figs. 1-52, plates 1-5.—Domantay, 1962:276.—Mendoza
and del Rosario, 1967:50.—den Hartog, 1970:232, fig.
61.—Cordero, 1981:323, fig. 3.

CHARACTERISTICS.—Plants of moderate size,
consisting of a robust rhizome, 2-4 mm in diam-
eter; internodes 3-6 mm long; roots clothed with
dense filiform laterals, one per node if present.
Erect shoot short, with 2-6 leaves enveloped by

SQ

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NUMBER 21

Ficure 24.—Halophila spinulosa: top, world distribution; bottom, Philippine distribu-
tion.

35

36 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

Ficure 25.— Thalassia hemprichii: a, habit of plant, X 1.0; B, upper portion of leaf showing
perpendicular nerves connected by cross veins, X 4.0.

3-8 cm-long sheaths; erect shoots sparsely distrib- | than 25 cm long), apices rounded, occasionally
uted along rhizome. Leaf blades linear and dis- uneven, margins entire; 10-16 nerves connected
tinctly falcate (particularly in less grazed areas), by cross-veins; median nerve often conspicuous.

4-10 mm wide, 7-40 cm long (commonly less Natura History.— Thalassia hemprichii is com-

NUMBER 21

Ficure 26.— Thalassia hemprichii: top, world distribution; bottom, Philippine distri-
bution.

37

38 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

mon on mud-coral-sand or coarse coral-sand sub-
strates, in sheltered habitats in the Philippines.
The plant has been observed growing from the
base and through fingers of corals at 6 m deep. T-
hemprichit may be found mixed with Syringodium
isoetifolium, Cymodocea serrulata, C. rotundata, Enhalus
acoroides, Halophila ovalis, and Halodule uninervis. At
several sites in Negros, Philippines, 7: hemprichi
appeared to be “mowed down” in large patches
and there was evidence of fish grazing on the
leaves of the plant. Algal epiphytes observed on
the leaves include species of Centroceras, Cladophora,
Enythrotrichia, Foshella, Giffordia, Herposiphonia, and
Polysiphoma. Flowering of T. hemprichi occurs dur-
ing January, February, and December in the
Philippines.

PHILIPPINE DistriBuTIoN.—Luzon (Manila
Bay, Pangasinan, Catanduanes, Albay, Batangas,
Cagayan, Mindoro, La Union, Ilocos Norte); Pa-
lawan (Quezon); Visayas (Samar, Siquijor, Cebu,
Leyte, Bohol, Negros); Mindanao (Surigao, Da-
vao, Cavili Island).

RancE.— Thalassia hemprichu is common in the
tropical region of the Indian Ocean and western
part of the Pacific. It occurs in Sudan, Eritrea,
Somalia, Kenya, Tanzania, Mozambique, Sey-
chelles, Aldabra Islands, Madagascar, Saudi Ara-
bia, Yemen, Andaman Islands, Thailand, Viet-
nam, Ryukyu Islands, Malaysia, Indonesia,
Papua New Guinea, Caroline Islands, Marshall
Islands, Bismarck Archipelago, New Caledonia,
and Queensland.

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