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ASIAN MA
BIOLO

ASIAN MARINE BIOLOGY 12

ASIAN MARINE BIOLOGY 12
1995

The Marine Biological Association
of Hong Kong

Hong Kong University Press
# m * # da % *i

© Hong Kong University Press 1996
ISBN 962 209 408 2

Printed in Hong Kong by Condor Printing Company Limited

THE MARINE BIOLOGICAL ASSOCIATION OF HONG KONG
The Association was founded on 12 February 1982 and has the following objects:
(1) To encourage and assist marine biological research and its beneficial applications in Hong Kong
...

(3) To co-operate with other scientific bodies and to seek such affiliations as may be appropriate
...


Membership of the Association is available in six classes:
Full Members, Fellow Members, Student Members, Associate Members, Honorary Members and
Sustaining Members
...
Full members shall be persons who are or have been engaged in or directed research
in a branch of marine science and who either possess appropriate academic qualifications or because
of knowledge and experience occupy positions that ordinarily would, in the opinion of Council,
require academic qualifications
...
Fellow members shall be Full Members who in the opinion of Council are
distinguished for their research in marine sciences in Hong Kong and will be entitled to use the
designation FMBAHK
...
Student members shall be persons who are studying relevant subjects at postsecondary but not postgraduate level
...
Associate members shall be persons who are interested in the objects of the
Association, but do not qualify for full membership
...
Honorary members shall be persons distinguished for their research in marine
science or who shall have rendered meritorious service to the Association
...
Sustaining members shall be persons, organizations or institutions who wish
to support the objects of the Association
...
Christine Loh, L
...
B
...
H
...
Lee, Ph
...
, I
...
O
...
P
...
B
...
D
...
B
...


Council
Chairman

: Dr S
...
Lee

Vice-Chairman

: Dr S
...
Leung

Secretary

: Dr A
...
C
...
Y
...
Cheung

Meeting's Secretary

: Dr S
...
Cheung

Membership Secretary

: Mr K
...
Leung

Librarian

: Dr LJ
...
S
...
K
...
Brian Morton
: c/o The Swire Institute of Marine Science, The University of Hong Kong,
Cape d'Aguilar, Shek O, Hong Kong

ASIAN MARINE BIOLOGY
Editor
Prof
...
Y
...
Cheung, Department of Biology and Chemistry, City University of Hong Kong, Tat Chee
Avenue, Kowloon, Hong Kong
Editorial Board
Prof
...
Birkeland, University of Guam Marine Laboratory, UOG Station, Mangilao, Guam 96913, USA
Dr Y
...
Chien, Department of Aquaculture, National Taiwan College of Marine Science and Technology,
Keelung, Taiwan
Dr L
...
Eldredge, Pacific Science Association, PO Box 17801, Honolulu, HI 96817, USA
Dr Y
...
Go, Department of Oceanography, College of Ocean Science and Technology, Cheju National
University, Cheju 590, South Korea
Prof
...
D
...
D
...
P
...
K
...
Ng, Department of Zoology, National University of Singapore, Kent Ridge, Singapore 0511
Prof
...
Okutani, Tokyo University of Fisheries, 4-5-7, Konan, Minato-ku, Tokyo 108, Japan
...
J
...
Ong, School of Biological Sciences, Universiti Sains Malaysia, 11800 Penang, Malaysia
Prof
...
K
...
E
...
Veron, Australian Institute of Marine Science, P
...
B
...
3, TownsviUe MC, Queensland
4810, Australia
Dr Yu
...
Yakovlev, Institute of Marine Biology, Far East Branch, Russian Academy of Sciences,
Vladivostok 690032, Russia

INSTRUCTIONS TO AUTHORS
Submission of manuscripts
...
The submission of a manuscript will be taken to imply that the material is original and that
no similar paper is being, or will be, submitted for publication elsewhere
...

Theses and serialized studies should not be submitted
...
Papers should be written in clear, concise English with the minimum number of tables
and illustrations
...
A charge will be made for changes introduced after an article has been
typeset
...
This should be concise, specific and typed on a separate cover sheet together with the
name(s) of the author(s) and that of the institution in which the research was carried out
...

A short title, of not more than 40 characters (including spaces), should be provided for page
headings
...
This should generally not exceed 200 words and must be intelligible to the reader
without reference to the main text
...

(c) Introduction, followed by the subject matter of the paper
...

(e) Results
...

(g) Acknowledgements
...

(i) Illustrations—See Section 4
...

Presentation
...
5 cm all round
...
All pages should be numbered serially and
fastened together securely
...

The first subhead will be shouldered, the second italicized shouldered and the third run-on with
the copy
...
Generic and specific names must be italicized
...

If it is necessary to refer to various passages in the text, please ensure that the relevant
manuscript page number is given
...

Customary abbreviations will be used
...
When a paper has more than two authors, the style Smith et al
...
1980) should be used
...
Note that these
papers should be alphabetized by title, discounting an initial article
...
All illustrations will be reduced to a size not exceeding 19x14 cm
...

A metric scale should preferably be included on each illustration
...

Line drawings are referred to as Fig
...
2, etc
...

If a line drawing or halftone plate is composed of more than one subject, one should be identified
as (A), (B), etc
...

(a) Line drawings
...
5 cm) or A4 (30 x 21 cm) sheets
...
Photocopies of drawings cannot be reproduced satisfactorily, but may in the first instance
be submitted, and the original drawing sent if the paper is accepted
...

Graph curves may be distinguished by use of solid
pecked
or dotted
...
The
following symbols available to the printer also, can be used for experimentally determined points
and for keying in the legends: O, • , • , • , A, A, V, T
...
Best quality glossy prints should be submitted of a size
capable of reduction rather than enlargement
...

Numerical matter
...
Only the minimum of numerical matter
essential for clarity should be included
...

Important: Authors are requested to check the list of references against the text to ensure that:
(a) The spelling of authors' names and the dates given are consistent
...
If more than one reference
by the same author(s), published in the same year is cited, use, a, b, etc
...
g
...
Text citations can be given in either of two ways:
(i) Single author: Where more than one reference is given for a single author the publications
should be listed chronologically
...
For text citations, use both authors' names and the year
...
for two author references
...
For all citations in the text it is usually sufficient to
quote the surname of the first author, followed by et al
...

IX

Instructions to authors

Please note that the names of all authors should be given in the list of references and et al
...
The full title of the paper must be given
...
The volume number should be followed by the first and last pages of the paper
...


Examples
Journal references:
Beklemishev, K
...
1952
...
Trudy Vsesoyusnogo Hidrobiologicheskogo Obschestva 5:276-96
...
F
...
H
...
and Colman, M
...
1981
...
Journal of the
Malacological Society of Australia 3:41-64
...
Y
...
Size, maturation and the social control of sex reversal in the coral reef fish Anthias
squamipinnis
...

Tyler, P
...
and Gage, J
...
1982
...
Journal of the Marine Biological Association of the United
Kingdom 62:45-56
...
G
...
X
...
Marine Biofouling and Its Prevention
...

(In Chinese)
Johnson, R
...
and Ohlhorst, C
...
1981
...
In Ocean Dumping of Industrial Wastes (ed
...
H
...
R
...
Kilho Park)
...
12, Marine science, 175-91
...

(Note: 'ed
...
')
King, C
...
M
...
Beaches and Coasts
...

Lam, C
...
Y
...
The Assessment of Eutrophication in Estuaries: A Review of New Zealand and
Overseas Studies
...
79/38
...
New Zealand: Ministry
of Works and Development
...
and Tseng C
...
, eds
...
The Marine Flora and Fauna of Hong Kong and Southern China
...
Proceedings of the First International Marine Biological Workshop: The Marine Flora and
Fauna of Hong Kong and Southern China, Hong Kong 1980
...
(Note: 'eds
...
Y
...
T
...
A preliminary checklist of the marine Gastropoda and Bivalvia (Mollusca)
of Hong Kong and southern China
...
B
...
K
...
Proceedings of the First International Marine
Biological Workshop: The Marine Flora and Fauna of Hong Kong and Southern China, Hong Kong
1980
...

Unpublished material:
Cortes-Zaragoza, E
...
Morphometries and relative abundance of tunas (Perciformes: Scombridae)
caught off Darigayos Cove, La Union
...
M
...
thesis, University of the Philippines
...
K
...
1986
...

Paper presented in the International Symposium on Environmental Pollution and Toxicology, 9-11
September 1986, Baptist College, Hong Kong
...


Proofs
...
Corrections should be made on the proof
which should then be returned within 48 hours to: The Editor, Asian Marine Biology, The Swire
Institute of Marine Science, The University of Hong Kong, Cape d'Aguilar, Shek O, Hong Kong
...


Copyright/Offprints
...
Offprints may be ordered at the proof stage
...


9
...
Whenever a Latin name is employed it must be followed by the authority, e
...
,
Anomalocardia squamosa Lamarck, on first usage in the paper
...

(b) All primary types of new taxa must be quoted in the paper
...


10
...
All manuscripts must be provided on a floppy disk after the paper has been
accepted for publication, in addition to the final printed version
...
44M 3
...
25" disk for IBM personal
computer, or compatible, or a 1
...
5" disk for Apple Macintosh computer
...

(a) Ensure that the disk you send contains only the final version of the paper and is identical to
the typescript
...

(c) Supply the file as a 'text only' WordPerfect (5
...
0 file on IBM personal computer or a Microsoft Word (5
...

(d) Use no special formatting characters
...


XI

EDITORIAL
This year, 1995, is again significant for the development of marine science in Hong Kong and it reflects
the growing interest in the marine environment locally
...
This was finally passed into law by the
Council on 31 May 1995 and the Marine Parks Ordinance thus became effective on 1 June 1995
...
On 10
August 1995, the Country Parks Board was officially retitled the Country and Marine Parks Board with
new responsibilities for the proposed marine parks and reserves
...
Established in 1989, with representation on it by the Marine
Biological Association of Hong Kong, the working group presented its first report to the Government
in 1990
...
Subsequently,
extensive consultation took place between the Government and interested parties and the final
recommendation to gazette the three areas was made on 1 August 1995, at the last meeting of the working
group
...
The only action
needed now is designation of the proposed parks and reserves by the Government
...
Two books about the marine parks and reserves, Hoi Ha Wan and An Introduction to the
Cape d'Aguilar Marine Reserve, Hong Kong, can be obtained from the World Wide Fund for Nature
Hong Kong and The Swire Institute of Marine Science, respectively
...

In 1994, The Hong Kong Government sponsored two Ph
...
students based at the Swire Institute of
Marine Science (SWIMS), to study the Chinese White dolphin, elsewhere referred to as the Indo-Pacific
Humpback dolphin (Sousa chinensis), in view of the emerging concern regarding the large number of
dead animals (nine in 1994) recorded as strandings in northwestern waters
...
Other developments are also taking place in these waters, however, and
which, it is now known, are the almost exclusive habitat of the dolphin
...
It has become a matter of the greatest concern that Hong Kong — ranking as eighth in
the world league of trading economies, with the sixth highest per capita GDP in the world in 1993 and
a budget surplus in 1994 of HK$19
...
The dolphin particularly
prefers the shallow area of water around a group of three islands, Tree Island, Sha Chau and Lung Kwu
Chau, in the furthest northwestern waters of Hong Kong and this was proposed by the Swire Institute
of Marine Science researchers as a possible dolphin sanctuary
...
This temporary facility will require a dredged turning circle and
channel for the vessels
...
Notwithstanding, in an attempt to placate conservationists who
argued that such a facility was both unnecessary if plans had been made in time to build a permanent
facility direct to the airport and that its construction would spell the end of the dolphins, the Government
has proposed that a 1,200 hectare area of sea bed around the islands be designated as a marine park
...
If no developments were planned for the area, there might be a
chance that the dolphins could be saved
...
The only possiblity for their survival would be the establishment
of a much bigger park to the south, west and north of northwest Lantau, ideally extending into Chinese
waters also
...
It is a remarkable fact that, despite being
afforded all this protection, the dolphin will probably become locally extinct and that Hong Kong and
China cannot, together, protect it
...
Laudable though, therefore, that so much has been achieved, it is now
increasingly doubtful that any of the places proposed for designation as marine parks and reserves will
actually survive
...
Any Hong
Kong proposals, therefore, to protect sensitive marine environments from internal pressures will meet
external ones which will grow in ignorance of what the Hong Kong Government is attempting to protect
and why
...
To help
achieve this, the Third International Conference on the Marine Biology of the South China Sea is to be
convened in Hong Kong by the Marine Biological Association of Hong Kong and the Swire Institute
of Marine Science from 28 October to 1 November 1995 (inclusive), at the University of Hong Kong
...
Interested potential participants should contact the
Conference Secretary at the Swire Institute of Marine Science
...
F
...
F
...
G
...
K
...
S
...
Retiring from the council, Dr S
...
Chiu (Vice-chairman)
and Dr R
...
Ong Che and Dr K
...
Chu (Council members) are herein thanked for their contributions
to the work of the Association
...
Williams and her position has been filled by Dr J
...
N
...


Brian Morton

xv

CONTENTS
The Marine Biological Association of Hong Kong
Asian Marine Biology — Editorial Board

v
vii

Instructions to Authors

viii

Editorial

xiii

Competitive interactions between co-occurring limpets, Cellana toreuma (Reeve, 1855) and
Patelloida saccharina (Linnaeus, 1758), on a rocky intertidal shore at Cape d'Aguilar, Hong Kong
J
...
Liu

1

Field experiments of predation by the muricid gastropods Thais clavigera and Morula musiva
on the intertidal barnacle Tetraclita squamosa at Cape d'Aguilar, Hong Kong
J
...
Liu

11

Spatial variation in larval trematode infections of populations of Nodilittorina trochoides and
Nodilittorina radiata (Gastropoda: Littorinidae) from Hong Kong
Tang Chong-ti

19

Community structure of scleractinian reefs in the Baitylong Archipelago (South China Sea)
Yuri Ya
...
A
...
Seed, N
...
Brotohadikusumo and R
...
Ong Che and Brian Morton

53

Occurrence of the barnacle Balanus amphitnte Darwin on the shells of the predatory gastropod
Thais clavigera Kuster at Cape d'Aguilar, Hong Kong
J
...
Liu and L
...
C
...
Parsons, Mary L
...
Porter

79

Odysseylana sirenkoi: A new genus and new species of cirolanid isopod from the South China
Sea (Crustacea
...
Malyutina

101

Records of Cuvier's beaked whales (Ziphius cavirostris) from Taiwan, Republic of China
J
...
Wang, L
...
Chou, C
...
Yao, A
...
Neimanis and W
...
Chou

Ill

Contents

Records of small cetaceans in Chinese waters: a review
Zhou Kaiya, Stephen Leatherwood and Thomas A
...
H
...
The two species were caged at densities ranging from natural to
4 times this number on the shore and in different combinations of species, to elucidate possible
competitive interactions
...
toreuma showed increased mortality and reduced weight at increased
densities over a two-month period, due to intraspecific competition
...
saccharina showed a slightly
reduced weight but no increased mortality at increased densities
...
toreuma
was thus greater than that on P
...
Tissue weights of P
...
toreuma, but not mortality
...
toreuma by P
...
Food supply, i
...
, encrusting algae and microalgae, is thought to be the limiting
factor for which the species compete
...
Unlike sessile
organisms, grazing limpets are not expected to
compete for space but, at high densities, may
compete for food (Haven 1973; Underwood 1979;
Creese and Underwood 1982; Hawkins and
Hartnoll 1983)
...
Apart from this, little research has
been done on the ecology of Hong Kong limpets
...
P
...

toreuma is a non-homing limpet and migrates to
upper shore levels in winter (Liu 1993b)
...

It is hypothesized that: (1) increased mortality and
reduced weight may occur at in-creased densities
of C
...
saccharina due to
intraspecific competition for food, and (2)
interspecific competition for food between these
two species may be occurring due to closely
similar methods of feeding
...


J
...
Liu

Plate 1
...
Cellana toreuma and B-B1, Patelloida saccharina
...
Hong Kong Island
(Fig
...

The study area delineated was between 1 m and
2 m above chart datum
...
i
...
Cellana toreuma
(5
...
0
...
n = 20) and Patelloida
saccharina (6
...
0
...
n = 20)
...

?

Cages used in the experiments were made of
metal netting of 0
...
5 cm mesh size and were
of 23 x 23 cm
...
e
...
The cages were fastened
to the rock by metal screws and rawl-plugs
...
Cellana toreuma and
Patelloida
saccharina were enclosed in cages in various
combinations and different densities, i
...
from

Competitive interactions between co-occurring limpets

Fig
...
A map of the southeastern coastal waters of Hong Kong showing the study site
...
Treatments were assigned randomly to the
cages
...
Adult
limpets were used in the experiments, as such
individuals experience little growth (Liu 1994a)
...
toreuma and
between 14-16 mm for P
...
Original
individuals used in the experiments were tagged
using glue-on polyethylene mollusc tags, with a
protective coating of the instant glue Aron Alpha
...

Dead animals were replaced by similar-sized
conspecifics to maintain densities
...
At the beginning
and end of the experiment, a random sample of
uncaged adult individuals of each species was
collected and shell lengths and tissue weights were
obtained to compare with caged individuals in
control (natural) densities
...
H
...
Cellana toreuma (C) and Patelloida saccharina (P)
...

Type of interaction
Total number of limpets

5

10

20

Experimental treatments

5C
5P

10C
10P
5C + 5P

20C
20P
5C + 15P
5P + 15C

Mean shell length (mm)

C
...
saccharina

Range

27-33

14-16

the cages and shell lengths and tissue weights
were obtained
...
A
predatory snail, Morula musiva (Kiener)
occasionally entered the cages and was removed
when found
The available food supply for foraging
limpets, in terms of biomass and species
composition of algae colonizing the rock surface,
were investigated
...
Algal biomass
was determined by chlorophyll a analysis
...
Samples
were analysed within 24 hours of collection,
usually after overnight storage in an air-tight and
light-sealed container placed in a refrigerator
(4°C)
...
The absorbance of the
extract was measured at wavelengths of 665 nm
and 750 nm using a PYE UNICAM PU 8600 UV/
VIS spectro-photometer
...
Calculations followed the formula:
The chlorophyll a content of the sample
13
...
cm- 2 (HMSO 1986)

Intraspecific
Intraspecific
Interspecific
Interspecific

Where A = absorbance
v = volume of solvent in ml
V = area of rock sampled in cm2
d = cell path length in cm (4
...
9 = constant
It is hypothesized that mortality of the same
species occurred equally in all cages
...
The question
to be asked, then, is whether the observed
frequencies deviate significantly from the
expected frequencies if the hypothesis is true? A
chi-square test was used as an overall measure of
deviation to determine whether actual mortalities
in the cages differed significantly
...
2)
...
toreuma at increased
densities
...
toreuma
reached 30% at high densities (20 individuals
...
05 m2) but was 10% at lower densities (5 and
10 individuals
...
05 m2)
...
saccharina showed no
effect on C
...
Thus,
in the treatments of 5 C
...

saccharina, no mortality of C
...
2A)
...

toreuma from 12 cages (6 treatments) were
recorded and the observed frequencies (O) are
given in Table 2
...
25) (Table 2)
...
At the highest
density of P
...
0
...
2B)
...
saccharina by Cellana
toreuma at increased densities
...
toreuma, i
...
, 5 P
...
toreuma, no mortality of P
...
2B)
...
Again, the null
hypothesis (of the same mortality rate in all cages)
cannot, therefore, be rejected (P < 0
...


AQn

Cellana toreuma

X

o
O

E

20-

c

D
0)

—i—

5C

1

10C

20C

5C+5P

5C+15P

•

r

15C+5P

B
Patelloida saccharina

401

X

o
o

20-

E

c
o
a>
2

0-

T

5P

10P

20P

'1

5P+5C

Fig
...
A, Cellana toreuma (C) and B, Patelloida saccharina
control and experimental densities
...
Mean mortalities of individuals at

5

J
...
Liu

Table 2
...

Cages

Observed
frequency
0
1
0
1
1
7
5
0
0
0
0
3
5

5C-1
5C-2
10C-1
10C-2
20C-1
20C-2
5C+5P-1
5C+5P-2
5C+15P-1
5C+15P-2
15C+5P-1
15C+5P-2

n = 23

Probabi lity
(if Ho true)

Expected
frequency

7t

_
=
=
=
=
=
=
=
=
=
=
=

Deviation
(O-E)
E = nrc

0
...
042
0
...
083
0
...
167
0
...
042
0
...
042
0
...
125

1
...
966
0
...
909
1
...
841
3
...
966
0
...
966
0
...
875
2
...
966
0-0
...
909
1-1
...
841
5-3
...
966
0-0
...
966
0-0
...
875
5-2
...
0

5/120
5/120
10/120
10/120
20/120
20/120
5/120
5/120
5/120
5/120
15/120
15/120

0

Deviation squared
and weighted
(O - E)2/E
0
...
966
0
...
433
2
...
350
0
...
966
0
...
966
0
...
571
X2 = 10
...
25

Table 3
...

Cages

Observed
frequency
O

5P-1
5P-2
10P-1
10P-2
20P-1
20P-2
5P+5C-1
5P+5C-2
5P+15C-1
5P+15C-2
15P+5C-1
15P+5C-2

0
0
0
0
2
0
1
0
0
0
2
1
n=6

Probability
(if H o true)
=

=
=
=
=
=
=
=
=
=
=
=

0
...
042
0
...
083
0
...
167
0
...
042
0
...
042
0
...
125

1
...
73 for 20 Cellana, 2
...
252
0
...
498
0
...
002
1
...
252
0
...
252
0
...
750
0
...
252
0-0
...
498
0-0
...
002
0-1
...
252
0-0
...
252
0-0
...
750
1-0
...
252
0
...
498
0
...
994
1
...
220
0
...
252
0
...
083
0
...
638
p-value < 0
...
01 for 10
Cellana) (Fig
...
There was little effect on C
...
98)
...


Competitive interactions between co-occurring limpets

saccharina were not decreased at higher densities
(2
...
There was, however, a
strong effect on P
...
toreuma (5
Patelloida + 15 Cellana), with ratios being
smaller (0
...
3B)
...
!?
'©

b

10-

1

'—i—'

C1

C2

—i—'

20C

10C

i—
...
3
...
Mean dry weight: shell length
ratios (g-mnrr1) at various experimental densities (Mean ± S
...
C1, Uncaged control on 25
March and C2, Uncaged control on 21 May
...
H
...
For both Cellana toreuma
and Patelloida saccharina, the mean dry tissue
weight/shell length (x 103) ratios of the uncaged
control animals at the beginning of the experiment,
i
...
, 6
...
00, respectively, were significantly
greater than the mean ratios obtained for the
uncaged, i
...
, 4
...
863, respectively, and
caged control animals (5 Cellana, 5 Patelloida),
i
...
, 4
...
31, respectively, at the end of the
experiment
...
e
...
66 vs 4
...
toreuma
and 1
...
31 for P
...

The significant differences in the dry tissue
weight/shell length ratios of uncaged individuals
at the beginning and the end of the experiment are
probably attributable to seasonal changes in algal
production (Liu 1993a; 1994a)
...
,
blue green algae and diatoms
...
Table 4 shows
calculated values of chlorophyll a (ug-cnr2) at the
beginning and the end of the experiment
...
10)

than at the end (2
...
Such a change is thought
to be caused by variations in rates of algal
production, which are related to seasonal changes
in the physical environment, particularly with
regard to temperature and the degree of wetting
(Liu 1994a)
Algal biomass decreased abruptly in cages
under heavy grazing pressure (20 Cellana) after
a two-month period, i
...
, from 3
...
64
...


Discussion
This experiment shows that Cellana toreuma
suffered increased mortality and a loss in tissueweight due to increases in density above the
natural level
...

Liu (1993b) showed that P
...
Homing behaviour may reduce
such competition (Mackay and Underwood 1977)
...
toreuma, therefore, shows stronger intraspecific
competition
...

Interspecific interactions between Cellana
toreuma and Patelloida saccharina were also
examined
...
toreuma

Table 4
...
Mean ± SD
...

Treatment

Uncaged control

5C
10C
20C
5P
10P
20P
5C+5P
5C+15P
5P+15C
8

Chlorophyll a (ug-cnr2)
At the end
At the beginning
3
...
96
3
...
14
2
...
04
2
...
73
2
...
20

(0
...
77)
(0
...
32)
(0
...
64)
(0
...
81)
(0
...
02)

2
...
21
1
...
64
2
...
15
1
...
46
1
...
71

(0
...
27)
(0
...
17)
(1
...
37)
(0
...
36)
(0
...
28)

Competitive interactions between co-occurring limpets

affected individuals of P saccharina in terms of
a reduction in tissue-weight but little in terms of
mortality The former thus appears to be
competitively superior to the latter The absence
of C toreuma resulted in increases in the tissue
weights of P saccharina (Fig 3A, 20 Patelloida,
10 Patelloida and 5 Patelloida) Increasing the
density of P saccharina did not, however, affect
individuals of C toreuma Haven (1973) tested
interspecific competition in a habitat where both
Acmaea scabra and A digitalis were common but
where the former was smaller than the latter
Removal of A digitalis from fenced exclosure
plots resulted in a significant growth increase in
A scabra, correlated with pronounced increases in
algal food supply Field observations (personal
observation) also suggest that the absence of C
toreuma (or only a few present) results in an
increase in the density of P saccharina (30
individuals 500 cm2) The presence of C toreuma
may effect a reduction in the abundance of P
saccharina
On local shores, however, the
densities of C toreuma do not occur at high
enough levels to eliminate P saccharina because
of strong interspecific competition for food (Liu,
1994a) and predation (Liu unpublished data)
which reduce the numbers of C toreuma C
toreuma affects its own survivorship more than
that of P saccharina It appears that the major
factor
allowing coexistence between P
saccharina and the superior competitor, C
toreuma, is the effect of intraspecific competition
among the C toreuma which reduces density
below levels at which they can eliminate P
saccharina
'Asymmetrical competition', where one
species adversely affects another but the reciprocal
effect is less intense, has been documented for a
number of marine organisms (Connell 1961,
Dayton 1971, Menge 1972, Underwood 1984) It
has also been demonstrated for co-occurring
limpets using experimental manipulations (Haven
1973, Stimson and Black 1975, Underwood 1984,
Black 1979, Creese 1982, Creese and Underwood
1982, Fletcher and Creese 1985, Garnty and
Levings 1985) This study has shown that
Patelloida saccharina has a relatively small effect
on Cellana toreuma, while C toreuma has a great
effect on P saccharina and on individuals of its

own species
An earlier study by Liu (1992) showed that
both Cellana toreuma and Patelloida saccharina
have a docoglossan radula which is capable of
scraping into the rock surface, I e , they are able
to consume the algal crusts of Hildenbrandia
prototypus and Gomontia sp What are the
mechanisms by which C toreuma, in experimental
cages, outcompetes P saccharina1 There is no
major difference between the radulae of C
toreuma and P saccharina, 1 e , their abilities to
use algal crusts are probably equal and from this
point of view neither is a superior competitor The
results can only be explained by the fact that while
the smaller P saccharina has no effect on C
toreuma, the larger C toreuma disturbs the much
smaller P saccharina by interfering with foraging
P saccharina is also active only at night when
submerged and homes to a scar (Liu 1993b)
Liu (1992) showed that Cellana toreuma and
Patelloida saccharina share a common food
resource which is in limited supply Grazing by
both C toreuma and P saccharina at natural
densities has a great impact on the colonization
and growth of algae Removal of all herbivores
resulted in the establishment of opportunistic
species of algae, e g , Acrochaetium robustum,
Bangia fuscopurpurea, Porphyra dentata and
Ulva lactuca Shortage of food is the most likely
reason for both the intra- and interspecific
competition demonstrated in this study, leading to
reduced tissue-weight and mortality It is
concluded that, in Hong Kong, limpet grazing is
strongly limited by algal availability Inter- and
intraspecific competition for food increases the
mortality and tissue weight loss of C toreuma and
P saccharina

Acknowledgements
I am grateful to The University of Hong Kong for
awarding me a Rayson Huang Postdoctoral
Fellowship I thank Professor Brian Morton,
Director, The Swire Institute of Marine Science,
The University of Hong Kong, for the provision
of laboratory facilities and work space during the
course of this study and for his critical reading of
the first drafts of the manuscript of this paper
9

J
...
Liu

References
Black, R
...
Competition between intertidal
limpets: an intrusive niche on a steep resource
gradient
...
H
...
The influence of interspecific
competition and other factors on the
distribution of the barnacle
Chthamalus
stellatus
...

Connell, J
...
1983
...
American
Naturalist 122: 6 6 1 - 9 1
...
G
...
Distribution and abundance of
the acmaied limpet Patelloida latistrigata and
its interactions with barnacles
...

Creese, R
...
and Underwood, A
...
1982
...
Oecologia 53: 337-46
...
K
...
Competition disturbance and
community organisation: the provision and
subsequent utilization of space in a rocky
intertidal community
...

Fletcher, W
...
and Creese, R
...
1985
...
Marine Biology 86: 183-91
...
D
...
C
...

Interspecific interactions and scarcity of a
tropical limpet
...

Haven, S
...
1973 Competition for food between
the intertidal gastropods Acmaea scabra and
Acmaea digitalis
...

Hawkins, S
...
and Hartnoll, R
...
1983
...

Oceanography
and Marine Biology
Annual
Review 2\: 195-282
...
S
...
J
...
An
investigation of methods for sampling
microbial films on rocky shores
...

HMSO 1986
...
4
...
Section 2
...

10

Liu, J
...
1992
...
Ph
...

Liu, J
...
1993a
...
Asian Marine Biology 10: 2 7 - 4 0
...
H
...
Activity rhythms
and
'homing'abilities by two pairs of high and
low-zoned intertidal limpets in Hong Kong
...
The Marine Biology of the South China
Sea
...
B
...
Hong Kong:
Hong Kong University Press
...
H
...
The ecology of the Hong Kong
limpets Cellana
grata and
Patelloida
pygmaea:
distribution
and
population
dynamics
...

Liu, J
...
1994b
...
Journal of
Molluscan Studies 60: 9 7 - 1 1 1
...
A
...
J
...

Experimental studies on homing in the
intertidal patellid limpet Cellana
tramoserica
(Sowerby)
...

Menge, B
...
1972
...

Ecological
Monographs 46: 3 5 5 - 9 3
...
W
...
Field experiments on
interspecific
competition
...

Stimson, J
...
1975
...
Oecologia 18: 111-20
...
J
...
An experimental
evaluation of competition
...

Underwood, A
...
1979
...
Advances in Marine Biology 16:
111-210
...
J
...
Vertical and seasonal
patterns in competition for microalgae
between intertidal gastropods
...


Asian Marine Biology 12 (1995): 11-17

FIELD EXPERIMENTS OF PREDATION BY THE MURICID GASTROPODS
THAIS CLAVIGERA AND MORULA MUSIVA ON THE INTERTIDAL
BARNACLE TETRACLITA SQUAMOSA AT CAPE D'AGUILAR,
HONG KONG
J
...
Liu
The Swire Institute of Marine Science, The University of Hong Kong, Cape d'Aguilar, Shek O, Hong
Kong

Abstract
Field experiments were undertaken to determine the importance of predation by the muricid gastropods
Thais clavigera and Morula musiva on a population of the barnacle Tetraclita squamosa inhabiting a
sub-tropical rocky intertidal community in Hong Kong
...
clavigera and M
...
Closed cages containing prey
without predators were used to estimate non-predator mortality
...

Tetraclita squamosa declined in density in open plots throughout the experimental period
...
squamosa in closed cages showed almost no reduction in density between September 1993 and
June 1994 but declined in numbers between June and August 1994
...
28%
...
Temperature-induced mortality occurred between June and August in all control and
experimental plots and when the mean mortality rates in closed cages and open plots were 25
...
60%, respectively
...

This indicates that predation intensities in open plots were low during the summer months
...
55%)
...
78%
...
Predation and summer heat are
the major factors affecting the barnacle population and the former is thought to be more significant
than the latter
...
The effects of intertidal muricid
gastropods upon their intertidal prey populations,
largely mussels and barnacles, have been well
studied (Connell 1961, 1970; Menge 1976; Moran
1985a; Fairweather 1985; Abe 1989; Fairweather
and Underwood 1991; Morton and Britton 1993)
...

Higher zoned, intertidal, species, e
...
, Thais
clavigera Kuster and Morula musiva (Kiener) feed
mostly on oysters and barnacles (Morton and
Morton 1983; Taylor 1990)
...
clavigera occurs
on almost every intertidal shore while M
...

T
...
musiva differ in size with
maximum shell heights of 50 mm and 36 mm,
respectively (Taylor 1990)
...
H
...
Abe (1989) showed that small T
...

clavigera mainly on the large barnacle Tetraclita
squamosa japonica
...
musiva is the main
predator of the mussel Septifer virgatus
...
clavigera
showed a marked seasonal pattern of migration
...

Both Thais clavigera and Morula musiva are
known to feed on T
...
Although Tong (1986b) has
shown, using gut contents analysis, that T
...
musiva are important predators
of the intertidal barnacle T
...
squamosa as a result of muricid predation
...
Since high temperatures in mid
summer have also been causally linked to
mortality of this species (Liu and Morton 1994),
it is hypothesized that:
(1) there would be significant differences in the
mortality of prey with and without predators;
(2) higher mortality rates may occur in open plots
due to predation by muricid gastropods; and
(3) that in closed cages, mortality may occur
during the summer months because of heat
stress
...


Material and methods
Field experiments were undertaken on the
northern shore of Lobster Bay at Cape
d'Aguilar, on Hong Kong Island
...
Uncrowded barnacle areas (< 120
individuals-m2) were chosen to reduce the chance
of competition
...
5 cm mesh size and were
23 x 23 cm in diameter and 4 cm high
...
Gaps between the substratum and the
base of a cage were loosely sealed (to allow for
drainage) with a non-toxic DEVCON wet surface
repair putty
...
Open plots as
controls were also established using rawlplugs at
their corners to allow all predators to gain access
to the prey (predator plots)
...
Individuals of Tetraclita squamosa used
in the experiments were tagged using glue-on
polyethylene mollusc tags, with a protective
coating of the instant glue Aron Alpha
...
The numbers of
dead and living individuals of Tetraclita squamosa
in each experimental plot were recorded every
month
...
Numbers of each species
were recorded in the field
...
Experimental treatments
...


160

J2 1 2 0

Results
Figure 1 shows the mean daily maximum and
mean air temperatures between September 1993
and August 1994 (Royal Observatory, Hong Kong
1993,1994)
...
2°C to 30
...
0°C to 28
...
Peak temperatures were recorded
between June and August, i
...
, 30
...
0°C
for mean daily maximum and mean temperatures,
respectively
...
2
...
• Thais clavigera; V
Morula musiva

35 -,

25

30

20

25

15
20

15
5 10
S

1993

0

N

D

J

F

J

J

1994

Fig
...
Mean daily maximum and mean air
temperatures (°C) between October 1993
and August 1994 (Records from Monthly
Weather Summary, between October
1993
and
August
1994,
Royal
Observatory, Hong Kong)
...
2)
...
3) when the mean rate
was 4
...
50) (Table 2)
...
e
...
There was a decline in

s o
1993

N

D

J

F

1994

Fig
...
Mortality rates of Tetraclita squamosa in
closed cages throughout the experimental
period
...


numbers of T
...
This
was probably because some of the cages were
broken by wave action at this time
...
18% (±
8
...
This decline is thought to be associated
with the summer heat
...
40% (± 10
...

Figure 4 shows the mortalities of Tetraclita
13

J
...
Liu

Discussion

1993

1994

Fig
...
Mortality rates of Tetraclita squamosa in
open plots throughout the experimental
period
...


squamosa in open plots throughout the
experimental period
...
squamosa, respectively, throughout the
experimental period
...
60% (+ 6
...
All
individuals in the open plot with 9 T
...
Mortalities largely occurred between
September 1993 and June 1994 in all the plots and
the mean mortality rate during that period was
87
...
76)
...
78% (± 9
...


Mortality of Tetraclita squamosa due to predation
by Thais clavigera and Morula musiva is
confirmed by this study
...
squamosa in all closed cages were
slightly lower than those in open plots
...

Burrows and Hughes (1989) showed that the
natural cycle of foraging and sheltering in the
dogwhelk Nucella lapillus (Linnaeus) is closely
associated with changing weather conditions
...
Moran (1985b) showed that
Morula marginalba Blainville forms aggregations
in a pool or crevice when environmental
conditions are unsuitable for feeding
...
Foraging activities of the predators
were likely influenced by many physical factors,
e
...
, temperature and wave action
...
clavigera
and M
...
Liu
and Morton (1994) showed that T
...
Although the mean daily

Table 2
...

Experimental period

Mean % mortality rate (+ S
...
)
Closed cage

September 1993 - June 1994
June - August 1994
September 1993 - August 1994

14

4
...
50)
25
...
40)
28
...
84)

Open plot
87
...
76)
35
...
22)
91
...
66)

Predation by Thais clavigera and Morula musiva on Tetraclita squamosa

maximum air temperature recorded by the Royal
Observatory, Hong Kong, shows a peak of 30
...
2°C in the early afternoon of a hot and
sunny day at Cape d'Aguilar (Liu and Morton
1994)
...
squamosa
...
Such influences, as shown in this study, in
variations in the effects of the predators on their
prey population, i
...
, low predation intensity
correlated with low predator densities and harsh
environmental conditions in the summer months
...
This can be interpreted as
being due to the exclusion of the predators and
confirms that the high mortality rates in all open
plots were caused by predation
...
Hart and Palmer (1987) showed that large
individuals of Thais emarginata (>15 mm shell
length) exhibit little variation in the locations at
which they drill barnacles, and attack the
opercular plates more frequently and concentrate
their attacks at the suture between the scutal
plates
...
g
...
Although predation has a major
influence on community structure, predators are
not completely efficient and, occasionally,
individuals escape the attentions of the predators
...

Both Thais clavigera and Morula musiva were
largely found to form aggregations in rock
crevices during the low tide period but otherwise
ranged widely over the intertidal area
...
Both species were

abundant on the shore throughout the year with
slightly decreased numbers during late autumn and
early winter
...
Tong (1986a) has shown that
recruitment of T
...
musiva is recruited between
September and March
...

Mortality can be affected by both biological
(e
...
, predation and competition) and physical
(e
...
, summer heat and strong wave action)
factors
...
18% which accords with figures
obtained by Liu and Morton (1994)
...
e
...
06% for T
...
This study,
however, suggests that the structure of the
barnacle population is affected more strongly by
biological factors (e
...
, predation) than by
physical ones (e
...
, summer heat) since major
declines in abundance can only be explained by
mortality from predation
...
g
...
g
...
Other sources of mortality,
e
...
, competition, were not studied in this work
but since the barnacles in the cages were not
crowded, this seems unlikely
...
I thank Professor Brian Morton,
Director, the Swire Institute of Marine Science,
the University of Hong Kong, for the provision of
laboratory facilities and work space during the
course of this study and for his critical reading of
the first drafts of the manuscript of this paper
...
H
...
1989
...
Physiological Ecology
(Japan) 26: 1-38
...
T
...
N
...
Natural
foraging of the dogwhelk Nucella lapillus
(Linnaeus): the weather or whether to feed
...

Connell, J
...
1961
...
Ecological Monographs
31: 61-104
...
H
...
A predator-prey system in the
marine intertidal region
...
Balanus glandula
and several predatory species of Thais
...

Fairweather, P
...
1985
...

Journal of Experimental Marine Biology and
Ecology 89: 135-56
...
G
...
J
...

Experimental removals of a rocky intertidal
predator: variations within two habitats in the
effects on prey
...

Fairweather, P
...
, Underwood, A
...
and Moran,
M
...
1984
...

Marine Ecology - Progress Series 17: 14356
...
W
...
R
...
Stereotypy,
ontogeny and heritability of drill site selection
in
thaidid
gastropods
...

Liu, J
...
and Morton, B
...
The temperature
tolerances of Tetraclita squamosa (Crustacea:
Cirripedia) and Septifer virgatus (Bivalvia:
Mytilidae) on a sub-tropical rocky shore in
Hong Kong
...


Menge, B
...
1976
...
Ecological Monographs 46:
16

355-93
...
J
...
Effects of prey density , prey
size and predator size on rates of feeding by
an intertidal predatory gastropod Morula
marginalba Blainville (Muricidae), on several
species of prey
...

Moran, M
...
1985b
...
Journal of
Experimental Marine Biology and Ecology
93: 103-14
...
and Britton, J
...
1993
...
In
Proceedings of the Fifth International Marine
Biological Workshop: The Marine Flora and
Fauna of Rottnest Island, Western Australia
(eds
...
E
...
I
...
Kirkman and
R
...
Perth: Western
Australian Museum,
Morton, B
...
1983
...
Hong Kong: Hong
Kong University Press
...
T
...
Controlled manipulations in the
marine intertidal zone, and their contributions
to ecological theory
...
Academy of
Natural Sciences
...

Royal Observatory, Hong Kong (1993; 1994)
...
Hong Kong: The
Royal Observatory
...
D
...
Diets and habitats of shallow
water predatory gastropods around Tolo
Channel, Hong Hong
...
B
...
Proceedings of the First
International Workshop on the Malacofauna
of Hong Kong and Southern China, Hong
Kong, 1977 Hong Kong: Hong Kong
University Press
...
D
...
Field observations of prey
selection by the muricid gastropods Thais

Predation by Thais clavigera and Morula musiva on Tetraclita squamosa

clavigera and Morula musiva feeding upon
the intertidal oyster Saccostrea cucullata In
The Marine Flora and Fauna of Hong Kong
and Southern China (ed B Morton), 837-55
Proceedings of the Second International
Marine Biological Workshop The Marine
Flora and Fauna of Hong Kong and Southern
China, Hong Kong 1986 Hong Kong Hong
Kong University Press

Tong, L K Y 1986a The population dynamics
and growth of Thais clavigera and Morula
musiva (Gastropoda Muricidae) in Hong
Kong Asian Marine Biology 3 145-62
Tong, L K Y 1986b The feeding ecology of
Thais clavigera
and Morula
musiva
(Gastropoda Muricidae) in Hong Kong Asian
Marine Biology 3 163-78

Asian Marine Biology 12 (1995): 19-26

SPATIAL VARIATION IN LARVAL TREMATODE INFECTIONS OF
POPULATIONS OF NODILITTORINA TROCHOIDES AND NODILITTORINA
RADIATA (GASTROPODA: LITTORINIDAE) FROM HONG KONG
Tang Chong-ti
Parasitology Research Laboratory, Xiamen University, Xiamen, People's Republic of China 361005

Abstract
The incidence of larval trematode (xiphidiocercariae) infections in populations of Nodilittorina
trochoides and Nodilittorina radiata from shores in Hong Kong was investigated during October 1993
...
e
...
hezuiensis and C
...
Two of these species (Cercaria hezuiensis and C
...

Infection rates were low and varied between the sites examined
...
Populations at some
sites on the more exposed southeastern coast of Hong Kong Island had no infected individuals
...


Introduction
Littorinid snails are intermediate hosts to a variety
of trematode parasites which can influence growth
rates, reproduction, behaviour and subsequent
survival of their molluscan hosts (see Lauckner
1987 for a review)
...

The
consequences of trematode infections can,
therefore, also vary between populations, resulting
in demographic variations between sites
...

Infections by species of xiphidiocercariae
(Cercaria minus) have previously been recorded
from Nodilittorina vidua (= N
...

In this study, Nodilittorina were collected from
nine sites (eight on the southeast coast of Hong
Kong Island and one in the New Territories) and
the prevalence of parasites in these different

populations assessed to investigate spatial
variation in parasite loads
...


Materials and methods
Collections of Nodilittorina species were made
during October 1993 at nine sites in Hong Kong
(Fig
...
Animals were returned to the laboratory
where they were examined by crushing them and
dissecting out living larval trematodes
...

Results
Incidence of larval trematode infections in
Nodilittorina at different sites in Hong Kong
A total of 6341 specimens of Nodilittorina
trochoides (Gray) and N
...
1
...
The incidence of larval trematode infections of Nodilittorina trochoides and N
...

Mollusc
host

N°of
snails

Trematode
species

N trochoides

513

Cercaria minus
Cercaria hezuiensis
Cercaria tangi

Total

513

0
0
0
1

N radiata

471

Cercaria minus
Cercaria tangi

Total

471

0 637
0 425
1 062

TOTAL

984

N°

(%)
Infected

positive

12

97
194
194
365

1 22

Table 2
...
radiata
from different sites on the exposed southeastern coast of Hong Kong Island
...

vidua) (Reid 1992) collected from Starfish Bay in
April 1986
...
Infection rate was very low
...
vidua examined only one was
positive (0
...
In the present
investigation, this species was recorded from both
Nodilittorina trochoides and Nodilittorina radiata
collected from Starfish Bay
...
97% in N
...
64%
in N
...
millegrana
...
2
...

22

Plagiorchioidea
Cercaria hezuiensis sp
...

Specimens of the second xiphidiocercariae,
including cercariae and their daughter sporocysts,
were obtained from Nodilittorina trochoides from
Cape d'Aguilar and Lobster Bay on the
southeastern coast of Hong Kong Island and from
Starfish Bay in the New Territories
...
19%
...
55% and 0
...

Morphological characteristics of the cercaria
and daughter sporocyst
Daughter sporocyst (Fig
...
Cyst-shaped body
is between 0
...
71 mm in length and between


...
nov
...
1mm

Fig
...
A, Cercaria of Cercaria tangi sp
...
and B, sporocyst of Cercaria tangi sp
...


This species of xiphidiocercanal cercariae is
similar to the cercaria of
Maritreminoides
caridinae (Yamaguti and Nisimura, 1944) (Chen
1957) belonging to the Microphallidae
(Trematoda)
...
There
are, however, important differences between these
species, e
...
, the molluscan host of M
...
The two species
of cercariae also differ in the shape of stylet, the
length of tail and the structure of the excretory
bladder
...
caridinae, the tail
is shorter than the body and its excretory bladder
is V-shaped
...
nov
...
C
...


Discussion
To date, five species of xiphidiocercariae have
been obtained from marine molluscs in Hong
Kong
...
The other two species are
Cercaria spelotremoides (Tang 1990) parasitizing
the digestive glands of Cerithidea rhizophorarum
collected from Three Fathoms Cove in the New
Territories, and Cercaria armata (Tang 1990)
parasitizing Clypeomorus pellucida collected
from mangrove areas in the New Territories
...
Latypov
Institute of Marine Biology, Far East Branch, Russian Academy of Sciences, Vladivostok 690041, Russia

Abstract
Preliminary investigation of fringing reefs at nine localities in the western part of the Baitylong
Archipelago, Vietnam, were made along line transects to 15 m depth
...
Two morphological types of reef occur: structural and unstructural, both
showing distinct vertical zonation in terms of species composition and community structure
...

One hundred and forty-two species of Scleractinia belonging to 43 genera were identified, 80 species
and 15 genera of which have not hitherto been reported from the Gulf of Tonkin
...

As a result of marked erosive processes, they form
complex morphostructures with abrasive bays and
fiords deeply incising the islands
...
The underwater topography is a weakly
inclined, aggradative, plain alternating with
surface terraces at depths of 6, 9 and 11 m and
numerous patches of truncated bedrock
...

In the rainy season, salinity drops to between 2 8 30%o coinciding with periods of large tidal
fluctuations (to - 3
...
7 m)
...
Water
temperature also falls to 16-18°C in winter
(Gurjanova 1972; Yet 1989)
...
In the
western part of the gulf, however, there are the
well-known national reserves of Kat'ba Island and
Halong Bay about which virtually nothing is
known
...
There is one
publication on the distribution of coral reefs in the
northern Gulf of Tonkin (Latypov and Malyutin
1990)
...


Material and methods
A study of coral communities and reef structure
was conducted east of Kat'ba Island, the largest
of the islands of the Baitylong Archipelago
...
Distribution of corals was
studied on five islands using transect and quadrat
techniques (Loya and Slobodkin 1971)
...
l)
...
Qualitative samples of coral species were
collected from each physiographic zone at depths
of 1, 2-3, and 5-7 m
...
Corals were
identified by the author using the following
publications (Veron and Pichon 1976,1979,1982;
Latypov 1990,1992)
...
Small islands and their bays have
structureless coral reefs built by settlement of
individuals or groups of colonies, rarely by
bioherms (Fig
...
Reef deposits form only a
veneer; they do not form the reef framework and
match the natural profile of the bottom which
consists, for the most part, of large pieces of rock
rubble and, rarely, of dead coral fragments
...
1
...
B, Map showing
the study area and location (black circles)
of the transects
...
2
...
1,

Sargassum

polycystum; 2, Gracilaria verrucosa; 3, Padina australis; 4, Pontes; 5, Favia, Favites; 6,
Goniopora stokesi; 7, Cypraea tigris; 8, Holothuria atra; 9, Acropora nobilis; 10, Turbmana
peltata; 11, Galaxea fascicularis; 12, Diadema setosum; 13, corymbose Acropora; 14, Echinopora
lamellosa; 15, Fungia; 16, Lobophyllia hemprichii; 17, Cladiella sp
...
; 19,
Verrucella umbraculum; 20, Pavona cactus; 2 1 , Trochus niloticus; 22, Echinophyllia aspera
...
3
...
Ya
...
Distribution of Scleractinia on reefs of the Baitylong Archipelago and other regions of
the South China Sea
...

Species
1
...

3
...

5
...

7
...

9
...

11
...

13
...

15
...

17
...

19
...

21
...

23
...

25
...

27
...

29
...

31
...

33
...

35
...

37
...

39
...

41
...

43
...

45
...

47
...

49
...
)
P
...
- E
...

P
...
profundacella Gardiner
Astreopora myriophthalma (Lamarck)
+
Montipora aequituberculata Bernard
M
...
danae M
...
& H
...
digitata (Dana)
M
...
informis Bernard
M
...
spongodes Bernard
+
M
...
turtlensis Veron & Wallace
+
M
...
undata Bernard
+
+
M
...
samoensis (Brook)
+
+
A
...
robusta (Dana)
+
A
...
formosa (Dana)
A
...
valenciennesi (M
...
& H
...
microphthalma (Verrill)
+
A
...
aspera (Dana)
A
...
millepora (Ehrenberg)
A
...
yongei Veron & Wallace
+
+
A
...
hyacinthus (Dana)
+
A
...
lutkeni Crossland
+
A
...
clavus (Dana)
+
+
P
...
explanulata (Lamarck)
+
Leptoseris mycetoseroides Vaughan
+
Pachyseris rugosa (Lamarck)
P
...
) +
Coscinaraea columna (Dana)
+
Fungia repanda Dana
+

2

3

4

5

6

7

+

+
+

+
+
+
+

+
+
+
+

+
+
+

+

+

+

+
+
+

+

+

+
+
+
+

+

+
+

+
+

+
+

+

+
+

+
+

+

+

+

+

+

+

+
+

+
+

+
+
+

+
+
+

+

+
+
+

+
+

+
+

+
+
+

+
+
+

+
+
+

+
+

+

+

+

+
+
+

+

+
+

+
+

+
+

+
+
+

+

+

+
+
+
+

+

+

+
+

+
+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+
+
+
+
+

+

+

+

+

+

+

+
+
+

+
+

+

+

+

+

+
+

+

+
+
+

+
+

+
+

+

+

+

+

+

+

+
+

+

+

+

+
+

+
+
+

+
+

+

+

+

+

+

+
+

+

+
+

+

+

+
+
+
+

+
+

+

+

+

+

+
+

+
+
+

+
+
+

+
+
+

+

+

+

+
(to be continued)

30

Community structure of Vietnamese reefs
50
...

52
...

54
...

56
...

58
...

61
...

63
...

65
...

67
...

69
...

71
...

73
...

75
...

77
...

79
...

81
...

83
...

86
...

88
...

90
...

92
...

94
...

96
...

98
...

100
...

102
...

104
...

106
...


F
...
corona Doderlein
F
...
danai M
...
& H
...
mokai Hoeksema
L
...
djiboutiensis Vaughan
G
...
- E
...

G
...
lobata Dana
P
...
- E
...

P
...
mayeri Vaughan
P
...
stephensoni Crossland
P
...
species 1
Barabattoia amicorum E
...
& H
...
pallida (Dana)
F
...
favus (Forskal)
F
...
rotumana (Gardiner)
F
...
maxima Veron, Pichon & Best
Favites abdita (Ellis & Solander)
F
...
flexuosa (Dana)
F
...
pentagona (Esper)
Goniastrea retiformis (Lamarck)
G
...
favulus (Dana)
Platygyra daedalia (Ellis &
Solander)
P
...
pini Chevalier
P
...
- E
...

Australogyra zelli (Veron,Pichon & Best)
Oulophyllia crispa (Lamarck)
Hydnophora exesa (Pallas)
H
...
-E
...

Plesiastrea versipora (Lamarck)
Leptastrea purpurea (Dana)
L
...
pruinosa Crossland
Cyphastrea chalcidicum (Forskal)
C
...
Ya
...

109
...

111
...

113
...

115
...

117
...

119
...

121
...

123
...

125
...

127
...

129
...

131
...

133
...

135
...

137
...

139
...

141
...


C
...
gemmacea (Lamarck)
Oulastrea alta Nemenzo
O
...
astreata (Lamarck)
Merulina ampliata (Ellis & Solander)
Symphyllia recta (Dana)
S
...
-E
...

S
...
-E
...

S
...
-E
...

Lobophyllia corymbosa (Forskal)
L
...
hattai Yabe, Sug
...
-E
...
)
Pectinia lactuca (Pallas)
P
...
)
E
...
orphensis Veron & Pichon
Oxypora lacera (Verrill)
Mycedium elephantotus (Pallas)
Turbinaria frondens (Dana)
T
...
peltata (Esper)
T
...
stellulata (Lamarck)
T
...
cornigera (Lamarck)
D
...
-E
...

Tubastrea aurea (Quoy & Gaimard)
T
...

fascicularis, P
...

The reef slope community is divided distinctly
into two facies, i
...
, Acropora cytherea + Galatea
fascicularis on the upper part of the reef slope and
Cladiella sp
...
Towards the base of the reef slope (to a
depth of between 8-10 m) scleractinian richness
and individual numbers decreased
...

Structural Reefs
...
The inner reef flat consisted of
coralogenic flat cemented crusts and branching
colonies of Acropora, Montipora or Pavona and
massive heads of Porites, which here formed
monospecific strands
...

Macrophytic algae occurred among the corals
...
Colonies of
branched Pavona cactus, Acropora pulchra,

Community structure of Vietnamese reefs

Acropora millepora and Montipora
digitata
formed small areas of monospecific settlements
and massive P
...
There were also crusts of Palythoa
with areas of up to several square metres and
small algal mats of Gracilaria up to 1
...
The
external reef flat was cut with longitudinal and
cross grooves occupied mainly by encrusting and
massive colonies (between 10-14-nr 2 ) of G
...
cactus, F
...
lobata,
Hydnophora exesa
...
The
external part of some reef flats were between 2 1 60% covered by monospecific stands of P
...
digitata
or an up to 54% covering of different species of
Acropora,
i
...
, A
...

cytherea,
A
...
nobilis
...
Species of Pectinia, Mycedium,
Merulina, Turbinaria, Sinularia and Cladiella also
occurred
...

The reef slope was steep, with numerous
grooves and spurs densely inhabited by different
corals
...
e
...
cytherea, A
...
humilis, A
...
e
...
danae, M
...
hispida were
dominant
...
Goniopora stokesi occupied areas of
hundreds -mr2, and accumulations of Fungiidae, up
to 15-45-m-2, were dominated by Fungia danai
...
The lower part of
the reef slope (> 3-5 m) was occupied mainly by
massive scleractinians and encrusting, plate-like,
forms
...
At a depth
of between 9-10 m, only single scleractinian and
gorgonian corals occurred
...
In spite of such
extreme conditions, the reefs of this region have
a relatively high species richness and a high
degree of similarity with other Vietnamese reefs
and those of the Indo-Pacific (Table 1 and Fig
...

The reefs studied had 65
...
4% of species in
common with the reefs of southern Vietnam (Tchu
and Con Son Islands and others)
...
More than two-thirds
of the coral species are common to the coral
faunas of the Gulf of Tonkin and the Gulf of
Thailand (62
...


0
...
4
...
1-7:
localities are the same as in Table 1
...
5
...
6
...
1972
...
Marine Biology 13: 100-23
...
1976
...
Nature 259:47880
...
and Slobodkin, L
...
The coral reefs
of Eilat (Gulf of Eilat, Red Sea)
...

Maragos, J
...
and Jokiel, P
...
1976
...
Local distribution
...
Naval Undersea Research
...

Pichon, M
...
Dynamics of benthic
communities in the coral reefs of Tulear
(Madagascar); succession and transformation
of the biotopes through reef tract evolution
...

Pichon, M
...
Dynamic aspects of coral reef
benthic structures and zonation
...

Porter, J
...
1976
...
American Naturalist 110:
731-42
...
C, Done, T
...
, Isdate, P
...
and Fisk, D
...

1985
...
Marine Ecology
Progress Series 23: 79-84
...
H
...
Coral reefs of St
...
The Caribbean Journal of Sciences 12:
170-90
...
, Yeemin, T
...
, Yamazato,
K
...
1986
...
Galaxea 5: 21-1 A
...
1977
...
Helgolander wissenchaften
Meersuntersuchungen 3: 97-9
...
R
...
1979
...
Marine Ecology Progress Series
1: 237-47
...
R
...
1982
...
Marme
Ecology Progress Series 7: 83-115
...
,Stoddart, D
...
and Sigre, D
...

1971
...
Symposium of the Zoological Society
of London 28: 217-59
...
I
...
Coral reefs ecosystems
...

Taylor, J
...
1968
...
Philosophical
Transactions of the Royal Society of London
254: 129-206
...
A
...
Ya
...
D
...
Vliyanie sveta na raspredelenie
skleractinii v sublitorali ostrovov YuzhnoKitaiskogo morya
...
(In Russian)
...
W
...
H
...
A
description of the high latitude shallow water
coral communities of Miyakejima, Japan
...

VeronJ
...
N
...
1976
...
I
...
Australian
Institute of Marine Science
...

VeronJ
...
N
...
1982
...
IV
...

Australian Institute of Marine Science
...

VeronJ
...
N
...
1984
...
V
...

Australian Institute of Marine Science
...

Wainwright, S
...
1965
...

Bulletin of the Sea Fisheries Research Station,
Israel 8: 40-53
...
H
...
Research results of corals and
coral reefs in western Tonkin Gulf
...

Zhuang, Q
...
,Lu, B
...
1983
...
Studia Marina
Sinica 20: 1-50
...

C
...
Richardson, R
...
A
...
Owen
School of Ocean Sciences, University of Wales - Bangor, Menai Bridge, Gwynedd, LL59 5EY, U
...


Abstract
Age and growth of Septifer virgatus (Wiegmann) from six physically contrasting sites on a wave-exposed
rocky shore at Cape d'Aguilar, Hong Kong were determined from the pattern of tidally induced
microgrowth bands present in the outer nacreous layer of the shell
...
Two obvious groups of narrowly spaced bands are
deposited annually, one during the winter (February to March) when seawater temperatures are at their
lowest, the other during the summer (July to August) when mussels are probably subjected to elevated
levels of physiological stress
...
The maximum
estimated age of S
...
Significant
intersite variations in shell allometry are reported whilst within each population shells become
proportionately wider, more elongate and lighter with overall increase in body size
...
Wherever conditions for
attachment and growth are favourable, mussels
frequently become the dominant space-occupying
organisms (Seed and Suchanek 1992)
...
1987) and as structurally
complex entities also provide extensive habitats
for a wide range of associated fauna (Suchanek
1985; Ong Che and Morton 1992; Peake and
Quinn 1993; Lintas and Seed 1994)
...
Unlike their temperate
counterparts, however, these warmer-water
mussels have generally been much less intensively
studied
...
Thus, whereas P
...
1985),
S
...

In this paper we present some preliminary
data concerning the age structure and growth
characteristics of Septifer virgatus populations
from several contrasting habitats on a waveexposed shore at the southernmost tip of the Cape
d'Aguilar peninsula in Hong Kong
...
1
...
2
...
Photomicrographs of acetate peels of Septifer virgatus shell sections
...
Growth lines (arrowed)
in the umbo region (U)
...
C
...
D
...
Groups of
narrowly spaced tidal growth bands in the outer nacreous shell layer (NL)
...
Narrowly spaced
growth bands (left of figure) and a sudden growth disturbance resulting in a prominent cleft in
the shell (arrowed), (P), penostracum
...


were intermediate between these values (Table 1)
...
Shell height was
negatively allometric with respect to shell length
in all six populations
...
The growth constant (k) and asymptotic shell length (L°°) estimated from the von
Bertalanffy growth function for Septifer virgatus from Cape d'Aguilar, Hong Kong
...
Coefficient of allometry1 for various combinations of size parameters together with interpopulation comparisons4 in Septifer
virgatus from Cape d'Aguilar, Hong Kong
Dependent
variable2
Cog 10 )

Independent
variable3

y

x

Sheltered

F values4

Wave -exposed

('°9io>

high

low

high

low

subtidal

pool

slopes

intercepts

Height

Length

0 804*

0 775*

0 717*

0 712*

0 775*

0 822*

7 11t

-

Width

Length

1 060 ns

1 030 ns

1 020 ns

0 997 ns

1 030 ns

1 050*

315r

-

Width

Height

1 250*

1 270*

1 350*

1 340*

1 270*

1 200*

0 97

0 62

Shell Weight

Length

2 770*

2 710*

2 770*

2 450*

2 710*

2 680*

9 57f

-

Dry Flesh Weight

Length

3 250*

2 960 ns

2 500*

3 010 ns

2 990 ns

2 930 ns

6 19f

-

Dry Flesh Weight

Shell Weight

1 170*

1 090*

0 878*

1 150*

1 100*

1 080*

7 62f

-

1

The coefficient b in the allometric equation
The variables y and x respectively in the allometric equation
* Relationships which depart significantly from isometry at p < 0 05
ns
not significant
4
ANOVA with covariate, t analysis significant at p < 0 05 (F = 2 21 at 5,229 df)
23

Table 3
...

Dependent
variable 1
(i°g,o)
y

Independent
variable 2

Wave-exposed

Sheltered

( | 0 9io)
x

high
a

3

b

4

low
5

high

r

a

b

r

a

b

low
r

subtidal

pool

a

b

r

a

b

r

a

b

r

Height

Length

0
...
804

0 942

0
...
961

0
...
947

0 165

0
...
959

0
...
775

0
...
004

0
...
946

Width

Length

-0
...
060

0
...
444

1
...
969

-0
...
020

0 963

-0
...
997

0
...
444

1
...
902

-0
...
050

0
...
449

1
...
464

1
...
934

-0
...
909

-0
...
340

0
...
464

1
...
884

0
...
200

0
...
800

2
...
983

-3
...
710

0
...
880

2
...
981

-3
...
450

0
...
640

2
...
969

-3
...
680

0
...
610

3
...
966

-5
...
960

0
...
580

2
...
916

-5
...
010

0
...
050

2
...
950

-5
...
930

0
...
140

1
...
974

-1
...
090

0
...
070

0
...
882

-1
...
150

0 910

-1
...
100

0 965

-1
...
080

0
...

The constants b (slope) and a (intercept) respectively in the allometric equation
...


Age and growth of Septifer

3 4

5

High shore sheltered
n=1491

" V J M jj , 1 M V- W l1tyV , r - , r r , r r ,

-V-Vl

0 ]l
0-2

8-10

16-18

24-26

32-34

40-42

3 *

48-50

56-58

64-66

4

r

*

0-2

un)ln N
1 T 1
8-10

T^

16-18

High shore pool
n=551

r

1

I

I

24-26

-

r

T

j—f-

32-34

I

40-42

48-50

T

T

T

56-58

1

l

r

I

64-66

c
456

Low shore exposed

n=1352

0-2

8-10

16-18

24-26

tlVM

32-34

40-42

48-50

Pr-,
56-58

64-66

Shell length (mm)

3
...
Asterisks denote mean values of
individual size classes, arrow symbols the estimated age (yrs) determined from microgrowth
bands in population sub-samples
...
A
...
Seed, N
...
Brotohadikusumo & R
...
This, however, is usually possible only
when annual recruitment is relatively restricted
and where growth rates of individuals within each
cohort are fairly uniform (Cerrato 1980)
...
1990) ( Fig
...
Moreover,
although such distributions have previously been
used to estimate mussel growth (Bayne and
Worrall 1980; Rodhouse et al
...

Morton (1995) followed successive cohorts of
Septifer virgatus which recruited onto the shore at
Cape d'Aguilar and concluded that although this
population comprised six size (year) classes, the
majority of individuals within the population
probably had a life span of between four and five
years
...
Similarly, in the
present investigation, no individuals over six years
old were found, strongly suggesting that this could
be the normal longevity of Septifer in Hong Kong
waters
...

The absence of old Septifer
virgatus,
particularly from the high shore populations,
stands in marked contrast to previous reports for
...
No
S
...
3)
...
Such temperatures
are close to those which induce heat coma
(44
...
virgatus (Liu and Morton 1994)
...
Liu and Morton (1994), however, have
shown that mortality of S
...

Surface growth rings resulting from seasonal
changes in shell deposition have previously been
used with varying degrees of success to determine
the age of mussels (Seed and Richardson 1990)
...
1980; Richardson 1993)
...
Radial shell sections revealed a clearly
marked annual pattern in the deposition of these
tidally induced bands, with two groups of
narrowly spaced bands representing periods of
reduced linear growth, produced each year
...
Our data are,
therefore, wholly consistent with earlier reports
that two surface growth rings are deposited
annually in S
...

Although coupling between reproductive activity
and periods of reduced shell growth has
previously been described for other bivalves, e
...
,
Spisula solidissima (Dillwyn) (Jones et al
...

1995), no such correspondence between
reproduction and growth was observed in

Age and growth of Septifer

Chamelea gallina (Linnaeus) (Ramon and
Richardson 1992)
For many intertidal marine organisms,
including mussels (Seed and Suchanek 1992 and
references therein), growth rate is typically
inversely correlated with tidal level, thus
reflecting the progressively reduced feeding times
with increasing aerial exposure Maximum growth
rates of Septifer virgatus at Cape d'Aguilar
occurred amongst low intertidal and shallow
subtidal individuals at the more wave-exposed
locality, whereas the slowest growth rate, which
was comparable to that previously reported by
Morton (1995) from analysis of size-frequency
data was recorded amongst individuals from the
shallow high-shore pools (Fig 3) Although
permanently submerged, these pool individuals
will presumably periodically experience high
seawater temperatures, fluctuating salinities and
reduced oxygen levels during periods of
particularly calm weather in the summer months
Significant intersite variability in the
allometric relationships between Septifer virgatus
populations has been demonstrated Furthermore,
the general pattern of growth at each of the study
sites is for individuals to become proportionately

wider and more elongate with an overall increase
in body size, strongly suggesting that such
changes in shell form have adaptive significance
The development of a more elongate shell elevates
the posterior feeding currents above those of
neighbouring conspecifics, whilst increase in shell
length and width relative to shell height will
effectively enhance physical stability Our results
are therefore strikingly similar to those previously
described for Mytilus (Brown et al 1976, Seed
1978, 1980) which, like S virgatus, also occurs
in densely packed assemblages on wave-exposed
rocky shores

Acknowledgements
The authors are grateful for the following financial
support C A R (Royal Society travel grant), R S
(British Council), R O (British Council and the
Swire Educational Trust) and N A B (Unesco)
We are particularly grateful to Professor Brian
Morton, Director of the Swire Institute of Marine
Science (SWIMS), The University of Hong Kong,
for his hospitality and provision of facilities
during our visit

References
Anwar, N A , Richardson, C A and Seed, R 1990
Age determination, growth rate and
population structure of the horse mussel
Modiolus modiolus Journal of the Marine
Biological Association of the United Kingdom
70 441-56
Bayne, B L and Worrall, C M 1980 Growth
and production of mussels Mytilus eduhs from
two populations Marine Ecology Progress
Series 3 317-28
Bhattacharya, C G 1967 A simple method of
resolution of a distribution into Gaussian
components Biometrics 23 115-35
Brown, R A , Seed, R and O'Connor, R J 1976
A comparison of relative growth in
Cerastoderma edule, Modiolus modiolus and
Mytilus eduhs (Mollusca Bivalvia) Journal
of the Zoological Society of London 179 297315

Cerrato, R M 1980 Demographic analysis of
bivalve populations In Skeletal Growth of
Aquatic Organisms (ed D C Rhoads and R
A Lutz), 417-68 New York Plenum Press
Huang, Z G , Lee, S Y and Mak, P M S 1985
The distribution and population structure of
Perna viridis (L ) in Hong Kong In The
Malacofauna of Hong Kong and Southern
China II (ed B Morton and D Dudgeon),
465-71
Proceedings of the Second
International Workshop on the Malacofauna
of Hong Kong and Southern China, Hong
Kong 1983 Hong Kong Hong Kong
University Press
Jones, D S , Thompson, I and Ambrose, W 1978
Age and growth rate determinations for the
Atlantic surf clam Spisula
solidissima
(Bivalvia Mactracea), based on internal
growth lines in shell cross-sections Marine
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C
...
Richardson, R
...
A
...
Owen

Biology 47: 63-70
...
1982
...
Marine
Biology 68: 117-33
...
Y
...
The population dynamics of the
green mussel, Perna viridis (L
...
Asian Marine Biology
2: 107-18
...
Y
...
1985
...
In The Malacofauna of Hong Kong
and Southern China II (ed
...
Morton and D
...
Proceedings of the Second
International Workshop on the Malacofauna
of Hong Kong and Southern China, Hong
Kong 1983
...

Leigh, E
...
, Paine, R
...
, Quinn, J
...
and
Suchanek, T
...
1987
...
Proceedings of the
National Academy of Sciences of the United
States of America 84: 1314-18
...
and Seed, R
...
Spatial variation in
the fauna associated with Mytilus edulis on a
wave-exposed rocky shore
...

Liu, J
...
and Morton, B
...
The temperature
tolerances of Tetraclita squamosa (Crustacea:
Cirripedia) and Septifer virgatus (Bivalvia:
Mytilidae) on a sub-tropical rocky shore in
Hong Kong
...

Lutz, R
...
1976
...
Journal of the Marine
Biological Association of the United Kingdom
56: 723-31
...
1991
...
Malacological
Review 24: 115-18
...
1992
...
In The
Mussel Mytilus : Ecology,
Physiology,
Genetics and Culture (ed
...
Gosling), 21-52,
Amsterdam: Elsevier
...
1995
...
Journal of Zoology 235:
50

485-500
...
G
...
1992
...
Asian Marine
Biology 9: 217-33
...
and Quinn, G
...
1993
...
Ecography 16: 269-77'
...
, Abello, P
...
A
...
Population structure and growth of
Donax trunculus (Bivalvia: Donacidae) in the
western Mediterranean
...

Ramon, M
...
A
...
Age
determination and shell growth of Chamelea
gallina (Bivalvia: Veneridae) in the western
Mediterranean
...

Rodhouse, P
...
, Roden, C
...
, Burnell, G
...
,
Hensey, M
...
, McMahon, T
...
and
Ryan, T
...
1984
...
Journal of the Marine
Biological Association of the United Kingdom
64: 513-29
...
A
...
An analysis of the
microgrowth bands in the shell of the
common mussel Mytilus edulis
...

Richardson, C
...
1993
...
In
The Marine Biology of the South China Sea
(ed
...
Morton), 419-34
...
Hong Kong: Hong
Kong University Press
...
A
...
J
...
W
...
Factors influencing shell growth in
Cerastoderma edule
...

Richardson, C
...
, Seed, R
...
1990
...
Marine Ecology Progress Series
66: 259-65
...
1969
...

(Lamellibranchiata) on exposed rocky shores
...
Growth and mortality
...

Seed, R
...
The systematics and evolution of
Mytilus galloprovincialis Lmk
...
B
...
Beardmore), 447-68
...

Seed, R
...
Shell growth and form in the
Bivalvia
...
D
...
Rhoads and R
...
Lutz),
23-67
...

Seed, R
...
Taxonomic and evolutionary
relationships within the genus Mytilus
...

B
...
Hong Kong: Hong Kong
University Press
...
1992
...
American Malacological Bulletin 9: 123-137
...
and Brotohadikusumo, N
...
1994
...
In
The Malacofauna of Hong Kong and Southern
China III (ed
...
Morton), 427-44
...
Hong
Kong: Hong Kong University Press
...
and Richardson, C
...
1990
...

In The Neurobiology of Mytilus edulis (ed
...

B
...
Manchester: Manchester
University Press
...
and Suchanek, T
...
1992
...
In The Mussel
Mytilus : Ecology, Physiology, Genetics and
Culture (ed
...
Gosling), 87-169
...

Suchanek, T
...
1985
...
In The
Ecology of Rocky Coasts (ed
...
G
...
Seed), 70-96
...

Taylor, J
...
, Kennedy, W
...
and Hall, A
...

The shell structure and the mineralogy of the
Bivalvia
...
Bulletin of the British Museum (Natural
History) Zoology, Supplement 3: 1-125
...
Ong Che and Brian Morton
The Swire Institute of Marine Science, The University of Hong Kong, Cape d'Aguilar, Shek O, Hong
Kong

Abstract
This paper presents the results of a second phase of investigations of the subtidal macrobenthic
community at Tai Tam Bay, Hong Kong
...
Species assemblages were segregated by
hierarchical classification into three communities, i
...
, those supported by a silt-clay bottom at the inner
bay station, a sand bottom at the outer bay station and a mixture of silt-clay and sand at the middle of
the bay
...
1- Hyale grandicornis Branchiostoma belcheri community and the transition zone as the Chaetozone spp
...
Dominant species at each station are listed
...
Species diversity during the sampling period
averaged 2
...
46 and 4
...
Evenness and
species richness averaged 0
...
90, 0
...
62, 4
...
73 at the three stations, again respectively
...

Water quality of Tai Tam Bay showed a general improvement in 1989-1991, as compared with 19871989, evidenced by the increased species diversity index for the bay as a whole, the lack of inflection
in log-normal plots of species abundance and the increased number of low abundance species
...
Continued monitoring of the area is recommended and a list of indicator
species which occur in the intermediate abundance groups and which could be the focus of monitoring
is provided
...
It was gazetted as a Site of Special
Scientific Interest (SSSI) in 1974 (Agriculture and
Fisheries Department 1974) to supposedly protect
the diversity of intertidal communities occurring
there
...
At the tip of the latter is the Cape
d'Aguilar SSSI, Hong Kong's first proposed
marine reserve
...
G
...
Morton

productivity (Chan et al
...
The studies found
the bay waters to be generally 'clean'
...
The studies recommended
continued monitoring to delineate long-term
trends in the area, but at a reduced number of
stations, i
...
, three instead of five, in part to
determine if such a minimized data set could still
monitor the bay's benthic community effectively
...
Ecological studies
have been undertaken in the area over a period of
15 years and their findings are discussed in an
attempt to define overall trends in environmental
quality
...

Dominant species were also selected by means of
the Biological Index of Dominance (B
...
D
...
This method determines the relative
abundance of the species using rank-score analysis
on a 5-point system
...
The points for
each species over the entire study period were
totalled and the species ranked
Seasonal trends
Species diversity (H'), Evenness Index (J')
(Pielou 1966) and Species Richness (D')
(Margalef 1958) were calculated as follows :

1
Materials and methods
J' =

Collection and treatment of samples
Bimonthly sampling was carried out on board the
marine monitoring vessel of the Environmental
Protection Department of the Hong Kong
Government, for two years from March 1989 to
January 1990 (designated 1989 in this paper) and
from March 1990 to January 1991 (designated
1990)
...
5 m2) from three stations, i
...
,
Stations 1, 4 and 5 of the 1987-1989 study (Ong
Che and Morton 1991) in Tai Tam Bay and
situated in the inner, middle and outer regions of
the bay, respectively
...
1) as SM31,
SM34 and SM35, respectively
...
5 mm sieve and the
residue preserved in 5% neutral formalin stained
with Rose Bengal
...

Specimens were identified using the reference
collection lodged at The Swire Institute of Marine
Science, The University of Hong Kong
...

Seasonal variations in diversity indices were
examined by plotting them against time for each
station
...

The data were logarithmically transformed (In
(x + 1) ) before clustering
...
Clustering utilized the group
- average linkage which defines the distance
between two clusters as the average of the
distances between all pairs of cases in which one
member of the pair is also a member of one of the
clusters
...


-

—
• — • stm
O — O Stn4
• — • Stn5

0

H

1

1

h

1
...
1
...


neighbour technique) (Clifford and Stephenson
1975; Norusis 1988)
...
Log-normal plots of the cumulative
percent of species in different geometric

classes (x 2 scale) were constructed for inner,
middle and outer bay stations, i
...
, stations 1, 4
and 5, using the combined faunal data for each
year
...
1983)
...
G
...
Morton

Results
Faunal analysis
The 5741 individuals collected from three stations
within Tai Tam Bay were distributed among 85
species of which annelids, molluscs and
arthropods together comprised more than 82% of
the total number of individuals and more than 90%
of the total number of species recorded
...

Table 1
...

A
...
3% to 32
...
5% to
8
...
This may be due to the bias
arising from high catches of some species in some
seasons
...
I
...
scale
...
In 1989, all
of the dominant species except one were
polychaetes, whereas in 1990, polychaetes
comprised only half of the dominant species list
...
The remaining 72% of the individuals
recorded comprised species occurring in the low
abundance spectrum
...
Half of these were
amphipods
...
While absolute ranking changed
between the two sampling years, the dominant
species selected by the two methods remained
largely the same
...


B
...
I
...
) obtained for stations 1, 4 and 5 in 1989 1990
...
Dominance
ranking, based on the two methods used, differed,
however
...
6 % to 16
...
and Sigambra
tentaculata
56

Table 3 shows the mean, median, maximum and
minimum values of Species Diversity, Evenness
and Species Richness at the three stations in Tai
Tam in 1989 and 1990
...
52 and 3
...
44 and 4
...
T - tests comparing diversity indices
between years indicate, however, that differences
were not significant
...
76, 4
...
36 at Stations 1,
4 and 5, respectively
...
(1989), Station 1 within
Tai Tam Bay is moderately polluted ( H' = 1 - 3)
while Stations 4 and 5 enjoy clean waters ( H' >
3 )
...
90) as
compared with 0
...
84 at

Tai Tam subtidal benthos

Table 2
...
I
...
A, 1989; B, 1990
...
G
...
Morton

Table 3
...
T-test results
comparing diversity values between years are also presented
...
52
2
...
26
1
...
28

3
...
96
3
...
49
n
...


0
...
76
0
...
41
-0
...
73
0
...
82
0
...
s
...
37
2
...
75
2
...
89

2
...
95
3
...
29
n
...


4
...
58
5
...
01
•0
...
44
4
...
93
4
...
s
...
92
0
...
94
0
...
32

0
...
91
0
...
74
n
...


4
...
59
5
...
14
-1
...
12
4
...
26
4
...
s
...
44
4
...
92
3
...
50

4
...
28
5
...
70
n
...


0
...
87
0
...
63
-1
...
87
0
...
97
0
...
s
...
03
6
...
69
4
...
28

5
...
71
8
...
19
n
...


Station 4
Mean
Median
Maximum
Minimum
t-test
Station 5
Mean
Median
Maximum
Minimum
t-test

Station 5
...
69 and 5
...
62)
...
Species
diversity indices at the outer stations (4 and 5)
were consistently higher throughout the year, as
compared with the inner station (1)
...

Evenness was relatively constant at Station 4, with
only a slight decline in September 1990 and
March 1991
...
The reductions in evenness
which occurred in summer (July to September) at
these stations were more pronounced in 1989 than
in 1990
...
g
...
at
Station 1, and Corophium sp
...
1, Ong Che and Morton
1991) at Station 5 accounted for the decrease in
evenness during this period
...
scabra
and Minuspio sp
...
Relative abundance of
Corophium sp
...
grandicornis reached 174
and 132 individuals-sample-1, respectively
...

2 (188 individuals-sample -1 , Musculus nanus
(140), Podocerus sp
...

sagamiensis (168) all of which are recruited in
winter
...
Species diversity, as reported in the
first study on the Tai Tam benthos (Shin 1977),
averaged 3
...
Diversity decreased to 3
...
2
...


study (H' = 3
...
Following the criteria specified
in Jhingran et al
...

Hierarchical clustering
Results of the hierarchical classification of the
1989 - 1990 species abundance data are presented
in Figure 3
...

The present analysis of data obtained from
three stations with three replicates each has
verified the community patterns defined by the
classification of the 1987 - 1988 faunal data
obtained from five stations with five replicates
each (Ong Che and Morton 1991)
...
station column, Figure 3),
indicating differences in the species composition
of the three areas of the bay, i
...
, Station 1 in inner
Tai Tam, Station 5 in outer Tai Tam and Station
4 in between these two areas
...
e
...


Some interspersion of contiguous stations
occurs, however, reflecting an overlap in species
distributions between stations
...
This suggests that the benthic
communities of Tai Tam Bay are not strictly
delimited and conform to the concept of
communities as a continuum of species
distributions (Gray 1981)
...

The analysis of station, replication and time
(month, year) groups shows that replicates were
representative of the area sampled
...
replicate
column, Figure 3)
...
Seasonal cycles in benthic communities
59

R
...
Ong Che & B
...
3
...
Squared Euclidean distances on the ordinate have been rescaled
to fall within the range 1 - 25
...

60

Tai Tam subtidal benthos

are strongly influenced by annual variations in
temperature, light, primary production and related
factors (Gray and Mirza 1981)
...
5-21
...
Distinctly
higher temperatures occur in summer (25
...
1°C)
...
season and date columns, Figure 3)
...
A single, steep, slope was obtained for
all the stations in both years
...

Figure 5 presents log-normal plots of the
number of species against the number of
individuals-species-1 in geometric classes for the
consolidated faunal data (1987 - 1990) from
Stations 1, 4 and 5 at Tai Tam
...
The range
of geometric classes decreased from between 8 to
10 in the 1987 - 1988 sampling period to between
6 to 8 in 1989 - 1990, indicating that fewer species
in the higher abundance classes were recorded
during the second phase of sampling
...
4
...

61

R
...
Ong Che & B
...
5
...


When the lower abundance groups were
compared over the two sampling periods, more
species were recorded in low numbers (geometric
classes I to IV) in 1989 - 1990
...

Similarly, the percentage of species in the
intermediate abundance group (geometric classes
V and VI) decreased over the two sampling
periods at both Stations 1 and 5, i
...
, from 5% to
1% and from 8% to 1%, respectively
...

62

Pearson et al
...

The Tai Tam log-normal plots reflect this
situation, especially in 1989-1990
...
None of these species has been
reported to be pollution indicators
...
Examples of such
related species are Chaetozone
setosa,
Lumbrinereis latreilli and Prionospio malmgreni
(Food and Agriculture Organization 1986)
...
Species in abundance classes V and VI at the three stations in Tai Tam Bay sampled in
1989-1990
...

Pseudopolydora sp
...

Chaetozone spp
...

Sigambra tentaculata
Aglaophamus toloensis

Veremolpa scabra
Prionospio saccifera

Station 1

Station 4
1989
1990

Mediomastus californiensis
Chaetozone spp
...
1
Glycera alba (Fauvel, 1923)
Hyale grandicornis
Loimia medusa
Lumbrinereis nagae (Gallardo, 1968)
Minuspio sp
...

Prionospio sp
...
1
Amphipoda sp
...
1
Podocerus sp
...

Colelepis sp
...


Discussion
Community structure
Benthic communities have traditionally been
characterized by the dominant species occurring
in them
...

In their study of the spatial distribution of the
63

R
...
Ong Che & B
...

They declined to name these five communities
after the three most dominant species, however,
because of low fidelity
...

In the present study, low fidelity was also
observed, resulting from communities not being
delineated by rigid boundaries
...
The
abundance of these species at the different stations
differed, however
...
, for example,
comprised ~ 20% of the total individual numbers
at Station 1 but less than 3% at Station 5
...
comprised = 18% of the
total abundance at Station 5 in contrast to ~ 4%
at Station 1
...
According to Brown, a
species is most abundant at the centre of its niche
and decreases in numbers towards the edge
...

Communities can be named after the dominant
species which occur at the centre of their niches
...
I
...
)
ranking of Fager (1957) selects the numerically
important and recurring species sensu Petersen
...
I
...
ranks of the five
most dominant species over the four sampling
years at each station, the communities at the three
stations within Tai Tam Bay can be named (Table
5)
...

of decreasing dominance), Station 4, in the
transition zone, as the Chaetozone spp
...
1 - Hyale grandicornis —
Branchiostoma
belcheri
community
...

64

In a study on the benthic communities of Tolo
Harbour and Mirs Bay, Shin (1990) delineated
three major location groups by agglomerative
hierarchical classification, representing different
infaunal communities in Tolo Harbour, Tolo
Channel and Mirs Bay
...
Thus, the urchin Schizaster
lacunosus, the ophiuroid Ophiura kinbergi and the
venerid bivalve Paphia undulata were confined to
Mirs Bay, the polychaetes Prionospio ehlersi and
Lagis koreni to Tolo Channel and the polychaetes
Sigambra tentaculata, Minuspio cirrifera, Tharyx
sp
...

Of the polychaetes dominant in Tolo Harbour,
two are associated with areas of environmental
stress
...
Sigambra tentaculata has also
been reported in great numbers in an anoxic basin
at Golfo Dulce, Costa Rica by Nichols-Driscoll
(1976)
...
1977;
Thompson et al
...

In the present study, both Sigambra
tentaculata and Minuspio sp
...
This interesting parallel in the pollution
indicating polychaete species occurring at inner
Tolo Harbour and inner Tai Tam Bay suggests that
the environmental changes which have been
affecting the Tolo Harbour infaunal community
may be operating at inner Tai Tam Bay as well,
albeit at a less intense degree
...
Lui (1991) reported
temperature stratification of the water column in
Tai Tam during the summer months and that this

Tai Tam subtidal benthos

Table 5
...
The five most dominant species and
the mean B
...
D
...
Ties were given average ranks and species not among
the top five that year were given an arbitrary rank for calculation of the mean B
...
D
...

Station 1

1987

1988

1989

1990

Veremolpa scabra
Minuspio sp
Aglaophamus toloensis
Chaetozone spp
Theora lata
Copepoda sp
Nephtys polybranchia (Southern, 1921)
Pnonospio saccifera
Mensca sp
Diastolopsis sp 1

1
25
3
5

45

1
25
25
5
6

1
68
4
5
66
6

2

1
45
2
6

Mean
35
34
80
98
62
94
88
98

Station 4
Sigambra tentaculata
Aglaophamus toloensis
Sternaspis scutata
Chaetozone spp
Mediomatus californiensis
Ampelisca cyclops
Amphipoda sp 8
Caprella aequilibra
Pnonospio saccifera
Golfmgia misakiana
Stenothoe sp
Prionospio sp
Magelona crenulifrons
Nephtys polybranchia
Diastolopsis sp 1

3

3
1
2

1
3
2
5
4

6
15
15
3
4
6
6
8

1
2
5

4
5
4

11 5
98
11 8
31
81
60
122
11 5
11 5
120
11 5
11 8
122
100
122

Station 5
Corophium sp 1
Stenothoe sp
Diastolopsis sp 1
Ampelisca cyclops
Prionospio sp
Branchiostoma belcheri
Hyale grandicornis
Veremolpa scabra
Golfmgia misakiana
Podocerus sp
Ophiura kmbergi

3
2
1
55
55
4

was more pronounced in the inner bay as
compared with the outer Stratification of the
water column restricted vertical oxygen transfer,
resulting in a depletion of bottom water oxygen
levels Periodic subtidal anoxia, exacerbated by
poor tidal flushing in the inner bay could be

1
2
3
4
5

2
5
5
4
1

3
4

3
1
2

42
65
90
52
80
58
43
92
90
85
88

limiting for the community there and thus account
for the lower species diversity recorded from this
station
The Tai Tam subtidal macrobenthos provides
a rich fauna for taxonomic study Jiang and Xu
(1992) reported upon a new mollusc collected
65

R
...
Ong Che & B
...
nov
...
may also be an as yet undescribed species
(Mackie, personal
communication)
...

Ecological monitoring
The water quality of Tai Tam Bay, as reflected in
the data presented here of an overall increase in
benthic species diversity, apparently showed a
general improvement during the second phase of
benthic sampling in 1989-1990
...
In addition,
most of the species occurred in the low abundance
groups
...

The Environmental Protection Department of
the Hong Kong Government reported that the
Water Quality Objectives (WQO) in the Southern
Water Control Zone, which includes Tai Tam Bay,
had largely been achieved in 1989 and 1990
...
O
...
Levels of inorganic nitrogen, organic
nitrogen, inorganic phosphate and mean surface
chlorophyll a during this period were all below the
cut-off criteria for eutrophication, i
...
, 0
...
3 mg-1-1, 0
...

The mean B
...
D
...
coli counts were similarly low
...

In a concurrent study of the phytoplankton
dynamics of Tai Tam Bay, Chiu et al
...
40 to 32
...
g-1-' at the surface and
from 0
...
91 u
...
Two of
66

their study sites (SM31 and SM34) are the same
as Stations 1 and 4 of this study
...
e
...
The
only substantially higher chlorophyll a level was
measured at the last sampling date of their study,
i
...
, May 1991 (> 30 fig-1 ' at Station 1 and > 15
(ig-1-1 at Station 4 for both surface and bottom
waters)
...
No unusual
increase in May 1991 was obtained for nitratenitrogen and ortho-phosphate content in the study
of Chiu et al
...

Although Tai Tam Bay waters are considered
to have exhibited a general improvement in water
quality during this second phase of monitoring, the
continued dominance of Sigambra tentaculata, an
indicator of moderate pollution (Shin 1990) and
the marginal species diversity index recorded in
this study for the inner bay (Station 1) remain as
points of concern signalling the fragility of the Tai
Tam Bay system
...
(1994) in
May 1991 indicate fertile water conditions
...

Much of the baseline information on the Tai
Tam subtidal macrobenthic community have been
collated in this and a previous study (Ong Che and
Morton 1991)
...

The present study has demonstrated that the
three-station-three-replicate programme yielded
results comparable with those obtained in the first
study using a five-station-five-replicate protocol
...
G
...
Morton

Patna, Bihar, India
...

Jiang J
...
and Xu F
...
1992
...

Journal of Oceanography of Taiwan Strait
11:291-93
...
H
...
Water quality and sediment
chemistry of Tai Tam Bay, Hong Kong
...

Margalef, D
...
1958
...
General Systems Yearbook 3:36-71
...
B
...
S
...
The fauna of
benthic sediments from the organically
enriched Oslofjord, Norway
...

Morton, B
...
Marine pollution induced
changes in Hong Kong — the Tolo Harbour
case history
...
M
...
W
...
W
...

Hoare, P
...
Holmes, R
...
S
...
B
...
Polmet '85, Hong Kong
...

Nichols-Driscoll, J
...
Benthic invertebrate
communities in Golfo Dulce, Costa Rica, an
anoxic basin
...

Norusis, M
...
1988
...
Chicago : SPSS
Incorporated
...
G
...
1991
...
Asian Marine Biology 8:193216
...
H
...
S
...
J
...
Objective selection of sensitive species
indicative of pollution-induced changes in
benthic communities
...
Data analysis
...

Petersen, C
...
J
...
On the animal communities
of the sea bottom in the Skagerrak, the
Christiania Fjord and the Danish waters
...


68

Pielou, E
...
1966
...
American Naturalist 100:463-65
...
K
...
1977
...
M
...

Shin, P
...
S
...
Benthic invertebrate
communities in Tolo Harbour and Mirs Bay :
a review
...
B
...
Proceedings of the Second
International Marine Biological Workshop :
the Marine Flora and Fauna of Hong Kong
and southern China, Hong Kong 1986
...

Shin, P
...
S
...
B
...
Spatial
distribution of the infaunal benthos of Hong
Kong
...

Thompson, G
...
and Shin, P
...
S
...
Sewage
pollution and infaunal benthos of Victoria
Harbour,
Hong
Kong
...

Thompson, G
...
, Wu, R
...
S
...
J
...

1982
...
In The
Marine Flora and Fauna of Hong Kong and
Southern China (ed
...
Morton and C
...

Tseng), 745-60
...
Hong
Kong : Hong Kong University Press
...
H
...
F
...
S
...
N
...
A preliminary survey of
organic pollution of shellfish in Tolo Harbour,
Hong Kong
...

Wu, R
...
S
...
Periodic defaunation and
recovery in a subtropical epibenthic
community in relation to organic pollution
...


Asian Marine Biology 12 (1995) 69-78

OCCURRENCE OF THE BARNACLE BALANUS AMPHITRITE DARWIN ON
THE SHELLS OF THE PREDATORY GASTROPOD THAIS CLAVIGERA
KUSTER AT CAPE D'AGUILAR, HONG KONG
J
...
Liu1 and L
...
H
...
Lu

squamosa suffers heavy predation pressure on the
shores of Cape d'Aguilar, Hong Kong (Liu,
unpublished data)
...

amphitrite settling on the shells of T
...
This study examines the occurrence of
Balanus amphitrite on a wide variety of molluscan
shells at Cape d'Aguilar, Hong Kong, to determine
habitat choice between different hosts
...
Site 1 was
located on the northern shore of Lobster Bay on
the westen shore of Cape d'Aguilar and is
exposed to moderate wave action
...
1)
...
5 m and
2
...
e
...
1
...
Areas of 0
...
The numbers of
individuals of each species
...
09 m2 were recorded
and the shell length of each individual was
measured to the nearest 0
...
Barnacle settlement on each shell was
recorded in terms of either the presence or absence
of individuals of Balanus amphitrite
...
amphitrite on each host were counted and
their shell lengths measured, again to the nearest
0
...
This work was undertaken during
October 1993
...
The percentage frequency
of occurrence of one individual of B
...
e
...
Only one shell from the exposed site
hosted 12 individuals of B
...

Figure 3 shows the relationship between
Thais clavigera host size and numbers of attached
barnacles
...
e
...
At site
2, i
...
, the exposed shore, there was no strong
relationship between host size and numbers of
barnacles
...
A total of 46 individuals
of Thais clavigera were examined and 14 of these
(30%) were used as a substratum by B
...

The total number of B
...
Additionally, 641 individuals
of Tetraclita squamosa, 29 individuals of Cellana
toreuma,
159 individuals of
Patelloida
saccharina, 2 individuals of Nerita albicilla and
18 individuals of Liolophura japonica were
examined, but only 1 individual of C
...
amphitrite
...
clavigera were examined and
27 of these (58%) were used as a substratum by
B
...
The total number of B
...
Additionally,
1436 individuals of T
...
toreuma, 3 individuals of P
...
japonica were examined but
none of these was used as a substratum by B
...
A narrow range of gastropod species
was settled upon by B
...
e
...
clavigera
...
amphitrite were 5
...
9 mm on the exposed shore
...
One individual may
occur on one host shell but there may also be a

Discussion
The present work shows that Cape d'Aguilar
supports a population of Balanus amphitrite
which largely occurs on the shells of the
predatory snail Thais clavigera
...
Bourget (1989) listed 12
environmental factors which have been shown to
influence the settlement of intertidal barnacles
...

From an examination of 37 large clusters of
the gooseneck barnacle, Pollicipes polymerus,
Chia (1989) showed that 81% of juveniles are
attached to the peduncles of adults
...

Mapstone et al
...
This
preference is not explained by the relative shell
area of A
...
mufria would be in pools
during low tide, since this is the primary habitat
of this potential host
...
The percentage frequency of each species used as a substratum by Balanus amphitrite from a semi-exposed shore at Cape
d'Aguilar, Hong Kong
...
of host
examined

Total number of
Balanus found

Shell length of host (mm)
Mean ± S
...

Max
...


Size

14

30

46

60

16
...
15)

25
...
3

46

Tetraclita squamosa

0

0

641

0

23
...
43)

35
...
7

60

Cellana toreuma

1

4

29

2

17
...
30)

22
...
1

29

Patelloida saccharina

0

0

159

0

11
...
28)

15
...
9

33

Nerita albicilla

0

0

2

0

17
...
3

15
...
0

35
...
5

18

Shell length of Balanus

Mean ± S
...

5
...
71)

Max
...
5

(4
...

1
...
The percentage frequency of each species used as a substratum by Balanus amphitrite from an exposed shore at Cape
d'Aguilar, Hong Kong
...
of host
examined

Total number of
Balanus found

Shell length of host (mm)
Mean ± S
...

Max
...


Size

Thais clavigera

27

58

47

88

21
...
92)

27
...
7

47

Tetraclita squamosa

0

0

1436

0

17
...
68)

32
...
1

57

Cellana toreuma

0

0

53

0

13
...
35)

20
...
0

23

Patelloida saccharina

0

0

3

0

9
...
32)

9
...
2

3

Liolophura japonica

0

0

18

0

34
...
99)

44
...
0

18

Mean ± S
...


Max
...


3
...
36)

8
...
3

Shell length of Balanus:

Balanus amphithnte on the gastropod Thais clavigera

10i
Site 1

o
c

C
D
O"
C
D

5-

0
10
Site 2

o
c

CD

O"
C
D

0

—T—

10
No's of individuals of Balanus amphitrite
...
2
...
H
...
Lu

15n
<>
1

CtJ

CO

Site 1

y=1
...
254
r = 0
...
007X +3
...
007
•

&
cxj
CO
=3

10•

•S?

•

CtJ

CQ

o
CO

•
•

5-

•

a5

...
3
...


the shells of other gastropods
...
amphitrite settles randomly but only those
attached to the shells of T
...
These authors all record that limpets reduce
the survival of post-settlement barnacles
...
At high densities, L
...
Experiments on the larval settlement of
B
...
Settlement
of B
...
clavigera may help us understand
adaptation and natural selection
...
Table 3

Balanus amphithnte on the gastropod Thais clavigera

Table 3
...

Mollusca host

Barnacle

Reference

Pecten maximus (scallop)

Balanus balanoides

Barnes 1953

Chlamys opercularis (scallop)

Balanus balanoides

Barnes 1953

Anadara subcrenata (clam)

Balanus cirratus

Cai and Huang 1984

Anadara granosa (clam)

Balanus cirratus

Cai and Huang 1984

Perna viridis (mussel)

Balanus trigonus

Cai and Huang 1988

Mytilus crassitesta (mussel)

Balanus trigonus

Cai and Huang 1988

Xenostrobus atrata (mussel)

Balanus amphitrite

Cai and Huang 1993

Saccostrea cucullata (oyster)

Balanus amphitrite

Rainbow 1990

Septifer virgatus (mussel)

Balanus amphitrite

Liu, pers
...
Gastropod hosts used by other species
...


Crepidula walshi
(limpet-like snail)

Yipp 1980

Calliostoma ligatum

Crepidula adunca
(limpet-like snail)

Vermeij et al
...
) lampanicola
(limpet)

Morton 1980, 1988

Austrocochlea

Patelloida mufria
(limpet)

Mapstone et al
...
The molluscan hosts are
typically bivalves, e
...
, scallops, clams, mussels
and oysters
...
g
...

We did not investigate the differences between
the shells of Thais clavigera and the shells of
other gastropods in terms of surface texture,
surface contour and bacterial film
...

Acknowledgements
JHL is grateful to The University of Hong Kong
for awarding her a Rayson Huang Postdoctoral
Fellowship
...

75

J
...
Liu & L
...
1953
...

Bourget, E
...
Barnacle larval settlement: the
perception of cues at different spatial scales
...
M
...
Chelozzi)
...

Branch, G
...
1981
...
Oceanography and Marine
Biology Annual Review 19: 235-380
...
X
...
G
...
Studies on the
orientation of cirripedes
...
The orientation of
Balanus cirratus on the hosts
...

Cai, R
...
and Huang, Z
...
1988
...
II
...
Oceanologia et
Limnologia Sinica 19: 321-7
...
X
...
G
...
The orientation
of cirripedes on their hosts from Hong Kong
waters
...
B
...

Proceedings of the First International Conference on the Marine Biology of Hong Kong
and the South China Sea, Hong Kong 1990
...

Chia, F
...
1989
...
In Proceedings
of the 23rd European Marine Biology
Symposium (ed
...
S
...
A
...
Fredensborg, Denmark: Olsen and
Olsen
...
H
...
Effects of competition,
predation by Thais lapillus, and other factors
on natural populations of the barnacle
Balanus balanoides
...
A
...
Shallow water barnacles from
Hong Kong
...
B
...
K
...

Proceedings of the First International Marine
Biological Workshop: The Marine Flora and
Fauna of Hong Kong and Southern China,
Hong Kong 1980
...

Harding, J
...
1962
...
Bulletin of the
British Museum (Natural History) Zoology 9:
273-96
...
S
...
Occurrence of the barnacle
Balanus trigonus Darwin on the asteroid
Anthenea flavescens Gray
...
B
...
Proceedings
of the Fourth International Marine Biological
Workshop: The Marine Flora and Fauna of
Hong Kong and Southern China, Hong Kong
1989
...

Lewis, J
...
1954
...

Liu, J
...
1995
...
Asian Marine Biology 12: 11-17
...
S
...
Costlow, J
...
and
Mitchell, R
...
Inhibition of attachment of
larval barnacles, Balanus amphitrite, by
bacterial surface films
...

Mapstone, B
...
, Underwood, A
...
and Creese,
R
...
(1984)
...
Marine Ecology Progress Series 17: 85-100
...
1980
...
(C
...
Journal
of Experimental Marine Biology and Ecology
47: 149-71
...
1988 Partnerships in the Sea: Hong
Kong's Marine Symbioses
...

Morton B
...
1983
...
Hong Kong: Hong
Kong University Press
...
S
...
Grazing patterns of the

Balanus amphithrite on the gastropod Thais clavigera

periwinkle and their effect on sessile intertidal
organisms
...

Rainbow, P
...
1990
...
In The Marine Flora and Fauna
of Hong Kong and Southern China II (ed
...

Morton), 567-8
...
Hong
Kong: Hong Kong University Press
...
D
...
Diets and habitats of shallow
water predatory gastropods around Tolo
Channel, Hong Hong
...
B
...
Proceedings of the First
International Workshop on the Malacofauna
of Hong Kong and Southern China, Hong
Kong 1977
...

Taylor, J
...
1990
...
In
The Marine Flora and Fauna of Hong Kong
and Southern China II (ed
...
Morton), 837-

55
...
Hong Kong: Hong
Kong University Press
...
K
...
1986
...
Asian Marine
Biology 3: 163-78
...
1970
...
IX
...
Publications
of the Seto Marine
Biological Laboratory 17: 339-72
...
J
...
B
...
J
...
A
...
Good hosts and their
guests: relations between trochid gastropods
and the epizoic limpet Crepidula adunca
...

Yipp, W
...
1980
...
M
...
C
...
Parsons1, Mary L
...
Porter1
' The Swire Institute of Marine Science The UniversiU of Hong Kong Cape d Aguilar, Shek O Hong
Kong
Ecosystems Ltd Hong Kong

Abstract
There are records for Hong Kong territorial waters either suggesting or confirming the presence of fifteen
species of Cetacea three members of the Balaenoptendae (Minke whale, Bryde's whale and Fin whale),
one of Phocoemdae (Finless porpoise), one of Physeteridae (Pygmy sperm whale), and ten of Delphinidae
(Bottlenose dolphin, False killer whale, Fraser's dolphin, Indo-Pacific humpbacked dolphin, Long-beaked
common dolphin, Pantropical spotted dolphin, Spinner dolphin, Striped dolphin, Rough-toothed dolphin
and Risso's dolphin) Of these, only two appear to be present year round in Hong Kong waters the
Indo Pacific humpbacked dolphin and the Finless porpoise
Introduction
Hong Kong is situated adjacent to the mouth of
the Pearl River on the south coast of the Chinese
province of Guangdong (Fig 1) The first western
records of cetaceans in Hong Kong date from
1637, when the adventurer Peter Mundy reported
that 'the porpoises here are as white as milke,
some of them ruddy withall' (Carnac-Temple
1919) This was what is now known to be the
'local' population of Indo-Pacific humpbacked
dolphin, Sousa chinensis (Osbeck, 1765) Based
on his own observations during voyages in the
eighteenth century, Osbeck (1765) classified
similar animals he found in Hong Kong as a
species new to science, the Chinese white dolphin
Since then and until relatively recently, little has
been done to study Hong Kong's cetacean
populations Official sighting and stranding
reports first began to be assembled by the
Agriculture and Fisheries Department of the Hong
Kong Government in 1973 Body measurements
(albeit incomplete) of stranded carcasses date from
1985 Reports were made largely by government
employees until the World Wide Fund for Nature
Hong Kong established, in 1989, a scheme to
prompt members of the public to report details of

their casual sightings The initial findings from
these various programmes suggested that the
waters of Hong Kong, surprisingly, were
frequented by several cetaceans, including at least
two resident species (Morton et al 1992) the
Indo-Pacific humpbacked dolphin, known locally
as the Chinese white dolphin, and the Finless
porpoise, Neophocaena phocaenoides (Cuvier,
1829) It was in the climate of awareness created
by WWF's high-profile public reporting
programme, together with lobbying from
conservation organizations, that the Hong Kong
Government initiated a study of cetaceans in Hong
Kong waters, including the threats facing them
and possible mitigations to those threats This
study is being undertaken by a full-time research
team at the Swire Institute of Marine Science The
University of Hong Kong
The following is a list of species occurring in
Hong Kong The correct classification ot sightings
and stranding data can only be contnmed,
however, in cases wheie there are either voucher
specimens or unambiguous photographs ot the
animals The reliability ot other reports is
unconfirmed The taxonomy used here follows
Honacki etal (1982), Barnes et al (1985), Pernn
(1989) and Heyning and Pernn (1994)

E C M Parsons M L Felley & L J Porter

English common names follow Pernn (1989)
Details on which CITES appendix the species
appear in are from Khnowska (1991)

Species Accounts
Order Cetacea Bnsson, 1762
Suborder Mysticeti Flower, 1864
Baleen whales
Family Balaenopteridae (Gray, 1864)
Rorquals
Subfamily Balaenopterinae (Gray, 1864)
Balaenoptera acutorostrata Lacepede, 1804
Minke whale
[CITES Appendix I]
Minke whales occur throughout the world's seas
They are mainly oceanic but sometimes come
close to shore Wang (1984a, 1991a, 1993) has
reported Minke whales from the Bohai, Yellow,
East China and South China Seas As would be
expected from their cosmopolitan distribution,
Minke whales have been reported from Hong
Kong waters There are currently three recognised
populations of this species (Omura 1975) and the
whales described in Hong Kong were probably
members of the population known to winter off
the coast of Japan (Balaenoptera acutorostrata
davidsoni Scammon, 1872) There have been
several unsubstantiated reports of Minke whales
in Hong Kong waters and one confirmed sighting
of two individuals in Sai Kung (Anonymous 20
May 1991) There have been only two Minke
whale strandings since records began to be kept
(Fig 1, Table 1) The first was near Crooked
Harbour, Lai Chi Wo (26 February 1978) and the
second at Tin Sam, Tai Po (3 February 1982)
Balaenoptera edeni Anderson, 1878
Bryde's whale
[CITES Appendix I]
This whale occurs in oceanic and, occasionally, in
coastal waters in warm temperate and tropical
climates In the North Pacific, the species is
limited to waters south of 30°N (Vidal et al 1993)
Chinese researchers report Bryde's whales in
theYellow Sea, the East and South China Seas,
80

and the waters around Taiwan (Yang 1976, Wang
1984a, 1991a,b, 1993, Chou 1994)
One cetacean carcass washed up in Hong
Kong has been tentatively identified as a Bryde's
whale from photographic evidence (S
Leatherwood pers comm ) The carcass was found
in Tung Wan on Cheung Chau Island (18
September 1992) (Fig 1, Table 1) A second,
confirmed, Bryde's whale carcass was discovered
in Tolo Harbour Channel on 22 November 1994
(Plate 1)
Balaenoptera physalus (Linnaeus, 1758)
Fin whale
[CITES Appendix I]
This is a cosmopolitan, but usually pelagic,
species It occurs in cooler waters during the
summer months and warm waters in winter The
species shows less distinct migrations than some
other types of great whale
Fin whales are known to winter in the Bohai
Sea , the East and South China Seas and the
Yellow Sea (Wang 1978, 1984a, 1991a, 1993) as
well as in the Sea of Japan There are six
recognized populations worldwide The East
China Sea Fin whales are thought to be a resident
and distinct population, as are those in the Yellow
Sea (Evans 1987) These whales presumably
migrate northwards during the summer months
There has been only one reported Fin whale
stranding (Table 1) , at Ming Sang wharf, Victoria
Harbour, on 12 April 1955 (Romer 1958) The
whale was an infant, only six to seven weeks old,
and the preserved skeleton of the whale is now
mounted and resides outside the Swire Institute of
Marine Science (Morton et al 1992)
Family Eschrichtiidae
Ellerman and Morrison-Scott, 1951
Eschrichtius robustus (Lillejeborg, 1861)
Gray whale
[CITES Appendix I]
Although there have been no recent sightings,
Gray whales are believed to have frequented Hong
Kong waters in the past 'Dragon' bones in several
temples in the territory are believed to come from

Cetaceans recorded from Hong Kong

Lung Kwu Chaa

KEY
Family Balaenopteridae
Balaenoptera physalus
Cj
Balaenoptera acutorostrata %
Balaenoptera edent
O
Family Delphinidae
Delphmus capensts
±
Grampus grtseus
•#•
Pseiidorca crassitiens
T^Sousa chinensis
D
Stenella sp
V
Stenetta attenuata
AV
Stenella coeruleoalba
CV
Steno bredanensis
*
Turstops truncatus
&
Family Phocoenidae
Neophocaena phocaenotdes
•
Family Physeteridae
Kogia brevtceps
rt
...
1
...


Gray whales historically hunted by locals
...

The most abundant occurs in the Eastern Pacific
...
1985;
Brownell and Chun 1977; Wang 1984a, 1991a,
1993)
...
Hunted until 1966, the Korean
population is now estimated to number between
200-300 individuals (Evans 1987)
...

Wang (1984b) documents a Gray whale captured
from nearby Guangdong Province, which was at
that time a breeding ground for whales
...
2), to the east of Hong Kong, was once a
calving site for Gray whales
...

Suborder Odontoceti Flower, 1867
Toothed whales
Superfamily Delphinoidea (Gray, 1821)
Family Delphinidae Gray, 1821
Marine dolphins
Subfamily Delphininae (Gray, 1821)
Delphinus spp
...
Common dolphins occur
throughout the Indian Ocean and, in the Pacific,
81

E
...
Parsons, M
...
Felley & L
...
Porter

Plate 1
...
Credit:
E
...
Parsons
...

Common dolphins have been seen in the Bohai
...

1991a
...
1993; Chou 1994; Zhou et al
...

The genus Delphinus currently contains two
species: Delphinus delphis (Linnaeus
...
Delphinus capensis (Heyning and Perrin
...
Ft should be noted that for stranding and
distribution records prior to 1994, all Common
dolphins were assumed to be D
...

Although there was a highly publicised visit
of a pod of approximately 50 Common dolphins
to Victoria Harbour (20 February 1978) and one
individual was captured near Kwun Tong in 1954
(Romer 1958)
...
l
...
It occured at Shap Long
...

From photographs of the dolphin it was suggested
(S
...
Perrin pers
...
) that
the animal was a member of the newly defined
species Delphinus capensis or the putative species
Delphinus iropicalis
...
Cuvier
...
The range
extends southwards from the Kuril Islands in the
North Pacific, to New Zealand, southern Australia
and Tasmania
...
1984;
Ailing 1986), especially in the northern and
western reaches and deeper coastal waters (Kruse
et al
...
extending southwards to Australia

Table 1
...
Parsons, M
...
2
...


Plate 2
...
Credit: Agriculture and Fisheries Department, Hong Kong Government
88

Cetaceans recorded from Hong Kong
Table 2
...

Spotted, Striped and Spinner dolphins
[CITES Appendix II]
Due to the difficulty in identifying stranded
stenellids to species it is highly probable that
several
stranded
carcasses
have
been
misclassified Bearing this in mind, the complete
records of Stenella spp strandings are here treated
as a single genus rather than three separate
species
All three pantropical stenellids occupy warm
temperate and tropical waters and share much the
same distribution Stenella attenuata (Pantropical
spotted dolphin, Gray, 1846) is primarily oceanic
and is one of the most commonly occurring
species in its range It is recorded from the South
and East China Seas (Wang 1984a, 1991a, 1993,
Zhou et al 1995), Taiwan (Yang 1976, Wang
1991b, Chou 1994, Zhou et al 1995) and near
Japan In the Indian Ocean, it occurs from the
Seychelles southwards to southern Australia and
New Zealand There are several geographic
variants of the species and the coloration of IndoPacific animals is varied and complicated
(Leatherwood and Reeves 1983), making them
easily confused with other stenellid species when
not seen close up
Stenella coeruleoalba
(Meyen 1833), the
Striped dolphin, is also oceanic, but it also comes
inshore in some areas The species is found
throughout the Indian Ocean, while in the Pacific
it extends from Japan to Taiwan (Yang 1976,
Wang 1984a, 1991a,b, 1993, Chou 1994), the East
and South China Seas (Zhou et al 1995) and
southwards to Australia's southern tip
The third stenellid species recorded from
Hong Kong waters is Stenella longirostris (Gray,
1828) This species is usually oceanic but can be
coastal Its range in the Indian Ocean extends from
the northernmost reaches to southern Australia and
South Africa Both China and Taiwan have

Cetaceans recorded from Hong Kong

records of the species ( Wang 1991a, b
...
1995)
...
illegally
caught Spinner dolphin carcasses were intercepted
by the Hong Kong Government
...
China, adding this
locale to their range
...
One was found on the south
side of Hong Kong Island, al Shek 0 ( 1 5 February
1985) and the second was at Tai Long Wan on the
south side of Lantau Island (13 March 1985)
(Fig
...
A carcass reported on 14
December 1987 at Po Chue Tam
...
Also on Lantau Island, the
first of the four known Stenella coeruleoalba
strandings was washed up at Pui O on 10 May
1988 (Plate 3)
...
Hoi Ha Wan (27 June 1992)
and Shek O (17 July 1994)
...

live animals have been sighted (L
...
Porter pers
...

Tursiops truncatus (Montagu
...
In the western
Pacific, the distribution extends from Japan to
Australia and New Zealand
...

They also occur in waters adjacent to Hong Kong
...

1991a,b
...
1995;
Zhou et al
...
At present, all bottlenose
dolphins are conservatively assigned to Tursiops
truncatus
...
A Striped dolphin (Stenella coeruleoalba) discovered on 10 May 1988 at Pui O, Lantau Island
...
CM
...
L Felley & L
...
Porter

different 'ecotypes' in many areas, including the
western Pacific (Walker 1981; Scott and Chivers
1990)
...
longer-snouted, and spotted coastal forms
(often refered to as Tursiops aduncus) and
heavier-bodied, larger-snouted and less spotted
offshore forms (often refered to as Tursiops gilli)
...
l)
...

Silvermine Bay (Lantau Island)
...
Deep Bay (26 March 1985)
...
Sai Kung on
8 August 1986
...
l
...
and
nearby Cape D'Aguilar (23 April 1994)
...
obs
...

One was an unconfirmed carcass in Tai Long
Wan
...
Clarke
pers
...
The other three carcasses were
found in Siu Sai Wan (25 November 1994) and
near Stonecutters Island (4 December 1994) and
Tolo Harbour (11 December 1994)(Fig
...
The
Siu Sai Wan and Stonecutters Island carcasses
were measured and necropsied and from
preliminary examination were deemed to be
members of an offshore ecotype (Leatherwood
pers
...
) (Plate 4)
...
A Bottlenose dolphin (Tursiops truncatus) reported on 25 November 1994 at Siu Sai Wan, Hong
Kong Island
...
Leatherwood
9?

Cetaceans recorded from Hong Kong

Subfamily Globicephalinae (Gray, 1866)
Pseudorca crassidens (Owen, 1846)
False killer whale
[CITES Appendix II]
This is a widely distributed species (Leatherwood
and Reeves 1983) which is found in offshore
warm temperate and tropical waters, including the
Indian Ocean (Leatherwood et al
...
The species' range extends from Japan
and the Red Sea south to New Zealand and South
Africa
...
They are
known to occur around Taiwan (Yang 1976; Wang
1991b; Chou 1994) Wang (1984a, 1991a, 1993)
and Zhou et al
...

There has been only one carcass found in Hong
Kong, in Tai Lam Chung on 19 April 1983
...
The first is Sousa
teuszii, the Atlantic Humpbacked dolphin
(Kukenthal 1892), occurring coastally in tropical
west Africa
...

The classification of the latter is confused and
disputed
...
Pilleri
and Gihr (1974) suggested that Sousa chinensis
should be split into several species: the freckled
Sousa lentiginosa (Owen, 1866), and Sousa
plumbea (Cuvier, 1829), both extending in range
from East Africa to Thailand; Sousa borneensis
(Lydekker, 1901), dwelling in the coastal waters
of Borneo and Australia; and Sousa chinensis in
Chinese waters
...
lentiginosa
was not valid, but that S
...
(in
press) argue, a sub-species
...
This
leaves the species Sousa chinensis (or sub-species
S
...
chinensis; Ross et al
...

The species usually occurs in warm, shallow
(less than 20 m deep) waters
...

Chinese
researchers
have
documented
Humpbacked dolphins in the East and South China
Seas (Wang 1982, 1984a, 1991a, 1993; Zhou et
al
...
They have also been known to
swim several kilometres up rivers, including the
Minjiang and Jiulong rivers in Fujian Province as
well as the Pearl (Wang 1982) and Shenzen rivers
(D
...
comm
...
They are reported to be abundant in
Xiamen, especially between February and May
(Z
...
Huang pers
...
; Wang 1984a)
...
Known locally as Chinese white dolphins,
the animals notable for their coloration
...
They lighten with age,
leading to white or pink adults which often have
either grey or blue mottling and spots
...
Ross pers
...
Their mating season appears to be from
August to September (E
...
Parsons and L
...

Porter pers
...
), which is later than the May to
June breeding season noted by Wang (1982) for
the Xiamen population
...
l)
and numerous sightings of live dolphins
...
To date, they
can consistently be found near Castle Peak (Tuen
Mun) and the islands of Lung Kwu Chau and Sha
Chau
...
CM
...
L
...
J
...
Regular sightings of identified individuals have confirmed that Sousa chinensis is resident
in Hong Kong
...
J
...
In
September 1994
...

Mui Wo and Peng Chau
...
Melville (1976) also records them off Pak
Nai
...

These sightings were, however, when the size of
the population was considerably larger and
possibly more widespread than it is at present
...
Cuvier, in Lesson
...
It is known to occur from northern

Japan to Australia in the Pacific Ocean and from
the Gulf of Bengal and the Gulf of Aden
southwards to South Africa and Western Australia
...
Rough-toothed dolphin distribution
includes both Chinese and Taiwanese waters
(Yang 1976: Wang 1984a
...
1993; Chou
1994; Zhou et al
...
A solitary carcass washed
aground on East Sharp Island, Port Shelter on 7
March 1982 (Fig
...

Family Phocoenidae (Gray
...
Cuvier
...
Although on

Plate 7
...
This stranding was reported on 13 December 1992 at Shek O, Hong Kong
Island
...
A Pygmy sperm whale carcass reported on 3 October 1992 at Marina Cove, Sai
Kung
...
Agriculture and Fisheries Department, Hong Kong Government

References
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...
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Andrews, R
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...
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...

Barnes
...
G
...
P
...
E
...

Status of studies on fossil marine mammals
...

Blokhin, S
...
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...
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...
M
...
On the Korean-Okhotsk population of
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...

Brownell
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L
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Asian Marine Biology 12 (1995) 101-109

ODYSSEYLANA SIRENKOI: A NEW GENUS AND NEW SPECIES OF
CIROLANID ISOPOD FROM THE SOUTH CHINA SEA (CRUSTACEA:
ISOPODA)
Marina V
...
Name is formed from the name of
the research vessel Odyssey, from which the
specimen was collected and the ending -lana to
indicate family affinity The gender is feminine
Remarks
Odysseylana
belongs to the
Comlera
group on the basis of general
morphology, body shape, antennae, pereopods,
elongate frontal lamina and pleopod 1 peduncle
The triangular pleotelson and uropod endopod
shape of Odysseylana
are similar to those of
Comlera (Montagu, 1804) Comlera differs in
having an indurate, operculate first pair of
pleopods, almost completely encapsulating
102

ventrum, flat, short uropod exopod (Fig 6 C, D),
more slender frontal lamina (about five times
longer than basal width) with a semicircular
anterior projection and pereopods 1-3 merus
distodorsally more prolonged Quadrate pleopod
1 peduncle, arrangement of pleopods, when the
first ones do not cover the following, slender, long
uropod exopod share this genus with Politolana
Bruce, 1981 Politolana differs from Odysseylana
in having frontal lamina more slender and rounded
anteriorly, ischium and merus of pereopods 1-3
distodorsally more strongly prolonged, appendix
mascuhna arising sub-basally or sub-medially,
endopod of uropod with a notch on the lateral
margin, pleotelson with lateral margins convex
Comlera and Politolana have no species with the
frontomedial margin of head extended and
covered basis of antenna 1 In both Comlera and
Politolana
antenna 2 has peduncle article 1-2
short, 3-5 long, article 5 is longest and differs
from antenna 2 of Odysseylana
with peduncle
articles 1-3 short, 4 and 5 long, with article 4
longest In Odysseylana pleonites in ventral view
are broader than pleopod 1 peduncle, whereas
Comlera and Politolana (and of most cirolanid
genera) have the ventral part of the pleonites
markedly narrower than pleopod 1 peduncle
Odysseylana is separated from the all other genera
of the Comlera group by the deep rounded notch
in the very narrow distal end of the pleotelson and
setation of its dorsal surface, conical uropod
exopod, 3 coupling hooks on maxilliped endite
Only one cirolanid genus has non-flat (cylindrical)
uropod exopod, l e , Atarbolana Bruce and Javed,
1987, but Atarbolana is otherwise very different
from Odysseylana
Odysseylana sirenkoi
(Figs 1-6)

sp nov

Material examined
Holotype N 1/69411, male,
10 45 mm long, South Vietnam, off Tkhu Island,
10 4 6 ' N 109 4 3 ' E , depth 300m 25 August 1984,
RV Odyssey, coll B I Sirenko
Description

of male

holotype

Body (Fig 1) Total body length 10 45 mm,
m a x i m u m width 2 70 mm, length 3 83 width

Odysseylana sirenkoi

Fig
...


Odysseylana
sirenkoi sp
...
Male,
holotype
...


Body elongate, dorsally vaulted, straight-side
...
8 times as wide as medial length;
frontomedial margin extended, covering basis of
antenna 1
...
Pereon outline smooth,
pereonite 6 longest, 2 and 3 shortest, remainder
subequal in length
...
Epimera of
pleonite 4 longer than preceding plate, posteriorly
extended, rounded and laterally bent
...
26 length, pleotelson
length 0
...
6
posterior margin width; posterior notch with 4
spines and plumose marginal setae; lateral margins
concave, denticulate, with plumose marginal setae
and without spines; dorsal surface with two broad,
dense rows of stout plumose setae
...
2 A, B) not visible in dorsal view, 2
...
Clypeus triangular, 4
...

Antenna 1 (Fig
...
Not extending to end of
antenna 2 peduncle
...
08 body length
...
Peduncle article 1 and
2 are subequal in length and fused, with two
distomedial setae, article 3 narrower but longer
than preceding ones, article 3 length 0
...
Ten flagellar articles with a
row of long and slender aesthetascs and a group
of short simple setae
...
2D)
...
Length 0
...

Mandibles (Fig
...
Spine row with 9 or 10 stout
spines, molar process long, triangular, with an
anterolateral row of small strong acute teeth
...
86 mandibular body length
...

Maxilla 1 (Fig
...
Outer endite with 13
spine-like setae, inner endite 0
...

Maxilla 2 (Fig
...
Innermost endite shortest
and broadest, with numerous simple and 3
circumplumose setae, medial endite with 14 and
outer endite with 5 spine-like setae
...
2
...


Odysseylana sirenkoi sp
...
Male, holotype
...


105

M V Malyutma

Fig
...


106

Odysseylana sirenkoi sp nov Male, holotype
...
5
...
nov
...
A-E, pleopods 1-5; F, penes on sternite 7
...
6
...
This species is known only from the
single specimen It is distinguished from other
Cirolanidae by the generic characters
Etymology
...
Y
...
-S Chou2, C
...
S
...
-H Chou4
1

2
?
4

Department of Biology, McMaster University, 1280 Main Street West, Hamilton, Ontario L8S 4K1
Canada
Department of Zoology, National Taiwan University Taipei, Taiwan, ROC
Department of Zoology, University of Guelph, Guelph, Ontario, N1G2W1, Canada
National Museum of Natural Science, Taichung, Taiwan 40419, ROC

Abstract
Recent strandings and previous records of Cuvier's beaked whale, Ziphius cavirostris, in Taiwan are
described The chronology and distribution of the events and the external morphology and appearance,
stomach anatomy and contents, and reproductive characteristics of specimens are presented

Introduction
Cuvier's beaked whale or the Goose-beaked whale
(Ziphius cavirostris Cuvier, 1823) is probably the
most cosmopolitan member of the Ziphndae
However, a global review revealed no published
accounts from the waters surrounding Taiwan,
even though there are numerous records from the
waters of Japan and parts of southeast Asia and
Australia (Heyning 1989) The paucity of
information from Taiwan more likely reflects
limited research effort or poor documentation of
specimens rather than a lack of animals We
present information on nine strandings in Taiwan
between late 1991 and late 1994, and mention
three specimens collected prior to 1990
Seasonality of occurrence, reproduction and diet
are discussed

Chronology and location of stranding events
On 26 November 1991, a live 575 cm male was
found on a beach at Houlung, Miaoh County,
northwest Taiwan Newspaper accounts stated that
the whale returned to the sea but stranded again
an hour later and died The eyes and both erupted

teeth were removed by local residents Five scars
or wounds on the body were reported, they may
have been inflicted by parasites (see later) The
complete skeleton (except the teeth) is at the
National Museum of Natural Science (NMNS-10)
Two animals stranded alive on a beach in
Huahen County, northeast Taiwan on 3 February
1993 Positive identification was made by the
authors from
high quality,
newspaper,
photographs Sexes are uncertain, but light
pigmentation around the head of one suggests that
it was male (see Heyning 1989)
After
unsuccessful rescue attempts with heavy
machinery, both animals died and were butchered
for sale by local residents The masses were
estimated by the residents at 3000 kg (the probable
male) and 2000 kg No samples were obtained
from either
Two more animals stranded alive on the same
beach early in the morning on 24 January 1994
The larger (approximately 7 m long), still partially
immersed, was pushed back to sea successfully by
a bulldozer It was probably a male judging from
rescuers' descriptions of visible teeth at the tip of
the lower jaws The smaller whale was stranded
higher on the beach and died after unsuccessful
attempts to return it to the sea (Plate 1 A) She was

J
...
Wang, L
...
Chou, C
...
Yao, A
...
Neimanis & W
...
Chou
120°

121°

122°

TAIWAN
STRAIT

25°

24°

PACIFIC
OCEAN

23c

O Lut&o (Green Island)

Lanyu (Orchid Island)

22°

21°
0

20 40
kilometres

Fig
...


Map of the study area with locations and
depth contours
...
This represents only the tenth
Cuvier's beaked whale for which mass has been
determined
...

Two weeks later, on 11 February 1994, a
112

whale stranded on Lutao (Green Island), Taitung
County, southeast Taiwan
...
Although length
and sex were not reported, light pigmentation
around the head and evidence from photographs
that teeth may have been removed from the tip of
the mandibles suggest it was a male
...
When we arrived the following day, she was
floating and her body was curled with the flukes
submerged to almost a metre depth making an
accurate measurement of the total length
impossible
...
5 cm long male
foetus (Plate IB)
...
The female's skull was
retrieved and is being prepared at the National
Museum of Natural Science (LY-94-01)
...
No other data are
available
...
She
was necropsied at the National Museum of Natural
Science, and the skeleton is being prepared for the
collection (TD-94-01)
...

An approximately 600 cm male stranded on a
beach at Lukang, Changhwa County, central
western Taiwan, on 19 October 1961
...
Yang also mentioned
that a specimen was captured near Hengtseng
(location details were not mentioned) but provided
no other information
...
Length and

Cuvier's beaked whales from Taiwan

Plate 1
...
A, HU-94-01;
B, foetus of LY-94-01
...
Some characteristics of Ziphius cavirostris stranded along the beaches of Taiwan
...


percentage of the total body length) of some
specimens are presented in Table 1
...
Four were greater
than two standard deviations
...
g
...

Stomach contents
The stomach anatomy of the Taiwan specimens
was consistent with the description by Mead
(1993)
...
Comparisons of morphometric measurements as a percentage of total length between
the Cuvier's beaked whales of Taiwan and reported means
...
Summary of the anatomy and contents of the stomachs of Cuvier's beaked whale from Taiwan
waters
...
Jefferson2
1
7

Department of Biology, Nanjing Normal University, Nanjing 210097, People's Republic of China
Ocean Park Conservation Foundation, Ocean Park, Aberdeen, Hong Kong

Abstract
There are confirmed records of 22 species of small cetaceans in Chinese waters (including Hong Kong,
Macau, and Taiwan) The list includes a few cold temperate species and nearly all of those warm
temperate and tropical species known from the Indo-Pacific region Three species previously reported
(Irrawaddy dolphin, Short-beaked common dolphin, and Harbour porpoise) are herein removed from
the list, based on a critical review of information supporting the identifications One or more of these
three, as well as additional species of beaked whales, may ultimately be shown to occur in Chinese
waters In China, small cetaceans of many species are suffering from problems associated with habitat
loss/degradation and kills in fisheries Effective conservation efforts are needed to avoid the loss of
stocks and even species

Introduction
China1 is bordered by the Bohai, Yellow, East
China and South China seas, adjacent to the
western North Pacific (Fig 1) China's Extended
Economic Zone is large, and it has a coastline of
over 18,000 km, not including that of the more
than 5,000 islands on the continental shelf and
slope and in oceanic waters Chinese fisheries rank
among the most productive anywhere, with
worldwide annual marine catches of > 10 million
tonnes Extensive fisheries are known to operate
in most coastal provinces and interactions between
fishery operations and cetaceans are known to
occur Figure 2 is a map of Taiwan
The extensive coastline of China is inhabited
by a large number of species of small cetaceans
Other than work on the population status and
biology of the Baiji (Lipotes vexilhfer Miller) and,
to a lesser extent, the Finless porpoise

(Neophocaena phocaenoides [Cuvier]), there has
been little directed work on the small cetaceans
of China In the past few years, however, many
research projects on marine cetaceans in mainland
China, Taiwan and Hong Kong have been initiated
and one for Macau is being planned (Perrin in
press) Parsons et al (1995) recently provided a
checklist of cetaceans in Hong Kong Several
species of China's marine dolphins and both its
species of freshwater small cetaceans are known
to be caught incidentally in fisheries throughout
the region (Zhou and Wang 1994, Chou et al
1995) However, very little is known about the
impacts of human activities on the small cetaceans
of China This paper reviews published
information on the occurrence of each of the
species recorded, presents some unpublished data,
and discusses the current conservation status of
small cetaceans in China

In this paper, we use the term China to refer to the geographical region including mainland China,
Taiwan, Hong Kong and Macau

Z h o u , K
...
L e a t h e r w o o d & T
...
J e f f e r s o n

110°E

130°E

120°E

/

rr
V

;"\
...
s

Panshan
Xincheng

•1
...
cW / C o u n t y ,

BOHAI SEA
—
Shandong
YELLOW SEA
Ganyu

<^

Guanne River
Jiangsu

Rugao County

Rudong

— Lusi

Zhenjiang City
...


120

1
...


Small cetaceans of China

Taoyur>4

-^Keelung

® -

Taipei

!

£3

Peikang »

VDashl

TAIWAN

Kaohsiung® ®Pmgtung

For most offshore species, we have only
scattered stranding records, which are not
indicative of their true ranges; we have therefore
not provided distribution maps for these species
...
Studies of stock structure are
badly needed for most species of cetaceans that
occur in the region
...
2
...


Known species and their distributions
Examination of the published and unpublished
literature indicates that 25 species of small
cetaceans have been reported from Chinese
waters
...
Of these, 13 have been recorded on both
sides of the Taiwan Strait, four from mainland
provinces and territories only, and five from
Taiwan only
...
However, all three are found along
adjacent sections of coastline (Short-beaked
common dolphins and Harbour porpoises to the
north (Perrin in press), and Irrawaddy dolphins to
the southwest (Lloze 1973) and may eventually be
found to be a part of the cetacean fauna of China
...
1995; Chen et al
...
There are also at least three stranding
records from Hong Kong (Parsons et al
...
It
is possible that some of these reported specimens
of K
...
simus
...

A female was found dead on the beach of Miaoli
County, Taiwan, in June 1986 (Chou 1989)
...
1991)
...
1995)
...
The skull of this specimen is now

The so-called Suao fish market is actually located in the small village of Nan Fang Au, just south of
Suao
...

121

Zhou K S Leatherwood & T A Jefferson
Table 1
...

Region
Species
Kogia breviceps

SCS

ECS

X

Ziphius cavirostris
M

densirostns

HK

X

X

X

X

X

X

X

X

T

X

Kogia simus

YS

BS

YR

X

Mesoplodon
ginkgodens

X

X

X

0rein us orca

X

X

X

Globicephala
macrorhynchus

X

Pseudorca crassidens

X

X

X

Feresa attenuata

X

X

X

Peponocephala electra

X

X

X

Sousa chinensis

X

X

Steno bredanensis

X

X

M

Lagenorhynchus
obliquidens

X
X

X

X

X

X
X

X

X

X

X

X

X
X

X

Grampus griseus

X

X

Tursiops truncatus

X

X

Stenella attenuata

X

X

X

S

longirostns

X

X

X

S

coeruleoalba

X

X

X

X

Delphinus capensis

X

X

X

X

Lagenodelphis hosei

X

X

X

X

Neophocaena
phocaenoides

X

X

X

X

X

X

Lipotes vexillifer

X
X

Region abbreviations SCS, South China Sea, ECS, East China Sea, YS, Yellow Sea, BS, Bohai Sea,
YR, Yangtze River, T, Taiwan territorial waters, HK, Hong Kong territorial waters and M, Macau territorial
waters
housed at the Tawian Fisheries Research Institute,
Kaohsiung (TFRIK) (Jefferson pers observ )
Several specimens were also obtained from Suao
and Hengchun, Taiwan (Yang 1976, Wang, P
1991) There are records of six recent strandings
from Taiwan, at Miaoh, Huahen, Lu Tao, Lan Yu,
and Taitung (Chou et al 1995, Chen et al 1995)
122

Baird's beaked whale: Berardius bairdu
Stejneger
A complete skeleton of a Baird's beaked whale is
housed in the Zhejiang Museum of Natural
History (Wang Yu pers comm ) A museum staff
member said it was collected by the Zhoushan
Marine Fisheries Company at the end of the

Small cetaceans of China

r5

i 2

Plate 1
...


1950s, but the collection locality is not known,
and it may not have come from China
...

Hong Kong
...
It may
...

waters
...
Local fisheries around the
Penghu Islands and Xiaoliuqiuyu Island
...
Another specimen was found at the
fish market in Kaohsiung
...
There is also a single record of a whale of
this species having been taken by commercial
whalers in southern Taiwan (Yu 1995)
...
A
specimen was found stranded at Changxin Island
...
data)
...
Taiwan, in 1968 (Kasuya and
Nishiwaki 1971; Yang 1976)
...
Liaoning Province, in August 1980 (Shi

Although there are confirmed records thus far of
only two species of Mesoplodon from Chinese
waters, it is likely that other species occur there
...
22), identified by
the author as a Blainville's beaked whale, show
what is probably another species of Mesoplodon
...
carlhubbsi
Moore) or a Stejneger's beaked whale (M
...

It should be noted that several sightings and
possible captures of Bottlenose whales
{Hyperoodon sp
...
1992; Perrin in
press; R
...
Pitman pers
...
), and it is feasible
that Bottlenose whales could occur in China
...
, S
...
A
...
3
...


126

Small cetaceans of China

Plate 2
...


Taiwanese records have been reported, from
Taipei
...
1995;
Chen el al
...
At the NTOU
...
Several small herds of
Risso's dolphins were observed off Hualien,
Taiwan, on 18 and 19 June 1995 (Jefferson
unpubl
...

Bottlenose dolphin: Tursiops truncatus
(Montagu)
Bottlenose dolphins are widely distributed in
Chinese waters (Fig
...
A larger form (truncatus-

or gilli-lype — Plate 2B) and a smaller form
[aduncus-lype — Plate 2C) are recognized (Zhou
1987; Wang et al
...
The former occurs in the
Yellow
...
and East China seas
...
Bottlenose dolphins have
been recorded from Liaoning
...

Zhejiang
...
1981; Shi and Wang 1983: Wang 1990; Zhou
1965, 1987; Zhou et al
...
1995)
...

observ
...

1976)
...

1994)
...
1995)
...
Taiwan (Yang 1976)
...
observ
...

Specimens were caught about 300 km southeast
of Shantou in 1978 (Zhou et al
...
Guangxi Zhuang
...

A dolphin carcass from Keelung was reported by
Yang (1976) to have been a specimen of Stenella
frontalis, but the published photos show that it is
was a Pantropical spotted dolphin
...

apparently of this species, was found in December
1987, on Lantau Island, Hong Kong (Parsons el
al
...
Spotted dolphins are also known from
Taiwanese waters, based on several recent
strandings (Chen et al
...
1995)
127

Zhou, K
...
Leatherwood & T
...
Jefferson

120°E

110"E

130°E

V
"\
...
4
...


128

Small cetaceans of China

and from captures by fishermen (Mizue 1988)
...
northeast Taiwan, on 19 June
1995 (Jefferson unpubl
...
This appears to be
one of the most common species of oceanic small
cetacean off the east coast of Taiwan
...

Guangxi Zhuang
...
Illegally caught Spinner dolphin
carcasses from Hainan Island were intercepted in
Hong Kong in 1993 (Parsons et al
...
For
Taiwan, there are stranding records of Spinner
dolphins from Kaohsiung (Yang 1976)
...
1995), and an unknown locality (Chen et al
...
Three skulls in the collection of the NTOU
were collected at the Suao fish market
...
Dolphins of this species were
frequently encountered just a few kilometres off
the coast of Hualien
...

data)
...
S
...


fc

•

- 4 v * > JT

*•

Striped dolphin: Stenella coeruleoalba (Meyen)
A male Striped dolphin was found at the fish
market in Suao
...

The collection of the NTOU in Taiwan also
contains the skull of a Striped dolphin collected
at the Suao fish market (Mizue 1988)
...
1989
...
1995)
...
capensis Gray —
Plate 3C) (Heyning and Perrin 1994)
...

Two specimens were captured from the northern
Yellow Sea in 1963 and 1978 and are housed in
the Dalian Natural History Museum (Shi and
Wang 1983)
...
1980)
...
A, Pantropical spotted dolphins observed off Hualien, Taiwan, on 19 June 1995, B, part of a
herd of Spinner dolphins off Hualien, Taiwan, on 18 June 1995, C, captive Long-beaked
common dolphin captured off the Penghu Islands, Taiwan; D
...
A, Finless porpoises captured in gillnets off mainland China; B, captive Baiji 'Qi Qi' at the Wuhan
Institute of Hydrobiology; C, free-ranging Baiji in the Yangtze River
...
observ
...
1995; Chen el al
...
Finless
porpoises, which are shallow water animals,
probably do not occur off the east coast of Taiwan
...

Baiji: Lipotes vexillifer Miller
Baiji (Plate 4B
...
6)
...
In the 1970s and 1980s, however, the upper
limit was no farther upstream than Zhicheng,
Hubei (Chen and Hua 1989; Zhou et al
...

Results of recent surveys of almost all the species'
previous range, Shanghai to Yichang, suggest that
131

Zhou, K
...
Leatherwood & T
...
Jefferson

Fig
...
Approximate distribution of the three described subspecies of the finless porpoise in Chinese
waters
...
p
...
p
...
p
...
p
...
p
...
, S
...
A
...
6
...


134

130°E

Small cetaceans of China

1989) Perrin and Brownell (1989) reported that,
in the past, some dolphins stranded on sandbars
were beaten to death by local residents No
directed exploitation is known, although capture
teams from the Institute of Hydrobiology and
Tonghng Seminatural Reserve are actively
searching for Baiji to live capture for captive
breeding programmes (Liu et al in prep )
There are no reliable population estimates for
Baiji and the best that can be surmised from the
various reports of surveys is that rates of
encounter and, presumably, also numbers, have
declined precipitously since the late 1970s, and
that the population is 'small and presumably still
declining' (Zhou et al 1994) The first rigorously
designed and conducted surveys covered most of
the river between Shanghai and Ytchang in the last
quarter of 1994 They used three vessels, a ship
in mid-channel and a ship along each shore, and
only five animals were seen This suggested to the
participants that the population numbered fewer
than 100 individuals, perhaps no more than a few
dozen (Liu et al in prep )
Live captures and removals of small cetaceans
must be included in assessments of the effects of
exploitation To date, as far as we know, only a
few Baiji, Finless porpoises, and Bottlenose
dolphins have been live-captured in mainland
China waters for either display or research In
Taiwan, although there has been no live-capture
fishery, Bottlenose dolphins and False killer
whales have been purchased for display in
association with the former drive fishery at the
Penghu Islands The number of species that have
been affected by live-removals is thus far
relatively small There is, however, the potential
for greatly increased future impacts, as we are
aware of at least nine dolphinariums under
construction in mainland China (one each in
Qmgdao, Shanghai, Guangzhou, Shenzen, Wuxi,
Xiamen, and Dalian and two in Beijing) and two
additional ones in Taiwan (Taichung and
Keelung)
The Chinese Government at various levels has

enacted a series of laws and provisions to protect
wildlife including small cetaceans (Zhou 1992)
The capture, killing, selling, or buying of the
national key protected aquatic animals, including
Baiji, is strictly prohibited Most ot the provinces
along the mainland coast and in the Yangtze River
have issued either their own laws or regulations
for the implementation of the national laws and
regulations In Taiwan, there is now legal
protection for all cetaceans But, in China, as
elsewhere, laws have little or no effect if people
do not know about them and if they are not
enforced humanely and consistently
The wide variety of cetaceans that occur along
the various coastlines and rivers of mainland
China, Taiwan, Hong Kong, and Macau represent
a valuable natural treasure They can only be
conserved for future generations with education
and laws suited to the needs of humans and
wildlife alike We hope that public officials and
citizens will take note of this fact and push for
strict and effective measure to conserve all
cetacean species in Chinese waters

Acknowledgements
Thanks to X Xu and G Yang for assistance in
Zhou's field work We appreciate the efforts ot G
Chen, R Chen, L Chou, J Y Wang, and H Yang
for their hospitality when Jefferson was in Taiwan
and E Huang for assistance in locating records
and for translation of several papers L Porter and
R Chen provided photos for the plates Support
for some of the work described in this paper came
from the World Nature Association, National
Taiwan Ocean University, National Taiwan
University, Hong Kong's Airport Authority,
Ocean Park Conservation Foundation, and the
IUCN/SSC Cetacean Specialist Group We thank
C Leatherwood for typing an earlier draft, B
Wang and C Mok for preparing the figures, and
Professor Brian Morton and an anonymous
reviewer for comments on the manuscript

135

Zhou K S Leatherwood & T A Jefferson

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Small cetaceans of China
Appendix I
...

Spelling used in Taiwan

Spelling used in Mainland China

Kaohsiung

Gaoxiong

Tungkang

Donggang

Keelung/Chilung

Jilong

Dashi

Daxi

Lukang

Lugang

Pingtung

Pmgdong

Peikang

Beigang

Taipei

Taibei

Hsmchu

Xmzhu

Hualien

Huahan

Ilan

Yilan

Taichung

Taizhong

Taitung

Taidong

Taoyun

Taoyuan

Yehliu

Yehu

Lu Tao

Ludao Island

Chmmen

Jinmen

1



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