Developments in Hydrobiology 98: Series Editor

Developments in Hydrobiology 98
Series editor
H. J. Dumont
Ecology and Conservation of Southeast Asian Marine
and Freshwater Environments including Wetlands
Ecology and Conservation of
Southeast Asian Marine and Freshwater
Environments including Wetlands
Edited by
A. Sasekumar, N. Marshall & D. J. Macintosh
Reprinted from Hydrobiologia, vol. 285 (1994)
Springer Science+Business Media, B.V.

Library of Congress Cataloging-in-Publicatio
n Data
A C.I.P. Catalogue record for this book is available from the Library of Congress.
ISBN 978-94-010-4414-1 ISBN 978-94-011-0958-1 (eBook)

DOI 10.1007/978-94-011-0958-1
The publication of this volume was funded by World Wide Fund for Nature Malaysia, Canada-
ASEAN Centre and Asian Wetland Bureau, and sponsored by AIDAB (Australia), UNESCO ,
IDRC, Ministry of Science (Malaysia), Petronas, ESSO
, University of Malaya and the Malaysian
Society of Marine Sciences.
Printed on acid-free paper
All Rights Reserved

© 1994 Springer Science+Busines s Media Dordrecht
Originally published by Kluwer Academic Publishers in 1994
No part of the material protected by this copyright notice may be reproduced or
utilized in any form or by any means, electronic or mechanical
including photocopying, recording or by any information storage and
retrieval system, without written permission from the copyright owner.
v
Contents
Preface vii
List of participants ix
ECOLOGY OF MARINE AND FRESHWATERS INCLUDING WETLANDS

Monitoring of sea surface temperature in the South China Sea
by KL. Tuen 1
Neap-spring tidal effects on dissolved oxygen in two Malaysian estuaries
by B.W. Nelson, A. Sasekumar & Z.Z. Ibrahim 7
The role of bacteria in nutrient recycling in tropical mangrove and other coastal benthic ecosys-
tems
by D.M. Alongi 19
Studies on the production of useful chemicals, especially fatty acids in the marine diatom
Nitzschia conspicua Grunow
by W.-L. Chu, S.-M. Phang & S.-H. Goh 33
Zoogeography and biodiversity of the freshwater fishes of Southeast Asia
by M. Zakaria-Ismail 41
The damselfishes (pisces: Osteichthyes: Pomacentridae) of Peninsular Malaysia and Singapore:
systematics, ecology and conservation
by T.M. Sin, M.M. Teo, P.KL. Ng, L.M. Chou & H.W. Khoo 49
The systematics and ecology of snakeheads (Pisces: Channidae) in Peninsular Malaysia and
Singapore
by P.G. Lee & P.KL. Ng 59
An annotated checklist of mangrove brachyuran crabs from Malaysia and Singapore
by C.G.S. Tan & P.KL. Ng 75
The ecology and biology of Southeast Asian false spider crabs (Crustacea: Decapoda: Brachyura:
Hymenosomatidae)
by C.T.N. Chuang & P.KL. Ng 85
Male courtship cycles in three species of tropical Ilyoplax crabs (Decapoda, Brachyura,
Ocypodidae)
by T. Kosuge, S. Poovachiranon & M. Murai 93
Distribution and biodiversity of Singapore gorgonians (sub-class Octocorallia) - a preliminary
survey
by N.KC. Goh & L.M. Chou 101
Distribution and abundance of marine wood borers on the west coast of Peninsular Malaysia
by H.R. Singh & A. Sasekumar 111
New records of Malaysian marine algae
by S.-M. Phang 123
The community structure of macroalgae in a low shore mangrove forest in Selangor, Malaysia
by S. Aikanathan & A. Sasekumar 131
vi
CONSERVATION AND POLLUTION

Marine environmental issues of Southeast Asia: state and development
by L.M. Chou 139
A review of otters (Carnivora: Mustelidae: Lutrinae) in Malaysia and Singapore
by N. Sivasothi & B.H.M. Nor 151
Hatch rates of green turtle eggs in Sarawak
by C.M.U. Leh 171
The use of artificial reefs in enhancing fish communities in Singapore
by C.Y.Y. Chua & L.M. Chou 177
Threats to the indigenous freshwater fishes of Sri Lanka and remarks on their conservation
by R. Pethiyagoda 189
Diversity and conservation of blackwater fishes in Peninsular Malaysia, particularly in the North
Selangor peat swamp forest
by P.K.L. Ng, J.B. Tay & K.K.P. Lim 203
Red tide phenomena in Brunei Darussalam - some implications for fisheries
by S. Subramaniam, S.M. Mahali & S.H.M. Taha 219
Water quality of Inanam River estuary and the Ko-Ne1ayan tiger prawn aquaculture ponds in
Sabah, Malaysia
by M. Mokhtar, A. Awaluddin & L.Y. Guan 227
Prolonged inundation and ecological changes in an Avicennia mangrove: implications for
conservation and management
by S.C. Choy & W.E. Booth 237
The remnant mangroves of Sei Kecil, Simpang Hilir, West Kalimantan, Indonesia
by R. Abdulhadi & Suhardjono 249
MANAGEMENT ISSUES, ECOLOGICAL ECONOMICS AND OTHER TOPICS

Marine living resources management in the ASEAN region: lessons learned and the integrated
management approach
by T.-E. Chua & L.R. Garces 257
Degradation of mangrove forests in South Sulawesi, Indonesia
by B. Nurkin 271
Value of mangroves in coastal protection
by M.A. Othman 277
A historical perspective of the resources and issues of Pak Phanang Bay, southern Thailand
by N. Srichai, S. Boromthanarat & B. Chaijaroenwatana 283
Management of coastal ecosystems in eastern Sumatra: the case of Berbak Wildlife Reserve,
Jambi Province
by G. Claridge 287
Mangrove conservation in relation to overall environmental considerations
by N. Marshall 303
Integrated planning and management of freshwater habitats, including wetlands
by P.R. Burbridge 311
RECOMMENDATIONS AND RESOLUTIONS 323

Hydrobiologia 285, 1994.
A Sasekumar, N. Marshall and D.J. Macintosh (eds), vii
Ecology and Conservation of Southeast Asian Marine and Freshwater Environments including Wetlands.
Preface
The regional seminar on the Ecology and Conservation of Southeast Asian Marine and Freshwater
Environments including Wetlands was organised by the Institute for Advanced Studies (University of
Malaya) and the Malaysian Society of Marine Sciences. The seminar was attended by 92 participants
from within and outside the ASEAN region (The Association of Southeast Asian Nations consists of
Indonesia. Malaysia, Philippines, Thailand, Singapore and Brunei). The Malaysian Deputy Minister of
Science, Technology and Environment Mr Peter Chin Fah Kui officiated at the opening ceremony.
During the 3 days (4 to 6 Nov. 1991) a total of 52 papers were discussed. Topics concerned the
ecology, conservation and management of Southeast Asian waters. The papers from the ASEAN region
exposed the inadequacy of training and research in ecology within the region. There appears to be little
emphasis on ecological research in Southeast Asia, and this probably stems from the inadequate teach-
ing of ecology in institutions of higher learning across Southeast Asia. Biology departments in these
institutions give too much emphasis to biology at the organismic level to the detriment of ecology or
environmental biology.
A major concern among ecologists in and outside the region is the degradation of the environment,
and the overexploitation of freshwater and marine resources. There is no indication that freshwater and
marine resources are being managed on a sustainable basis, making it apparent that ASEAN nations lack
the expertise to manage their resources on a sustainable basis. Sustainability demands that man treat
nature responsibly because the human species is only a part of nature, and other species have a right to
exist without interference. Unfortunately, economics has operated on the premise that the world's
resources are infinite or that any depletion is so far in the future that it should not concern the present
generation. This view is being discarded, and there is an attempt to integrate ecology and economics in
all development activities. The seminar initiated discussion on ecological economics, with one paper
examining the value of mangrove swamps. Ecological economics takes into account the true costs of
services and goods with regard to the environment.
The Malaysian Society of Marine Sciences is deeply concerned with developments in the marine
environment of Malaysia. One of the issues concerns the fate of marine turtles whose populations have
declined over the last decades. The number of female leatherback turtles coming ashore to lay eggs in
Rantau Abang, Terengganu has declined from a high of 1,500 in 1960 to a low of 80 in 1990. During all
these years many organisations had drawn the attention of the authorities to the problem, but action has
been very slow. Now the experts think it's too late to save the giant leatherbacks from local extinction.
There is serious concern over the fate of all species of turtles in the sea as modern methods of fishing
give them little chance of survival. There should be a total ban on the sale of all turtle eggs so that
greater numbers of their young find their way into the sea. Such a ban should be enforced in the ASEAN
region. In this environment conscious age, it is appalling to know that Japan buys turtle eggs from
Indonesia for human consumption!
Loss of wetlands whether freshwater swamps or mangrove swamps is a major problem in the ASEAN
region. Reclamation of mangrove swamps for aquaculture and agriculture seems to be a continuous
activity in the ASEAN region. Policy makers continue to ignore the ecological and economic impor-
tance of mangrove swamps as nursery and feeding places for marine life.
viii
The status of marine parks should be examined in the light of recent resort development activities on
small islands. The government has been progressive in declaring many small islands on the east coast of
peninsular Malaysia as marine parks. However, this protection does not extend to the land on the
islands. It appears the authorities have overlooked the fact that massive development of land on small
islands can change the ecology of fringing coral reefs. Excessive siltation and nutrient enrichment (from
sewage) can alter coral reefs into algal reefs which will be of no interest to tourists. Appropriate action
to rectify this anomaly in the status of marine parks is urgently required.
This proceedings also contains numerous recommendations for the promotion of ecological studies
and regional cooperation in marine, freshwater ecology and conservation. Special efforts should be
made to study common water masses like the Strait of Malacca, Gulf of Thailand and the South China
Sea. ASEAN should move in harmony with the rest of the world by promoting regional studies.
It is my pleasant duty to thank the organising committee members for their help in the organisation of
the seminar. A special thanks to Hydrobiologia and its Chief Editor Prof. Dumont (University of Ghent,
Belgium) for agreeing to publish the proceedings as a special volume in the series Developments in
Hydrobiology. Finally on behalf of the organisers, I thank the following sponsors who provided financial
assistance and support to organise the seminar. They include Worldwide Fund for Nature (Malaysia),
Australian International Development Assistance Bureau (AIDAB), Canada-ASEAN Centre, Interna-
tional Development Research Centre (IDRC) of Canada, The Ministry of Science, Technology and
Environment Malaysia, The Asian Wetland Bureau, Petronas, ESSO, and the Institute of Advanced
Studies (University of Malaya).
A. SASEKUMAR
(Chairman, Organising Committee) &
President of the Malaysian
Society of Marine Sciences
c/o Zoology Department
University of Malaya
59100 Kuala Lumpur
Malaysia
Hydrobiologia 285, 1994.
A Sasekumar, N. Marshall and D.J. Macintosh (eds), ix
Ecology and Conservation of Southeast Asian Marine and Freshwater Environments including Wetlands.
List of participants
ABDULLAH ABDUL RAHMAN, PKPS Aquaculture Sdn. D'CRUZ, R., Asian Wetland Bureau, Institute of Advanced
Bhd., 8th Floor, Plaza Perangsang Persiaran Perban- Studies, Univ. Malaya, 59100 Kuala Lumpur, Malaysia.
daran, 40675 Shah Alam, Malaysia. DAHURI, R., Fakultas Perikanan IPB, Kampus IPB
ADNAN KASRY, Jalan M.H. Thamrin, No. 96 Pekanbaru, Dermaga, 16610 Bogor, Indonesia.
28132 Indonesia. DAVIES, J., Dept. of Planning & Landscape Architecture,
AHMAD ISMAIL, Dept. of Biology, Faculty of Science, Queensland Univ. of Technology, GPO Box 2434,
Environmental Studies, Univ. Pertanian Malaysia. Brisbane, Qld 4009, Australia.
AIKANATHAN, S., WWF Malaysia, 3rd Floor, Wisma IJM DAVISON, G.W.H., WWF Malaysia, P.O. Box 911, Jalan
Annexe, Jalan Yong Shook Lin, 46200 Petaling Jaya, Sultan, 46200 Petaling Jaya, Malaysia.
Malaysia. DJAMARI, S.R., Directorate of Nature Conservation, JL.
ALINO, P., Marine Science Institute CS, Univ. of the Juanda 15 Bogor, Indonesia.
Philippines, Diliman, Quezon City 1101, Philippines. FATIMAH BT. MAJBID, Dept. of Zoology, Faculty of Life
ALONGI, D., Australian Institute of Marine Science, Sciences, Univ. Kebangsaan Malaysia, 43600 Bangi,
Townsville, MC, QLd 4810, Australia. Selangor, Malaysia.
ANTON, A., Dept. of Biology, Univ. Pertanian Malaysia, FATIMAH BT. MD. YUSOFF, Faculty of Fisheries & Marine
43400 Serdang, Selangor, Malaysia. Sciences, Univ. Pertanian Malaysia, 43400 Serdang,
ARUMUGAM, P.T., Faculty of Fisheries & Marine Science, Selangor, Malaysia.
43400 UPM Serdang, Malaysia. HAVANOND, S., Nakorn Sri Thamarat, Regional Forest
BOOTH, W., Biology Dept., Univ. of Brunei, Gadong 3186 Office, Amphoe Maung, Nakorn Sri Thamarat Province,
BSB, Brunei Darussalam. 8000 Thailand.
BURBRIDGB, P., House of Ross, Comrie, Perthshire, HO, C.Y., Institute of Advanced Studies, Univ. Malaya,
Scotland, DH6 2JS, U.K. 59100 Kuala Lumpur, Malaysia.
BURHANUDDIN NURKIN, Pusa Studi Lingkungan Hidup HUGHES, R., Asian Wetland Bureau, Institute of Advanced
(PSL) UNHAS, Jalan Perintis Kemerdekaan, Ujung Studies, Univ. Malaya, 59100 Kuala Lumpur, Malaysia.
Pandang, 90245 Indonesia. IDRIS ABDUL GHANI, Dept. of Biology, Univ. Pertanian
CHOO, P.S., Fisheries Research Institute, Glugor, Pulau Malaysia, 43400 Serdang, Selangor, Malaysia.
Pinang, Malaysia. IDRIS MOHD SAID, Asian Wetland Bureau, Institute of
CHONG, V.C., Institute of Advanced Studies, Univ. Malaya, Advanced Studies, Univ. Malaya, 59100 Kuala Lumpur,
59100 Kuala Lumpur, Malaysia. Malaysia.
CHOOSAK, R., Division of Environmental Impact ISHAK BIN ARIFFIN, WWF Malaysia, P.S. 10769, 50724
Evaluation, Office of the National Environment Board, Kuala Lumpur, Malaysia.
Rama 6 Road, Bangkok 10400, Thailand. IZUDDIN, K., Faculty of Agriculture, Sriwijaya University,
CHOU, L.M., Dept. of Zoology, National Univ. of Sin- Palembang, Indonesia.
gapore, Lower Kent Ridge Road, Singapore 0511. JINTONY, B., Fisheries Dept., 8th Floor, Menara Khidmat,
CHOY, S.C., Biology Dept., Univ. of Brunei, Gadong 3186 88628 Kota Kinabalu, Sabah, Malaysia.
BSB, Brunei Darussalam. KASIDJAH HARDJOPRANOTO, Directorate of Nature
CHU, W.L., Institute of Advanced Studies, Univ. Malaya, Conservation, JL. Juanda 15 Bogor, Indonesia.
59100 Kuala Lumpur, Malaysia. KHAIRUL OSMAN SALLBH, Dept. of Geography, Univ.
CHUA, Y.Y., CHRISTOPHBR, Dept. of Zoology, National Malaya, 59100 Kuala Lumpur, Malaysia.
Univ. of Singapore, Lower Kent Ridge Road, Singapore KOSUGB, T., c/o Dept. Biology, College of Science, Univ.
0511. of Ryukyus, Nishihara, Okinawa 903-01, Japan.
CHUANG, T.N., CHRISTIANA, Dept. of Zoology, National LB DIBN DUC, Dept. of Biology, Univ. of Hanoi, 19 Le
Univ. of Singapore, Lower Kent Ridge Road, Singapore Thanh Tong, Hanoi, Vietnam.
0511. LEB, G.P., BBTSY, Institute of Advanced Studies, Univ.
CLARIDGB, G., James & Claridge Environmental Manage- Malaya, 59100 Kuala Lumpur, Malaysia.
ment Consulting, 105 Honour Av., Chelmar, Brisbane, LBB, P.G., Dept. of Zoology, National Univ. of Singapore,
Qld 4068, Australia. Lower Kent Ridge Road, Singapore 0511.
x
LEELAVATHY RAJENDRAN, Pusat Asasi Sains, Univ. PHANG, S.M., Institute of Advanced Studies, Univ. Malaya,
Malaya, 59100 Kuala Lumpur, Malaysia. 59100 Kuala Lumpur, Malaysia.
LEH, M.U., CHARLES, Sarawak Museum, 93566 Kuching, PETHIYAGODA, R., Wildlife Heritage Trust, 95 Cotta
Sarawak, Malaysia. Road, Colombo 8, Sri Lanka.
LIM. R.P., Department of Applied Biology, University of ROSMAN ABDULLAH, Dept. of Zoology, Univ. Malaya,
Technology Sydney, P.O. Box 123, Broadway, NSW 59100 Kuala Lumpur, Malaysia.
2007, Australia. SAGATHEVAN K., Sunway College, 5, Jalan College,
LIM, L.H.• SUSAN, Institute of Advanced Studies, Univ. Bandar Sunway, 46150 Petaling Jaya, Malaysia.
Malaya, 59100 Kuala Lumpur, Malaysia. SALAHUDIN YAAKOB, WWF Malaysia, 3rd Floor, Wisma
LOPEZ, N.C., Institute of Biology, College of Science, UM Annexe, Jalan Yong Shook Lin, 46200 Petaling
Univ. of the Philippines, Diliman, Quezon City 1101, Jaya, Malaysia.
Philippines. SASEKUMAR, A., Dept. of Zoology, Univ. Malaya, 59100
MARIANA AHMAD, Dept. of Zoology, Univ. Malaya, Kuala Lumpur, Malaysia.
59100 Kuala Lumpur, Malaysia. SIN, T.M., Dept. of Zoology, National Univ. of Singapore,
MARSHALL, N., Adjunct Professor, Horn Point Environ- Lower Kent Ridge Road, Singapore 0511.
mental Labs, Cambridge, MD 21613, U.S.A. SINGH, H.R., Dept. of Zoology, Univ. Malaya, 59100 Kuala
MASHOR MANSOR, School of Biological Sciences, Univ. Lumpur, Malaysia.
Sains Malaysia, 11800 Pulau Pinang, Malaysia. SRICHAI, N., Faculty of Science & Technology, Prince of
MAZLIN MOKHTAR, Chemistry Dept., Univ. Kebangsaan Songkla University, Pattani Campus, Thailand 94000.
Malaysia, Kampus Sabah, Beg Berkunci 62, 88996 Kota SUBRAMANIAM, S., Fisheries Dept., Muara 4069, Brunei
Kinabalu, Sabah, Malaysia. Darussalam.
MOHD FAUZI B. ABDULLAH, Fisheries Research Institute, TAN, B.H., Institute of Advanced Studies, Univ. Malaya,
Glugor, Pulau Pinang, Malaysia. 59100 Kuala Lumpur, Malaysia.
MOHD HUSSAIN B. ABDUL RAHIM, Dept. of Biology, TAN, C.B., Institute of Advanced Studies, Univ. Malaya,
Univ. Pertanian Malaysia, 43400 Serdang, Selangor, 59100 Kuala Lumpur, Malaysia.
Malaysia. TAN, G.S., CHERYL, Dept. of Zoology, National Univ. of
MOHD ZAKARIA ISMAIL, Dept. of Zoology, Univ. Malaya, Singapore, Lower Kent Ridge Road, Singapore 0511.
59100 Kuala Lumpur, Malaysia. TAY, I.B., Dept. of Zoology, National Univ. of Singapore,
MUCHTAR AHMAD. JL., Gunung Kelud, No. I, Pekanbaru, Lower Kent Ridge Road, Singapore 0511.
28142 Riau, Indonesia. TAN, K.P, IRENE, Institute of Advanced Studies, Univ.
MUHAMMAD AKHIR OTHMAN, Dept. of Irrigation & Malaya, 59100 Kuala Lumpur, Malaysia.
Drainage, Jalan Sultan Salahuddin, 50628 Kuala TONG, S.L., Institute of Advanced Studies, Univ. Malaya,
Lumpur, Malaysia. 59100 Kuala Lumpur, Malaysia.
NATHER KHAN, Asian Wetland Bureau, Institute of TUEN, K.L., Malaysian Meteorological Service, Jalan
Advanced Studies, Univ. Malaya, 59100 Kuala Lumpur. Sultan, 46667 Petaling Jaya, Malaysia.
NELSON. B.W., Department of Environmental Sciences, VATID, C.S., Dep. Superintendent, Khao Sam Roi Yot
University of Virginia, Charlottesville, VA 22903, U.S.A. National Park, Khao Daeng, Kuaburi, Prachuab
NG, K.L.. PETER, Dept. of Zoology, National Univ. of Kirikhan, 77150 Thailand.
Singapore, Lower Kent Ridge Road, Singapore 0511. VIKINESWARY SABARATNAM, Institute of Advanced
NG, Y.L., Institute of Advanced Studies, Univ. Malaya, Studies, Univ. Malaya, 59100 Kuala Lumpur, Malaysia.
59100 Kuala Lumpur, Malaysia. WAN MOHD LOTFI W.M., Dept. of Zoology, Faculty of
NORZEDAH ALI, Asian Wetland Bureau, Institute of Life Sciences, Univ. Kebangsaan Malaysia, 43600
Advanced Studies, Univ. Malaya, 59100 Kuala Lumpur, Bangi, Selangor, Malaysia.
Malaysia. WOLANSKI, E., Australian Institute of Marine Science,
NUKUL RUTTANADAKUL, Dept. of Biology, Prince of Townsville, MC, Qld 4810, Australia.
Songkla University, Pattani, 9400 Thailand. YAP, S.Y., Institute of Advanced Studies, Univ. Malaya,
PANG, F.Y., Institute of Advanced Studies, Univ. Malaya, 59100 Kuala Lumpur, Malaysia.
59100 Kuala Lumpur, Malaysia. ZALINA IBRAHIM, Dept. of Zoology, Faculty of Life
PARR, I., National PKS Divn., Royal Forest Dept., Sciences, Univ. Kebangsaan Malaysia, 43600 Bangi,
Paholyonh Road, Bangkok, Thailand. Selangor, Malaysia.
PARRISH, F.D., Asian Wetland Bureau, Institute of ZAMRY KAHPUT, Dept. of Zoology, Faculty of Life
Advanced Studies, Univ. Malaya, 59100 Kuala Lumpur, Sciences, Univ. Kebangsaan Malaysia, 43600 Bangi,
Malaysia. Selangor, Malaysia.
PATHMARANEE. N., Institute of Advanced Studies, Univ. ZAKIR HUSSAIN, Regional Wetlands Coordinator, mCN,
Malaya, 59100 Kuala Lumpur, Malaysia. INROM, AIT, GPO Box 2754, Bangkok, Thailand.
Hydrobiologia 285: 1-5, 1994.
A. Sasekumar, N. Marshall & D. J. Macintosh (eds), Ecology and Conservation of Southeast Asian Marine and
Freshwater Environments including Wetlands.
© 1994 Kluwer Academic Publishers.
Monitoring of sea surface temperature in the South China Sea
K. L. Tuen
Malaysian Meteorological Service, lalan Sultan, 46667 Petaling laya, Malaysia
Abstract
Since the early 1980s, the Malaysian Meteorological Service (MMS) has made concerted efforts to build
up adequate meteorological data base for the South China Sea and the Straits of Malacca. Close rap-
port with local oil companies and universities has enabled MMS to widen its sources of such data.
This data base has been used to carry out a study on the influence of the sea surface temperature (SST)
variations in the South China Sea on the atmosphere during the northern hemisphere winter. Encour-
aging results of the study has prompted MMS to initiate real-time monitoring of the SST in the South
China Sea. These data are available to local scientists who are interested in studying the effects of such
variation on marine life in the Malaysian Exclusive Economic Zone (EEZ).
Introduction Malaysian EEZ, it is also responsible for the col-

lection of surface marine data in the South China
Malaysia, with its long coastline of 4800 kilome- Sea.
tres, lays claim to many of the rich marine re-
sources in the South China Sea. At present, there
is active exploration of petroleum and gas in the Data acquisition programme
continental shelf of the South China Sea and more
production fields are expected to be developed in The primary task of D MM 0 at its early stages of
the near future. The Fishery Department has also development was to create an awareness of the
conducted surveys of fish distribution in the need for marine meteorological and oceano-
country's EEZ. The information gathered is vital graphic data base. Approaches were made to
as the Malaysian Government is presently putting various agencies which carried out marine opera-
great emphasis on the development of deep sea tions such as the oil companies, fishery agencies,
fishing. shipping companies, the Royal Malaysian Navy
The Malaysian Meteorological Service (MMS) and harbour authorities to seek their cooperation
is well aware of the importance of the influence of in making regular observations of the marine en-
the neighbouring seas on the weather and climate vironment. The main targets of the efforts for the
of Malaysia. It also realises the importance of the creation of such data base are the merchant ves-
effects of atmosphere on the surface marine con- sels recruited by DMMO to participate in the
dition and the subsequent effects on marine op- Voluntary Observing Ships' programme (VOS).
erations and associated activities in the Malay- The recruitment is done through the Port Meteo-
sian EEZ. Hence in 1975, it established the rological Offices established at Port Klang, Port
Division of Marine Meteorology and Oceano- Kuching, Port Bintulu and Port Kota Kinabalu
graphy (DMMO). The role of DMMO is not (Fig. la). A total of 110 vessels are now contrib-
confined to the monitoring of the weather over the uting their observations in real-time by transmit-
2
Fig. 1a. Location of the Port Meteorological Offices in Malaysia.
ting their observations to the nearest coastal sta- quire their SST data to complement its own ma-
tions, which in turn transmits the data to DMMO rine meteorological and oceanographic data base.
through the national teleprinter network as well
as the Global Telecommunication System.
For data acquisition in the coastal waters, The surface circulation in the South China Sea
workers on board production oil/gas platforms
have been making regular observations and pass- The variations of the sea surface temperature
ing these data to DMMO through telex or telefax. (SST) in the South China Sea are dependent on
Figure 1b indicates the locations of the platforms the surface current flow. It is known that the
which straddle the Malaysian coastal zone. South East Asian waters are located in an ideal
The data acquired through the VOS programme monsoon region. As such the surface circulation
and the cooperation with the oil companies con- is dominated by these monsoon winds. Figure 2
sists of elements of the atmosphere as well as is a reproduction of Wrytki (1961) results on the
wave heights and sea surface temperatures (S ST). surface circulation during the northern hemi-
Cooperation with local universities which has sphere winter months.
carried out specific scientific studies in the Ma- The focus here is on the variations of the SST
laysian coastal water has enabled MMS to ac- during the winter months. During these periods,
3
• OIL RIG
• OILRIG ON GTS
• LIGHTHOUSE
SOUTH
CHINA
SeA
Fig. lb. Location of the oil platforms and lighthouses which are part of the observation network.
the steadiness of the north-easterly sets up a mon- marked intraseasonal and interannual variabili-
soon current flow in the South China Sea. The ties of the SST in the equatorial South China Sea
coriolis effects on these south-westerly currents and their interactions with local circulation and
cause a piling up of water towards the Vietnam weather. This result prompted MMS to start real-
coast. The pressure gradient then results in a time monitoring of the variation of SST in the
higher current speed in this part of the South equatorial South China Sea.
China Sea.
The South China Sea is also wide enough for
the development of a horizontal circulation Real-time monitoring of the SST
caused by the variation of the wind stress across
the basin during the winter months. The counter The real time monitoring of the SST uses data
current of this circulation flows northwards along within each 1 0 x 1 0 grid and the mean SST in
the coast of Borneo. each such grid is obtained by averaging over a 10
Meteorologists are interested on the variations days period. The SST data are derived from the
of the SST as set up by this circulation during the merchant ships' observations as well as the ob-
winter months. Lim & Tuen (1990) carried out servations from oil platforms. In each month,
one such study which revealed evidence of there are then three 1O-days mean SST profiles,
4
90· 100·
DECEMBER
ernh••c 12 f - - 6,mlday
1S f - - ,]
J4
50 +--
(-f- "
2(
15 .....- J6
>I()O +4-t- '"
- - - CURRENT BOUNDARIES
D DIVERGENCE
'0·
o·
"'.
Fig. 2. Surface circulation in the South China Sea during the northern hemisphere winter (Wyrtki, 1961).
with the last lO-days SST averaged over a period not show such sharp temperature gradients. The
of 8-11 days, depending on the months. central part of the South China Sea experienced
Figure 3 shows an example of a lO-day mean a much slower change in the SST. This was the
SST during the period 1-10 January 1991. The result of the north-westerly counter current of the
profile indicates relatively cold water moved horizontal circulation in the South China Sea
southwards towards the southern part of South which advected the warmer water from the equa-
China Sea. A distinct strong temperature gradi- tor.
ent in the water along the Vietnam coast indicated
the effect of south-easterly current in that area.
However, the consequences of the existence of Remarks
this 'frontal current' near to the Malaysian EEZ
off the east coast of Peninsular Malaysia is not The purpose of this paper is to highlight to the
known. local marine scientists on the existence of such
The whole basin of the South China Sea did distinct temperature gradients in close proximity
5
Fig. 3. 10-day mean sea surface temperatures in the South China Sea during period 1-10 January 1991.
to the Malaysian EEZ during the winter months. References

The Malaysian Meteorological Service hopes to
collaborate with these scientists on the study of Lim, J. T. & K. L. Tuen, 1990. Sea Surface temperature
these variations of sea surface temperature on the variations in the South China Sea during the Northern
Hemisphere Winter Monsoon. Presented in the TOGA
distribution of aquatic resources as well as other Conference, July 1990, Hawaii.
marine phenomena in Malaysian waters. Wyrtki, K., 1991. Scientific results of marine investigations of
the South China Sea and the Gulf of Thailand. N aga Re-
port Volume 2, 195 pp.
A. Sasekumar, N. Marshall & D. J. Macintosh (eds), Ecology and Conservation of Southeast Asian Marine and 7
Neap-spring tidal effects on dissolved oxygen in two Malaysian estuaries
Bruce W. Nelson t, A. Sasekumar 2 & Zelina Z. Ibrahim 3

1 Department of Environmental Sciences, University of Virginia, Charlottesville, Virginia 22903, USA;
2 Department of Zoology, University of Malaya, 59100 Kuala Lumpur, Malaysia;
3 Faculty of Environmental Sciences, Universiti Pertanian Malaysia. 43400 UPM, Serdang, Malaysia
Key words: dissolved oxygen, Malaysia, estuaries, neap-spring tidal cycles
Abstract
The longitudinal distributions of dissolved oxygen change dramatically during neap-spring tidal cycles
in the lower Selangor and Klang Rivers. An oxygen deficit develops in both estuaries when the tidal range
is high. The Selangor River inflow is nearly saturated with D.O., but during spring tides D.O. falls to
as little as 15% of saturation in the middle reaches of the estuary. The Klang River has low D.O.
freshwater input, an oxygen minimum develops during neap tides, and anoxic conditions are produced
by spring tides. These oxygen minima are attributed to the high oxygen demand of sediment that rests
on the bottom during neap tides and is resuspended during spring tides. The ecological effects of tidal
cycling patterns of dissolved oxygen in these Malaysian estuaries need further study, within the context
of land use patterns and other human activities.
Introduction the dissolved oxygen distribution in two Malay-

sian estuaries.
Dissolved oxygen is an important indicator of
water quality in rivers and estuaries. Its amount
and distribution are affected by atmospheric ex- Area description, materials and methods
change, the physical circulation, turbulence, and
water temperature, as well as organic activity such The Selangor and Klang rivers flow from the
as primary production, decomposition of organic western slopes of the Main Range of Malaysia to
matter, and other biological processes. The oxy- the Malacca Straits (Fig. 1). Both drainage ba-
gen demand of efiluents from human activities sins are underlain largely by granitic rocks and
consumes oxygen in natural waters, so that mea- the deep, residual, clayey soils developed from
surements of the percentage saturation are com- them in a tropical climate. These soils are subject
monly used to evaluate water quality. to erosion, and both rivers carry large sediment
While it is recognized that the sediments car- loads during the rainy seasons. The Selangor
ried by rivers have a chemical oxygen demand of River has a larger catchment (1450 km 2 ) than the
their own (Brewer et aI., 1977), the effects on oxy- Klang (712 km 2 ) and a proportionately larger dis-
gen dissolved in natural waters have not been charge. Both drainage basins contain upland
studied extensively. In this paper we show that tropical forest and some lowland swamp forest,
sediments carried in suspension materially affect but agriculture, largely oil palm and rubber, oc-
8
THE STRAIT~I~~~~~~
• URBAN
f;:rtl FOR EST
o OTHER LAND USE
101 0 30'
Fzg. 1. Location Map and Land Use Patterns.
cupy much of the lowlands. The Klang valley estuary (Fig. 5a) is similar. The limit of salt in-
between the federal capital, Kuala Lumpur, the trusion is near Klang town, 15 to 20 km from the
large urban area shown in Fig. 1, and Klang town, mouth. The mean neap and spring tides are 1.3 m
shown near the coast, is heavily populated and and 4.1 m, and the extreme range of tides is 5.1 m.
industrialized. Thus there are important differ- The mean tidal range is 3.0 m in the Klang and
ences in the pollution loads carried by the two a little less in the Selangor. Both would be classed
streams. as mesotidal estuaries. During neap tides, maxi-
Before they enter the sea, the lower portions of mum current speeds are 0.5 m s - 1 and sediment
each river are estuarine, and there ocean tides mix concentrations are less than 100 mg 1- 1. Current
freshwater and sea water. An arrested salt wedge speeds increase to 1.2-1.4 m s - 1 during a 4.0 m
is found during neap tides about 10 km from the spring tide and mean sediment concentrations rise
mouth of Selangor estuary, and at other times salt dramatically to 5000-6000 mg 1- 1. Variations of
intrudes as much as 15 or 20 km (Fig. 2a). Fresh- this kind follow the lunar cycle more closely than
water is always found at Kuantan. The Klang a seasonal cycle, an important distinction between
9
SUNGEI SELANGOR ESTUARY
• Anchor Stotion
§ Fluid Mud
·0
o, 2 3
!
I
Kllomet.,.
Fig.2a. StatIOn Locations III the Selangor River Estuary.
these and other estuaries. Variations in tidal en- with a water sampler and the oxygen probe.
ergy affect many aspects of the lower portions of Rented fishing boats were used to make the lon-
both rivers, such as the salinity distribution, the gitudinal surveys and anchor stations. We thank
current speeds, the sediments in suspension, the the institutions with which we are associated for
cycles of plankton abundance, and, as shown support throughout the course of this work.
below, the oxygen distribution.
In the sections that follow we discuss the varia-
tions in dissolved oxygen concentration that are Results
related to the neap-spring tidal cycle. The data
summarize observations made from 1987 to 1989, Oxygen in the Selangor River estuary
but the conclusions are consistent with other re-
sults obtained by the authors from 1983-1990. The water quality in Selangor River is generally
Dissolved oxygen was measured both by Winkler quite high from the headwaters to the head of the
titration and with a Yellow Springs oxygen meter. estuary, and dissolved oxygen is near saturation
Salinity and temperature were measured in the at the prevailing water temperatures. This trend
field with an Industrial Instruments induction continues into the estuarine portion of the river
conductivity meter and the data were checked by during periods of low tidal range. Figure 2b is a
standard chlorinity titration of water samples. The vertical section of dissolved oxygen along the
distribution offluid mud shown in Fig. 2a is based lower 15 km of the Selangor River obtained when
on echo sounder surveys and vertical profiling the tidal range was 1.1 m (neap). At the mouth the
10
o --MSL.----::::::::::==-=---==--------:-===::::---:::------------===::::-:e;;-:-_-
...
til
!G>
---6------ ~e----------------_--
:E
:i 10
.....
n. DISSOLVED O2 , mg./!.
w
o August II, 1989
20 SELANGOR RIVER ESTUARY A-l.Im.
o 10 IS
DISTANCE FROM MOUTH, Kilometers
Fig.2b. Dissolved Oxygen Concentrations for a 1.1 m Neap Tide.
surface water is saturated with dissolved oxygen. tration of 52 mg 1- 1. Only near the head of the
The degree of saturation decreases upstream, and estuary do they exceed 100 mg 1- 1.
the inflowing surface water, with dissolved oxy- During a spring tide of 3.7 m, eight days ear-
gen concentrations of 6.2 mg 1- 1, is about 80% lier, the measurements shown in Fig. 2c were
saturated. The water column is virtually isother- made. The changes are remarkable. At both the
mal at 28-30 °C (see Fig. 4). Because salinity sea and the river ends of the section the dissolved
differences between the surface and bottom es- oxygen is 4-5 mg 1- 1, sensibly 'less' than during
tablish a density stratification during neap tides, neap tide. Since spring tides minimize salinity
the bottom water has lower dissolved oxygen than stratification, a drop in surface dissolved oxygen
at the surface. The minimum is 4.4 mg 1- 1, about would be caused by mixing. But mixing cannot
40% of saturation. A plot of the data (Fig. 6a) explain the drop to as low as 2.2 mg 1- 1 in the
shows no systematic variations of dissolved oxy- central portion of the estuary! This oxygen mini-
gen with salinity at neap tide, except those due to mum extends through the water column and
stratification. The suspended solids are extremely moves up and down the estuary with the tide. It
low under these conditions with a mean concen- has a mean position 8-10 km from the mouth. A
.A 0 /~MS4Ly?-/~--~~<~'~~~-~~----------~~~---5----
•• 2.5 .. 3 ""-
~ / ",'\ ""-
:E
:i 10
.....
n. DISSOLVED O2, mg.lf.
w
o August 3, 1989
20 SELANGOR RIVER ESTUARY A-3.7m.
o 5 ~ 15

Fig.2e. Dissolved Oxygen Concentrations for a 3.7 m Spring Tide.
11
plot of the data (Fig. 6a) shows a dissolved oxy- appears in the surface water a little after the time
gen minimum at 5-10%0' It seems surprising to of maximum current. The peak sediment concen-
find a minimum during spring tides, when one tration at the surface was 7500 mg 1- 1, while near
expects the stronger currents and greater turbu- the bottom it was 10 700 mg 1- 1. The dissolved
lence 'to increase' the dissolved oxygen by ex- oxygen exceeds 5 mg 1- I, about 85 % of satura-
change between the atmosphere and water. How- tion, at high water slack. The dissolved oxygen
ever, the turbulence acts unexpectedly to produce falls as the sediment concentration rises, and it
an oxygen depression. We know of no sources of reaches a minimum when the sediment concen-
low oxygen water that might discharge into the tration peaks. The minimum dissolved oxygen is
estuary at high tides, so the oxygen minimum must 1. 7 mg 1- 1, about 20 % of saturation. After 1500
be created by internal processes. The main factor hours when the current decreases and suspended
that correlates with the oxygen minimum is the sediment concentration falls, the dissolved oxy-
sediment concentration. For this set of observa- gen increases again until low water slack. During
tions, the average for surface water was 1500 mg the next flood tide, a fresh supply of oxygenated,
1- 1, while the near bottom waters often contained high salinity water appears again at high water
more than 10000 mg 1- 1. Therefore, we attribute slack, and the cycle is poised to repeat itself.
the minimum to the oxygen demand of bottom [A sample collected at station S-4.8 twenty-
sediment that is resuspended during spring tides. four hours later is plotted at 0930 hours in Fig. 3b.
We illustrate this point below. It has high salinity and 5 mg 1- 1 dissolved oxy-
The correlation of sediment resuspension and gen, confirming the renewal of oxygenated water
oxygen depletion may be seen in Fig. 3a, which during the flood tidal phase. The sample marked
shows how the concentrations of dissolved oxy- 'BB' is the oxygen content of freshwater about
gen, salinity, and suspended sediment vary with 45 km above the mouth. The difference between
time during the ebb phase of a tidal cycle at a the two is attributable to the salinity difference.]
station 4.8 km from the mouth during a tide of Figure 3b plots data for an anchor station near
3.7 m. The salinity falls from 30%0 at high water the limit of salt intrusion during a 3.4 m tide. The
to 5%0 at low water, as the tide ebbs. The current surface and bottom water salinities drop during
speed increases to a maximum of about 1 m s - 1, the ebb until at 1400 hours little difference exists
sediment is entrained from the bottom, and it between the two. This is the tip of the salt wedge.
Station S·4.8 7 ·23 ·89 Stotion 5-12 7 ·24·89

"
'' .:
,.
.. ..
30 .0 0 B.8.
0\ 0 0
I
:.~ ~'+.' .
. +-__ ,
0. 0 . 0 5'4.6
-.......J Sur f oc.e O. 0.
o ",• ~ o ~ a Ii
i! 20 .0
~ l .o' o
o
~
•
,. ""'·'7
• Bortom 0. 0.
Z
J
15 ,/t'
',' 0 ,..- I.
....
~
0 ...... ° ,0.
z
o
..
3
U>
10
. '
.. ',
.. .. Surfoee 5. S .
Salinity
\ .
o
Su rface S. S.
I .... .. .... '. I 0
... . . . .~~......-. -' o . - . .. .. -•. .. . . ..

~
'0 12 13 14 I~ 16 11
10 12 13 14 16 17
TIME. ~ours "
TIME. hour,
Fig.3a. Dissolved Oxygen, Salinity, and Sediment Concen- Fig. 3b. Dissolved Oxygen, Salinity, and Sediment Concen-
tration from H.W.S. to L.W.S., Selangor River, Station 4.8, tration from H.W.S. to L.W.S., Selangor River, Station 12,
July 23, 1989,3.7 m Tidal Range. July 24, 1989, 3.4 m Tidal Range.
12
The suspended sediment concentrations rise in concentration remains high. The lack of stratifi-
the surface water to 4000 mg 1-) as the current cation promotes ventillation, and the system re-
speed increases. At high water slack, the dissolved covers completely in the freshwater reaches up-
oxygen, 2-4 mg 1- ), represents water depleted in stream.
oxygen and transported to S-12 from down- The oxygen demand of the freshwater sedi-
stream, i.e., such as that found near low water at ments at the head of the estuary may be less than
station S-4.8. As the tide ebbs and sediment con- that of sediments found farther downstream. The
centration rises, the dissolved oxygen falls. The upper part of the salt intrusion is a site of rapid
bottom water loses half of its initial oxygen; at accumulation of sediment. In higher salinity
1 mg 1- ) it is 15 % saturated. It is noteworthy that water, the sediment contains sulfate-reducing ma-
the oxygen minimum occurs before the peak in rine bacteria, and these probably increase the
sediment concentration and as stratification sediment oxygen demand. The high oxygen de-
disappears at the tip of the salt wedge. This means mand sediment might also be advected from
that the estuarine circulation is a necessary Kuala Selangor near station S-4 in Fig. 2a. We do
condition for maintaining the oxygen minimum. not know how much pollutant this small commu-
After the salt wedge passes downstream, the dis- nity discharges, but if it is the source the sedi-
solved oxygen increases, although the sediment ments must be very sensitive to such environmen-
DISSOLVED OXYGEN, m g./l. DISSOLVED OXYGEN, mo.!l.

o 2 4 6 o 2 4 6
o .' o
o ~
,
I \
\
,,
~ s ,,
'- S
,,
I
(
,
I " ...
\ " ...
\
\
,,
o "
.... 2 ~
,, \
,, ...
:I: \
...
:I:
Go , Go
\
\
I
;
.
1&1
1&1 3
o o 3 ~ !
,!
, !
,I .I
.................... ·····d .,....
\
--- i
4
,, i
4 o "FLUID MUD"
6 \ 7777777777777777
7777777777777777 5
o II' 20 30 o 10 20 30
SALI NITY (0100) a TEM PERATURE (Oe.) SALINITY (0100) a TEMPERATURE (OC.J
S-II 8'11'89 A =1.1 m. S - 10 8 '11' 8 9 A = I. I m.

Fig.4a. Dissolved Oxygen, Salinity, and Temperature Pro- Fig.4b. Dissolved Oxygen, Salinity, and Temperature Pro-
files, Selangor River, Station S-10, August 11, 1989. files, Selangor River, Station S-11, August 11, 1989.
13
tal impacts. We believe, however, that the net were measured near high water on the same day
movement of sediment, pristine or polluted, is as the longitudinal survey illustrated in Fig. 2a.]
downstream on an annual basis. Salinity increases with depth, temperature is con-
In any case, the oxygen minimum occurs in the stant, and oxygen and its percentage saturation
upper part of the estuary 6-10 km from the decrease with depth from 80 % to 60 %. Figure 4b
mouth, where the sediments have a high oxygen is typical of situations where fluid mud is present.
demand in a reach where fluid mud is found. The Salinity, dissolved oxygen, and temperature are
shallowest depths are formed at neap tides by similar to Fig. 4a, except in the fluid mud layer
patches of newly deposited sediment with very shown below the dotted line in the figure. There
high water content. During spring tides, the fluid the salinity (3%0) and dissolved oxygen « 1 mg
mud is resuspended and contributes to the high 1- 1) drop abruptly. The salinity must be inherited
turbidity conditions observed. These fluid mud from sediment deposited at the end of a previous
patches, therefore, play a crucial role in the dis- ebb tide. We believe that when such fluid mud is
solved oxygen budget. Figure 4a shows vertical resuspended by spring tides, it consumes the oxy-
profiles of temperature, salinity, and dissolved gen dissolved in the overlying water and leads to
oxygen where fluid mud is absent. [The profiles the oxygen minimum observed in Figs 2c and 3a.
o Miles
", 1
6 I
Kilometers
Fig. 50. Station Locations in the Klang River Estuary.

14
Oxygen in the Klang River estuary of dissolved oxygen exist throughout the depth.
The estuarine circulation brings saline water into
The location of sampling stations and the distri- the lower 10 km of the estuary along the bottom,
bution of dissolved oxygen in the lower 22 km of and flushes the mixed water seaward in the sur-
the Klang River is shown in Fig. 5a. Because this face layer. A surface plume of low oxygen water
river is subjected to a great deal of environmen- bounded by the two 1 mg 1- 1 isopleths may be
tal stress, the quality of the water passing through seen in Fig. 5b.
the catchment and entering the head of the estu- During a 4.2 m spring tide (Fig. 5c) the dis-
ary is much lower than in the Selangor River. solved oxygen is reduced to zero both at the sur-
Further, the polluted sediments transported by face and the bottom, and the estuary becomes
the river have a very high oxygen demand. anoxic at salinities below 25%0' Ordinarily we
During a 1.4 m neap tide (Fig. 5b), the river water would expect the higher turbulence of spring tidal
may contain as little as 2-3 mg 1- 1 of dissolved currents to ventillate the water, rather than the
oxygen (25-35% saturation) and the bottom reverse. However, the turbulence entrains large
water less than 0.5 mg 1- 1 (less than 10% satu- quantities of bottom sediment during spring tides,
ration). Plots of the data (Fig.6b) show some and the mean sediment concentrations are 6000-
surface waters, both near the head and at the 8000 mg I - I. The anoxia between 6 km and 22 km
mouth of the estuary, with 4-5 mg 1- 1 dissolved is attributed to the oxygen demand of highly pol-
oxygen, which must be due to exchanges with the luted bottom sediments that are resuspended by
atmosphere. Figure 6b has a very pronounced the spring tides and consume the small amount of
oxygen minimum similar to that developed by dissolved oxygen that persists during a neap tide.
spring tides in the Selangor River, but it is much Seaward of the mouth, mixing occurs and the
broader and more pronounced. Surface dissolved dissolved oxygen rises from 3 to more than 5 mg
oxygen concentrations of 1 mg 1- t, or < 15 % 1- I.
saturation, occur over the 10-15%0 salinity range. The results from anchor stations in the Klang
There are no external sources oflow oxygen water are similar to those described above for the Se-
along the reaches represented by the minimum, langor (Fig. 3a). They show decreases in dis-
which appears, therefore, to be due to processes solved oxygen coincident with increases in cur-
within the estuary itself, probably dynamic ex- rent velocities and sediment concentrations,
change with the bottom water. The bottom waters except that complete anoxia occurs in the Klang.
are nearly anoxic. At the seaward end 4-5 mg 1- 1 The effect is more extreme than in Selangor River,
o ~'4S~~2
' '~
... '. ~
.,E5 "' . ------=:

4 3
:Ii 10
°
~ 15
2 > mo.l l.
"-
Ul ~o Dissolved
Cl KL ANG R IVE R ESTUAR Y
Ju ly 22, 198 7
• • A- 1. 4m .
o 10 20
Fig. 5b. Dissolved Oxygen Concentrations for a 1.4 m Neap Tide.

15
E 5
CD
::it 10
~ 15
Dissolved 02. mg.!l.

Q.
ILl 20
a KLANG RIVER ESTUARY
July. 14. 1987
I I
A· 4.2m.
o ~ ~ ~ 20
DISTANCE FROM MOUTH. Ki lom elers
Fig. 5c. Dissolved Oxygen Concentrations for a 4.2 m Spring Tide.
because the Klang discharge is lower in dissolved lieved that ventillation due to the non-tidal, gravi-
oxygen and the polluted bottom sediments have tational circulation and turbulent exchanges pro-
a greater oxygen demand. duced by neap-spring tidal cycling tend to
increase dissolved oxygen, at least in microtidal
estuaries such as Chesapeake Bay (Webb &
Discussion and conclusions D'Elia, 1980; Kuo et al., 1991), where the dis-
solved oxygen in bottom water changed from
The source of oxygen demand in estuaries is gen- 0.5 mg 1- 1 to 4 mg 1- 1 during a neap-spring de-
erally discussed in terms of the consumption of stratification event. It is well known that pollut-
organic matter that comes from primary produc- ant discharges into tidal rivers, such as the Po-
tion (for example, Aston, 1980; Officer et al., tomac, the Rhine, the ScheIdt, and the Baltic, for
1984; Herman et al., 1991). Furthermore, it is be-
KLANG RIVER ESTUARY

SELANGOR RIVER ESTUARY
I
•
0 0 0,
o 7 - 22-87
-' 6
..... 0
0
o NEAP • 7-14-87
cP
..
0. 0 0 0 o?,
E5
..
0 000
•
°4 ••'\
N ..
•
•
•
••
II:>
o ~
/1'-4
0 NEAP 8 a.-
w • ... . '-0. o
>
...J
3
• .- •- • • SPRING
°
<f) 2-
<f)
0 ,
• 8 -3 -89 0 8 -11- 89
0
o 5 10 15 20 25 30 SALINITY. %0
SALINITY, %0
Fig. 6b. Plot of Dissolved Oxygen vs Salinity for Klang River.
Fig. 6a. Plot of Dissolved Oxygen vs Salinity for Selangor July 14, 1987 (closed circles), 4.2 m spring tide; July 22, 1987
River. August 3, 1989 (closed circles), 3.7 m spring tide; Au- (open circles), 1.4 m neap tide. Surface values = circles with
gust 11, 1989 (open circles), l.l m neap tide. bars.
16
example, reduce dissolved oxygen by microbial investigated experimentally by Edwards & Rolley
action on dissolved organic matter. Indeed, the (1965), Martin & Bella (1971), and others who
low oxygen input from the Klang River is an ex- found that entrainment increases the oxygen con-
ample ofthis effect. Seasonal variations have been sumption by a factor of 2-10. A review of this
observed, and Hoppema (1991) described small field is given by Hatcher (1986). However, the
variations in dissolved oxygen during a tidal cycle effects of sediment entrainment from the bottom
in the Wadden Sea without, however, explaining are difficult to analyze quantitatively, and there
its cause. But none of these cited causes of low are few applications of these studies to estuarine
dissolved oxygen in estuarine waters is respon- systems.
sible for the phenomena described above. The plots in Fig. 6 indicate that the circulation
The estuarine portions of the Klang and Se- pattern in a stratified estuarine system sustains
langor Rivers undergo periodic changes in their the depleted oxygen condition, making estuaries
dissolved oxygen budgets due to the oxygen de- fundamentally different in their oxygen budgets
mand of bottom sediment that is alternately de- from freshwater rivers. Neap-spring tidal cycling
posited and resuspended by the variations in tur- must be considered one of the fundamental pro-
bulent energy associated with neap-spring tidal cesses that control dissolved oxygen in an estu-
cycles. If the influent water quality and biological ary. By this mechanism the estuary purges itself
processes were the only controls, we would ex- of the high oxygen demand of sludge that it car-
pect the high turbulent exchanges of spring tides ries as sediment load. The estuarine circulation
to strengthen the circulation, enhance exchange discharges low oxygen and anoxic water at the
with the atmosphere, and increase the dissolved mouth, where mixing with coastal water com-
oxygen content of estuarine water; the stratifica- pletes the re-oxygenation process. The natural cy-
tion and reduced turbulence during neaps should clical variations of dissolved oxygen have a bear-
lead to less oxygenated conditions. The reverse is ing on the water quality standards that should be
observed. Even in the Selangor River, where the applied to estuaries. Furthermore, tropical rivers
freshwater input is well oxygenated and the sur- have higher sediment loads, and sediment with
face waters are near saturation at the ambient inherently higher oxygen demand, than rivers in
temperatures and salinities, a dissolved oxygen other climatic regions. Higher standards of sedi-
minimum develops near the head of the estuary ment and water quality may have to be applied to
on spring tides. This minimum is associated with such systems.
high turbidities and sediment resuspension.
Where the freshwater oxygen input is low and the
sediment has a high pollution load, as in the Klang References
River, neap-spring cycling gives rise to periodic
Aston, S. R., 1980. Nutrients, dissolved gases, and general
anoxia of the estuarine waters. A tide greater than biogeochemistry in estuaries. In E. Olausson and I. Cato
3 m is necessary to create anoxia in the Klang. (eds), Chemistry and Biogeochemistry of Estuaries. J. Wiley
One consequence of the periodic anoxia in the & Sons, New York: 233-262.
Klang River is the mobilization of trace metals, Brewer, W. S., A. R. Abernathy & M. J. B. Paynter, 1977.
such as iron (Nelson, 1988). Oxygen consumption by freshwater sediments. Wat. Res.
11: 471-473.
The influence of sediments usually is discussed Edwards, R. w. & H. L. J. Rolley, 1965. Oxygen consump-
in terms of oxygen consumption by the molecu- tion of river muds. J. Ecol. 53: 1-19.
lar diffusion of reduced substances from anaero- Hatcher, K. J., 1986. Introduction to sediment oxygen de-
bic bottom sediments. For example, the release of mand modelling. In K. J. Thatcher (ed.), Sediment Oxygen
sulfide and its oxidation in the water column ac- Demand: Processes, Modelling, and Measurement. Inst.
Natural Resources, Univer. Georgia Press, Athens: 113-
counts for up to half of the total oxygen consump- 138.
tion in Chesapeake Bay. The oxygen demand of Herman, P. M. J., H. Hummel, M. Bokhorst &
river and estuarine bottom sediments has been A. D. G. Merks, 1991. The Westerschelde: interaction be-
17
tween eutrophication and chemical pollution? In M. Elliott gen uptake rate of estuarine botom deposits. J. Wat. Pollut.
& J. P. Ducrotoy (eds), Estuaries and Coasts: Spatial and Cont. Fed. 43: 1865-1876.
Temporal Comparisons. ECSA 19 Symposium, Olsen and Nelson, B. W., 1987. Tidal regulation of dissolved iron in the
Olsen: 359-363. Klang estuary. EOS 68: 331.
Hoppema, J. M. J., 1991. The oxygen budget of the western Officer, C. B., R. B. Biggs, J. L. Taft, L. E. Cronin,
Wadden Sea, The Netherlands. Estuar. coast. Shelf Sci. 32: M. A. Taylor & W. R. Boynton, 1984. Chesapeake Bay an-
483-502. oxia: origin, development, and significance. Science 223:
Kuo, A. Y., K. Park & M. Z. Moustafa, 1991. Spatial and 22-27.
temporal variabilities of hypoxia in the Rappahannock Webb, K. L. & C. F. D'Elia, 1980. Nutrient and oxygen re-
River, Virginia. Estuaries 14: 113-121. distribution during a spring-neap tidal cycle in a temperate
Martin, D. C. & D. A. Bella, 1971. Effect of mixing on oxy- estuary. Science 207: 983-984.
A. Sasekumar. N. Marshall & D. J. Macintosh (eds), Ecology and Conservation of Southeast Asian Marine and 19
The role of bacteria in nutrient recycling in tropical mangrove and other

coastal benthic ecosystems
Daniel M. Alongi
Australian Institute of Marine Science, PMB No.3, Townsville Me, Qld, 4810, Australia
Key words: bacteria, sediments, nutrients, mangroves, tropics
Abstract
Sedimentary bacteria have generally been recognized as an essential food for protists and invertebrates,
forming the base of benthic food webs. This trophic role has been well documented, but bacteria play
an equally important role as mineralizers of organic detritus and recyclers of essential nutrients. Recent
evidence suggests that this latter role is more important than their trophic function in tropical mangrove
and coastal sediments. Bacteria in these systems are, on average, more abundant and productive than
their counterparts in higher-latitude systems. They account for a disproportionate share of nutrient
uptake to the extent that bacterial communities act as a sink for carbon, processing most of the energy
and nutrients in tropical aquatic systems. Most bacteria remain unconsumed in tropical deposits, dying
naturally and lysing, with the next generation of cells consuming, mineralizing and recycling this mate-
rial either into new biomass or dissolved material. Bacteria in tropical aquatic sediments are ultimately
controlled by inputs of dissolved and particulate detritus, natural mortality and recycling. To replenish
damaged ecosystems in the tropics, restoration of the natural geochemical profile in the sediments is
necessary to re-initiate the growth of bacteria in order to restore the essential recycling processes which
assist in the conservation of nutrients.
Introduction their bypro ducts (gases, etc.) leads to replenish-

ment and recycling of the biosphere.
Bacteria constitute the foundation of all of Earth's In this paper. I will focus on recent evidence
ecosystems, being responsible for the degradation which suggests that bacteria in soils and sedi-
and recycling of essential elements such as car- ments are more abundant and productive than
bon, nitrogen and phosphorus. The energetic supposed reasonable only a decade ago. Microbes
power of bacteria and other microbes is enor- in tropical ecosystems, in particular, are even
mous - greater than all other living organisms more productive and efficient in recycling
combined - and is the powerhouse driving ter- nutrients than their counterparts in higher lati-
restrial and aquatic food webs. Most of the tudes, indicating that they are proportionately
Earth's microbial activity is vested in its 'dump- more important in terms of energy flow in low-
ing grounds' - soils and sediments - simply be- latitude ecosystems. Maintaining the health and
cause this is where organic matter, derived from activity of sedimentary bacteria thus has severe
dead organisms, deposits. Decomposition and re- implications for restoration and conservation of
constitution of this matter into either microbes or endangered coastal systems in the tropics.
20
Bacterial abundance and productivity tent patterns (Table 1). Bacteria in these man-
groves are more abundant (range: 0.02-3.6 x 1011
Mangroves, seagrass beds and un vegetated tidalflats cells g- 1 DW sediment) than in most other
aquatic sediments. This suggests that many of the
Accurate estimates of bacterial densities in man- factors regulating bacteria (grain size, organic C
grove sediments are rare, as most of the early and N, etc.) also range greatly among different
microbial work utilized a variety of plate count- intertidal zones and estuaries over time (Alongi,
ing techniques that resulted in severe underesti- 1988a). Indeed, a partial correlation analysis in-
mates of true density (Ayyakkannu & Chan- dicated that different regulatory factors dominate
dramohan, 1971; Matondkar et al., 1980, 1981). in different intertidal zones. Vertical distribution
Two of the first studies that have estimated abun- of bacteria has been examined only in mangrove
dance using direct counting techniques indicated forests of northern Australia and Papua New
that densities in mangrove soils are generally Guinea (Stanley et al., 1987; Boto et al., 1989;
highest in more organic-rich deposits and in the Alongi. unpubl. data) where density variations
warmest months of the year (Dye, 1983; Hoppe with sediment depth are erratic to 20 cm. These
et al., 1983). erratic profiles are contrary to the decline in den-
More recent studies suggest that bacteria in sities usually observed in temperate sediments
tropical mangrove sediments are characterized by and are likely due to large populations of bacte-
very large temporal and spatial variations (Alongi, ria associated with mangrove roots and rhizomes
1988a, b). In tropical mangroves of Cape York and their exudates - similar to the situation in
peninsula (northern Australia), cell abundance seagrass beds (Moriarty et al., 1986) and salt
varies significantly with tidal height and season marshes (Christian & Wiebe, 1979).
among different systems with no clear or consis- Estimates of bacterial cell production in
Table 1. Bacterial densities, productivity and specific growth rates (Jl) in surface sediments (0-2 cm) in the low, mid and high
intertidal zones of four northeastern Australian mangroves during the austral dry (W) winter and summer (S) wet seasons (after
Alongi, 1988a). Values are means ± 1 S.D.
Estuaries Bacteria Productivity Growth rate

(cells·g- 1 DW x 10 10 ) (gCm- 2 d- l) (d -I)
W S W S W S
Morgan/McIvor
Low 3.4 ± 2.2 2.3 ± 0.3 0.8 ±0.2 5.1 ± 0.9 0.7 5.5
Mid 3.8 ± 0.6 1.6 ± 0.6 0.8 ±0.2 3.2 ± 1.3 0.6 5.5
High 3.1 ± 1.6 0.2± 0.08 0.6±0.2 3.6 ± 0.8 0.2 4.2
Lockhart
Low 35.9 ± 10.9 2.9±0.5 1.3 ± 0.2 2.1±0.7 0.2 3.8
Mid 25.0± 6.3 12.8 ± 3.9 1.7 ± 0.2 1.4 ± 0.1 0.3 0.6
High 14.5 ± 1.7 8.4 ± 2.3 1.7 ± 0.6 1.0 ± 0.2 0.4 0.5
Claudie
Low 28.1 ± 9.4 6.1 ± 0.5 1.4 ± 0.2 1.7 ± 0.2 0.2 0.8
Mid 20.3 ± 4.7 34.4±0.8 2.4 ± 0.4 2.0 ± 0.1 0.6 0.3
High 4.2 ± 1;1 1.6 ± 0.08 1.0 ± 0.5 0.2 ± 0.1 0.6 0.3
Escape
Low 9.1 ± 3.6 8.3 ± 3.4 0.9±0.2 1.6 ± 0.1 0.4 1.0
Mid 31.3±6.3 10.6 ± 2.7 0.9±0.1 1.1 ± 0.2 0.2 0.2
High 1.1 ± 4.1 2.9 ± 1.1 1.0 ± 0.1 1.3 ± 0.1 1.6 0.3
21
tropical mangrove sediments have been few with other than sulfate reducers. High rates of oxygen
nearly all studies conducted in tropical Australia consumption at Ao N am Bor support the idea
(Stanley et al., 1987; Alongi 1988a, b; Boto et al., that rapid bacterial activity in these sediments
1989). Using DNA synthesis methods (see review occurs mainly via non-sulfate reducing pathways
of Moriarty, 1990), these workers found very high (Kristensen et al., 1988, 1991).
rates of productivity (range: 0.2-5.1 g C m - 2 d - 1, Bacterial abundance and productivity in man-
0-2 cm depth) down to a sediment depth of grove sediments are comparable to those in sea-
20 cm. Specific growth rates (production divided grass beds (Moriarty & Pollard, 1982; Moriarty
by standing crop) range from 0.2-5.5 d - 1 in these et al., 1985, 1990; Pollard & Moriarty, 1991). In
deposits, equivalent to a turnover of the entire subtropical seagrass beds. Moriarty et al. (1985)
bacterial community from 3-83 hrs, averaging found production of anaerobic sulphate reducing
15 hrs (Table 1). bacteria much greater than carbon production by
Variations in production among estuaries com- the remaining bacterial flora (= 33 mg C m - 2
plicate generalizations oflarge-scale patterns. For d - 1). As with mangrove bacteria, in seagrass
instance, zonation patterns in the Claudie estuary sediments the bacterial flora exhibits diel varia-
were different between winter and summer. In tions linked to changes in primary production
summer, production rates were equivalent be- (Moriarty & Pollard, 1982). Bacterial growth in
tween low and mid-intertidal forests and much these sediments is stimulated by dissolved exu-
less in the high intertidal, but in winter, the val- dates from seagrass rhizomes (Moriarty et al.,
ues between low and high-intertidal forests were 1986). In the Gulf of Carpentaria, Moriarty et al.
not significantly different (Table 1). In the dry (1990) and Pollard & Moriarty (1991) observed
Australian tropics, bacterial numbers are just as production rates of 3-13 g C m - 2 d - lover a
high as in wet tropical mangroves, but rates of sediment depth of 12 cm. Highest rates of sul-
productivity are lower (Alongi, 1988b). At four phate reduction coincided with below-ground
intertidal sites, bacterial numbers, productivity biomass of roots and rhizomes as similarly ob-
and growth rates fluctuated greatly over time, but served in Thai mangroves by Kristensen et a!.
with no distinct seasonality (Fig. 1). Variations (1991). These rates are equal to 30-80% of net
occurred over tidal cycles suggesting changes in community productivity underscoring the domi-
bacterial activity over very short time scales. On nance of bacterial decomposition in detritus-
creek banks adjoining mangroves in other coun- based food chains.
tries, Moriarty (1986a) and Kemp (1988) ob- In unvegetated mudflats, the abundance and
served similarly high rates of bacterial production production of sedimentary bacteria are similarly
(200-700 mg C m - 2 d - I). variable to those in vegetated (mangroves, sea-
Other methods also indicate high rates of bac- grasses) areas (Alongi, 1988b, 1991; Pollard &
terial activity and nutrient turnover in mangrove Moriarty, 1991). Bacterial abundance is generally
sediments (Blackburn et al., 1987; Kristensen less on sandflats than in other adjacent habitats
eta!., 1988, 1991). In the Ao Yon and Ao Nam (Fig. 1), but rates of productivity and growth are
Bor mangroves in Phuket, Thailand, Danish roughly equal and, in some instances, greater than
workers (references above) found sulphate reduc- in adjacent vegetated sediments (Alongi, 1988b).
tion rates within the range found in coastal tem- On a tropical mudflat adjacent to a seagrass
perate sediments, but lower compared to salt (Enhalus acoroides) bed, production and growth
marshes. This was attributed to moderate redox rates were not significantly different between sites,
levels and high rates of bioturbation by crabs. but sulphate reduction was several times lower
However, the data of Blackburn et a!. (1987) show (0.32 vs 2.2 g C m - 2 d - 1; Pollard & Moriarty,
remarkably rapid turnover of the NH4+ pool in 1991). In the Fly Delta, Papua New Guinea, bac-
these muds indicating that most of the high sub- terial numbers and production are high in mud-
surface activity is due to anaerobes (fermenters?) banks and mangrove sediments compared to
22
• mangrove high Intertidal (station 1 ) • mangrove beach wrack (station 3)

o mangrove low intertidal (station 2) o sand flat (station 4)
40
OL-~--~----~---.--------
;'1750
t" 1500
\ 1250
o
~1000
c
.2
U
'0 "
a
0
500
Qj
.;:
.,
CD
U
III
0
2
';'
'0
..:-.,
~
z:
i0
I;,
~
u.,
Q.
<J)
Time ( months)
Fig. 1. Variations in bacterial numbers, productivity and specific growth rates in mangrove, sandflat and beach-wrack sediments
in the Australian dry tropics.
(From Alongi. Journal of Marine Research. 1988b. With permission.)
most other intertidal deposits (Alongi, 1991. un- «6:1 versus 10-25:1) than in the mangroves,
published data). A comparison of these data indicating that higher quality material deposits
(Table 2) indicate equivalent rates of productivity onto the mudbanks and perhaps offsetting the
and biomass in mangrove versus unvegetated lower absolute concentrations of nutrients. As in
habitats. Dissolved and particulate nutrients in salt marshes, a close relationship undoubtedly
the mudbank are in lower concentration than in exists among sedimentary bacteria, porewater
the mangrove muds, but the C:N ratio of the nutrients and the plants per se, but such complex
mudbank deposits is, on average, much lower synergistic-antagonistic interactions are hard to
23
Table 2. Grand mean baeterieal densities and rates of DNA subtidal studies have focused on coral reef la-
and protein synthesis averaged over a sediment depth of goons and in coastal and shelf sands and muds
20 em at various mudbank and mangrove areas of the Fly
Delta, Papua New Guinea (After Alongi, 1991 and unpub- in the central Great Barrier Reef province
lished data). Ranges are in parenthesis. (Moriarty, 1983; Hansen et al., 1987; Alongi
et al., 1989; Alongi, 1989a; Ducklow, 1990).
Numbers Production (mg C m - 2 d - I) Summarizing these data, Alongi (1990) found that
(cells g-I OW)
DNA Protein subtidal communities are dependent on sediment
temperature, although to a much lesser degree
Mudbanks than in intertidal deposits (Fig. 3). Turnover of
AIB 1.5 x 10 10 (0.9-2.4) 229 (44-495) 41 (33-49)
URA 1.3 x 10 10 (0.7-1.6) 413 (184-903) 75 (44-95) bacteria was generally faster in the intertidal sedi-
UMA 1.4 x 10 10 (0.1-8.3) 304 (1l-697) 18 (1l-27) ments (':::;4 days). Both data sets indicate a simi-
SOB 5.0 x 10 9 (1.l-8.2) 187 (12-492) 34 (17-41) lar trend of more rapid turnover with higher tem-
Mangroves perature (Fig. 3). Bacteria in coral carbonates
'SAl 1.5 x 10 10 (0.6-2.0) 334 (0-916) 55 (19-104)
'SA2 9.0 x 109 (5.6-17.0) 223 (66-604) 34 (8-132)
exhibit equivalent specific growth rates compared
'NFl 2.9 x 10 10 (1.4-5.5) 240 (86-498) 73 (34-217) to communities in coastal subtidal muds and
'NF2 1.0 x 10 10 (0.6-2.3) 180 (0-349) 31 (0-58) sands, but bacterial numbers and total produc-
'RBI 1.8 x 10 10 (1.3-2.1) 305 (0-585) 96 (20-298)
*RB2 9.7 x 109 (1.5-17.1) 251 (63-412) 34 (2-106)
tivity are generally less on coral reefs (Hansen
et al., 1987; Ducklow, 1990). Bacterial biomass
* SA = Sonneratia-A vicennia, NF = Nypa fruticans, RB = and growth rates in stromatolites formed from
Rhizophora-Bruigeria forests. Mudbank station abbreviations
four different types of benthic cyanobacterial mats
are explained in Fig. 2 legend.
are similar to those in soft carbonates on coral
reefs (Moriarty, 1983).
unravel (see bacteria-nutrient interactions sec- In coastal and shelf sediments of the central
tion). Great Barrier Reef province, bacteria are very
Bacterial carbon production at all of the New abundant compared to temperate subtidal depos-
Guinea stations was estimated from rates of pro- its, n,'mging from 0.5-20.8 x 1010 cells g-I sedi-
tein eH-leucine incorporation) and DNA eH- ment dry weight in inshore areas and from 0.4-
thymidine incorporation) synthesis. Both meth- 2.7 x 10 10 cells g-I OW in middle and outer shelf
ods did not agree well with significantly higher sands (Alongi, 1989a; Alongi et al., 1989). Of par-
rates of DNA synthesis than protein synthesis ticular interest is the fact that large amounts
(Table 2). Nevertheless, they did agree in that ( '" 25 000 tonnes C yr - I) of mangrove litter are
both methods revealed very active, subsurface exported from adjacent mangrove forests and de-
bacterial communities (Fig. 2), in agreement with posited along a narrow band in subtidal areas
the high subsurface rates of thymidine uptake along the north Queensland coast. Sampling of
observed in the Australian mangroves (Stanley surface sediments in this region indicates that this
et al., 1987). High rates of subsurface production litter is highly refractory and of poor nutritional
and growth in these and other tropical sediments quality, enhancing bacterial activity only in semi-
(seagrasses, Moriarty et aI., 1990) indicate that enclosed channels where the litter can deposit in
these habitats are important sites of organic large quantities (> 4 kg OW litter m - 2 of sea-
matter decomposition and nutrient diagenesis. bed). However, bacteria in this shallow area are
very productive and actively growing even to
20 cm depth, suggesting that factors (physical dis-
Coastal subtidal environments turbances from tidal scouring and cyclones) other
than subsurface burial and accumulation
The dynamics of bacteria in coastal subtidal habi- of litter, over the long-term, supports this high
tats in the tropics has rarely been investigated, if productivity (Alongi, 1992). Whether or not
one excludes submerged seagrass beds. Most such events occur in shallow coastal areas in
24
A DNA SYNTHESIS (llgC.cm·3.h· l ) B. Il (h· l )

8 16 24 32 40 0 0.04 0.08 0.12 0.16 0.20
t
.60
e=AIB I
0= URA 3.2
2.2
ll.=UMA
" =SOB
1.0
f 6.8
16
4 3.5
;. 3.5
20
E'
i2.
~ C. PROTEIN SYNTHESIS (llgC.cm· 3.h· l ) D. Il (h· 1)
fu
Q 0.8 1.6 2.4 3.2 4.0 0 0.02 0.04
00
(o>~>ll.)
4
0.13
8
t ,
0.21
12
<O~
, 0.26
16
t 0.32
,;. 0.14
20
Fig. 2. Vertical profiles of bacterial DNA and protein synthesis in intertidal muds (4 stations) on different mangrove islands of
Papua New Guinea. The 4 stations are abbreviated as sites AlB, URA, UMA and SOB. Station symbols in parentheses in
graphs C and D represent statistically significant differences among the sites.
(From Alongi, Journal of Erperimental Marine Biology and Ecology. 1991. With permission of Elsevior Science Publishers.)
South-east Asia is problematical as no similar the Fly and adjacent rivers. Within the Fly Delta
studies have been conducted in this region. It is and out into the adjacent Gulf of Papua, bacte-
likely that this phenomenon occurs in Asian areas rial numbers vary widely among sediment types
where quiescent conditions permit the accumula- and with sediment depth (Alongi et al., 1992). On
tion of plant detritus adjacent to massive man- average, numbers of bacteria are higher (range:
grove forests. 0.9-6.5 x 1010 cells g- 1 DW) in the more stable
In contrast to the reef-dominated shelf of the seabed of the Gulf of Papua than in the eroded,
central Great Barrier Reef, the Gulf of Papua in disturbed bottom of the delta (range: 0.007-
the northern Coral Sea is dominated by massive 1.3 x 1010 cells g-l DW). These densities are
freshwater and suspended material runoff from greater on average than off the Amazon River
25
• •
• subtidal muds - sands
50
• o intertidal muds - sands
• o coral reef carbonates
• • •• • • seagrass muddy sands
• •
20
~ • • • •
• •
10
• • •
•
•
0
subtidal y = 46.4 - 1.44x

• r' =-0.20
0 •
,..co
"iii
•
:!!. 7
• •
•
~
'0>"
E 6
• •
•• • •
:J
fo-
•
•• • _ _•_• • _ _ _ _ _ _ _ _ _ _
0
0
• •
4 __________ 1... _ _ _ _ _ _
o 0 0 ••
o [] • []
intertidal y = 5.55 - 0.146x
• 0
• r' = -0.48
0
0 • 0 0
00 0
0 0
•
• 0
i
15 20 25 30 35
Degrees (OC)
Fig. 3. Relationship between turnover of bacterial populations and surface sediment temperature in various benthic habitats within
the central Great Barrier Reef province.
(From Alongi. Oceanography and Marine Biology Annual Reviews. 1990. With permission.)
(Aller & Aller, 1986), but equivalent to values in available information, albeit scarce, indicates very
the central Great Barrier Reef shelf (Alongi, abundant and productive bacterial assemblages
1989a). Rates of DNA and protein synthesis were in tropical estuarine and marine sediments.
very erratic ranging in surface deposits from
0-387 mg C m - 2 d - , (DNA synthesis) and from Bacteria- Nutrient interactions
0-854 mg C m - 2 d - , (protein synthesis). As for
bacterial numbers, production and specific Regardless of habitat and sediment type, the
growth rates were generally higher in the Gulf of growth and division of bacterial cells depends
Papua, mirroring rates of benthic respiration (63- upon sustenance from dissolved and particulate
780mg C m- 2 d-', Alongi etal., 1992). nutrients. For this reason, bacterial numbers and
More data are needed on the distribution, rates of production and specific growth usually
abundance and productivity of bacteria in sedi- correlate with standing amounts and fluxes of dis-
ments from a variety of tropical habitats. The solved and particulate organic matter {Meyer-
26
Reil, 1984: Moriarty et al., 1986; Herndl et al., water interface, but when sediments were poi-
1987). soned m the chambers to kill the flora and fauna,
The studies of Stanley et al. (1987) and Boto rates of amino aCId flux ranged from 27-69 mg N
et al. (1989) indicate a close coupling between m - 2 d - 1. Concurrent measurements of bacterial
bacteria and dissolved nutrients in tropical man- growth indicated that these fluxes accounted for
groves. Stanley et at. (1987) examined the com- between 9-38% and 5-19% of the nitrogen and
position and bacterial utilIzation of dIssolved free carbon required to support the observed levels of
amino acids in mangrove sediments on Hinchin- bacterial productivity.
brook Island, Australia. They found that in high More extensive nutrient flux experiments were
intertidal sediments, bacterial growth rates cor- conducted by Boto et al. (1989) at the same
related positively with total amino acid concen- sites. Using more sophisticated chambers (Fig. 4;
trations. Flux chamber experiments revealed complete descriptIon in legend), they found
negligible ammo acid flux across the sedIment- that dissolved orgamc carbon behaved simIlarly:
Fig 4 Photograph of glass chambers used to measure DOC fluxes across the sediment-water mterface m mangroves Chambers
have a sediment surface area of 0 007 m2 and are of 1 liter volume Arrows depict p = port for DOC samples, m = self-con tamed
motors to stir propellers m (g =) glass chambers Complete deSCriptIOn of the chamber assembly can be found m Boto et af (1989)
27
despite a high concentration gradient of DOC tropical subtidal sediments (Alongi et al., 1989;
between sediment porewaters and overlying tidal 1992). In coastal sediments off Hinchinbrook
waters, significant efflux of DOC was rarely de- Island, DOC fluxes across the sediment-water
tected unless sediments were poisoned. The dif- interface provided 51 % of the carbon required to
ference in flux between poisoned and unpoisoned sustain the measured rates of bacterial produc-
chambers was equated to uptake by bacteria tivity. Efficient DOC-bacteria recycling was also
(Fig. 5). Efflux rates of DOC varied widely with observed in nearshore sediments within the Fly
tidal height and season (range: 0-2.4 g C m - 1 d). River and in the Gulf of Papua. In unpoisoned
but on average provided 35% of bacterial carbon chambers, DOC fluxes were usually into the sedi-
requirements at the sediment-water interface. ment (range: -797 to + 514mg C m- 2 d- l ) in-
Similar experiments conducted on mudbanks in dicating uptake by bacteria at the sediment-water
Papua New Guinea indicate a similarly close cou- interface. When these sediments were poisoned,
pling between bacterial growth and DOC flux the fluxes either stopped or reversed direction.
(Alongi, 1991). Based on thymidine uptake rates, these fluxes
This close relationship appears to hold true for supplied, on average, 60% of the bacterial carbon
requirements. The leucine-derived rates suggest
5.0
the percentage was as high as 74 % (Alongi et al.,
1992).
There is some evidence that sedimentary bac-
teria in the tropics are phosphorus-limited
(Alongi, 1991). In mudbanks of the Fly River,
4.0 specific growth rates of bacteria correlated sig-
nificantly with phosphate and dissolved organic
1 phosphorus in the porewaters, and with total
0 phosphorus. Densities of bacteria also related
.s'" positively with total P. It is not unreasonable to
~
~ 3.0 expect P limitation in the tropics. For example, a
~
"" positive response to phosphate enrichment in the
"
.5
growth of mangrove trees (Boto & Wellington,
c
0
:; 1983) and of the seagrass Syringodium filiforme
:e 11)
(Short et al., 1990) has been observed.
g 2.0
8 It is difficult to conduct enrichment experiments
0
0
0
on sedimentary bacteria to test for nutrient limi-
tation. Addition of dissolved glucose, for example,
may stimulate cell division but the addition of the'
1.0 water per se may be the stimulatory factor rather
than the DOC, as found by Christian & Wiebe
(1979) in salt marsh soils. Another vexing prob-
lem is that the results of such an experiment may
well depend upon the methodology used to mea-
OL-----r-----~----+-----+-----;_
o 60 120 160 240 300
sure bacterial activity and biomass.
Time(Min)
Fig. 5. Variations of DOC concentrations with time for wa- Bacteria as a sink for carbon: implications for con-
ters in flux chambers (see Fig. 4) overlying untreated (.) and servation and management
poisoned (.) mangrove sediments. Hinchinbrook Island,
Australia. The fate of bacteria in aquatic sediments is
(From Boto et al. Murine Ecology Progress Series, 1989. With permission.) poorly understood, and has been the focus of
28
considerable controversy for many years. Inad- isms. This trophic linkage decreases with sedi-
equate methodology and multiple roles are the ment depth (and/or away from oxidized linings of
major reasons for much of this disagreement. Es- biogenic structures) where sediments are more
sentially, bacteria in sediments serve two roles: anaerobic and less populated by microbial preda-
(1) as food for other organisms and (2) as miner- tors. Assuming that grazing intensity lessens in
alizers of organic matter (Blackburn, 1988; these deposits, unconsumed subsurface bacteria
Tenore, 1988). Both roles are closely intertwined. die naturally and lyse, with the next generation of
Early trophic studies showed that most benthic bacteria consuming and mineralizing this mate-
organisms feed on bacteria. Subsequent research rial into new biomass or as dissolved material.
dealt with the role of bacteria (and other microbes) This activity maintains the cycling of essential
in detritus-based food webs, particularly their nutrients.
participation in coprophagy and detrital aging. A variety of observations from tropical man-
This work led to the concept that bacteria enrich groves supports this scenario: (1) densities of
the protein content of detrital plant material and protozoans, meiofauna and small macroinfauna
fecal pellets by decomposing refractory compo- are low with extrapolation oflaboratory ingestion
nents over time, with this microbially-enriched rates indicating that only a small proportion of
material being ingested (or re-ingested) by detri- bacterial biomass is consumed (Tietjen & Alongi,
tivores to derive a considerable amount of their 1990); (2) bacterial biomass is large (> 200 g C
nutrition from digestion of the bacteria and their m - 3 of sediment) and productivity exceeds the
mucus. highest rates of mangrove primary production;
This trophic role has been over emphasized (3) micro algal biomass is low with little measur-
because of the popUlarity of a seminal paper by able benthic net primary production; (4) the dis-
Gerlach (1978) who estimated the effect of graz- solved and particulate carbon pools are large;
ing by larger organisms on bacteria in a hypo- (5) tidal cycle measurements indicate that bacte-
thetical benthic community. He concluded that rial growth is regulated by temperature and not by
grazing pressure and other invertebrate activities grazing; (6) bacterial abundance and production
(e.g. excretion) stimulate bacterial growth and usually correlate with physicochemical factors,
detrital decomposition in sediments. More recent but rarely with other benthos; (7) as explained
microbial-detritus studies have indicated that this earlier, dissolved free amino acids and DOC in
concept is overly simplistic (see reviews of Alongi, the sediment porewater are rapidly utilized by
1989b and Kemp, 1990). Bacteria are not always bacteria at the sediment-water interface and
tightly linked trophic ally within benthic food (8) there is a close similarity between the compo-
chains, being not only ingested by larger animals, sition of the free amino acid pool free in the pore-
but having an unknown. but probably large, pro- water and that found within the intracellular pools
portion of biomass left unconsumed to partici- of some sulfate-reducing bacteria (Stanley et al.,
pate in biogeochemical cycles. 1987; Alongi, 1988a, b; Boto et at., 1989).
Alongi (1989b) constructed a benthic carbon In reviewing the literature, a pattern emerges in
flux model (the 'carbon-sink' hypothesis) based which studies that have found a grazing effect
on work in tropical mangroves which states that nearly always were conducted in organic-poor
sedimentary bacteria efficiently recycle nutrients habitats, such as on sandy beaches, intertidal
within microbial food webs in order to conserve sandflats, surfaces on plant leaves or on or near
essential elements within the ecosystem (that is to biogenic structures. Conversely, a significant
prevent its loss). This concept proposes that bac- grazing effect has rarely been observed in organic-
teria serve as a trophic link with the rest of the rich environments, such as in microcosms, muds
benthic food web in surface sediments and in and in muddy vegetated sediments. Based on this
burrow linings, tubes and other habitats that are observation, Alongi (1989b, 1990) proposed that
aerated and inhabited by larger benthic organ- a significant trophic loss in bacterial biomass (i.e.
29
a grazing effect) will be observable only if bacte- cling nutrients; high rates of primary production
rial growth rates are less than or equal to rates of by small, highly efficient plankton coupled with
ingestion by predators, for example, in organic- rapid rates of decomposition by microbes, appear
poor habitats where bacterial growth is compara- to be the most important factors responsible for
tively slow, mainly because they are nutrient- efficient nutrient recycling in these tropical sys-
limited. In organic-rich habitats, bacterial growth tems (Lewis, 1987).
is fast, greater than rates of consumption, so no Tropical aquatic and terrestrial ecosystems
grazing effect is discern able. In these latter envi- appear to be proportionally more dependent upon
ronments, bacterial abundance will be controlled bacteria and other microbes for their health and
by carrying capacity, nutrient supply and envi- survival than ecosystems of higher latitudes. In
ronmental conditions (Moriarty, 1989). The essence, microbial communities are responsible
amounts of bacterial biomass consumed thus de- for a greater share of the fluxes of energy and
pends upon not only how fast they are eaten, but nutrients through tropical food webs inhabiting
how fast they divide. Kemp (1990) has come to terrestrial soils and aquatic sediments. This im-
a similar conclusion regarding the fate of benthic plies that degradation and, conversely, restora-
bacterial production concluding that 'the impli- tion, of tropical ecosystems is greatly dependent
cation is that most of bacterial production is upon the extent to which benthic microbial com-
ultimately respired and remineralized by a communities and their geochemical environment are
plex microbial food web. Thus, a majority of or- disturbed. There is little information on the effects
ganic matter input to the benthos is not consumed of disturbance (e.g. pollution, dredging) on these
by larger metazoans, i.e., from the large metazoan communities, but some indication of possible
point of view, the microbial food web is a sink for effects can be gleaned from some studies con-
energy and major nutrients'. ducted in habitats which possess some charac-
Microbial food chains in other tropical eco- teristics of polluted environments. For example,
systems appear to be similarly highly efficient Moriarty (1986b) investigated the role of bacteria
in nutrient recycling (Lathwell & Grove, 1986; in the carbon cycle of ponds used for culture of
Vitousek & Sanford, 1986; Lewis, 1987; Ruess & penaeid prawns at Gelang Patah in Malaysia. He
McNaughton, 1987; Hatcher et al., 1989; Singh found that these aquaculture ponds support
et al., 1989, 1991; Furtado et aI., 1990). As de- greater numbers and production of bacteria than
tailed by Lathwell & Grove (1986) and Singh other aquatic habitats, accounting for most of the
et al. (1989, 1991), at least half of all tropical soils organic carbon (as chicken manure) added to the
are highly weathered and impoverished of nutri- ponds. A similar study was conducted in fish
ents, requiring the need for very efficient nutrient- ponds in India (Olah et al., 1987) where bacterial
conserving mechanisms. In dry tropical savanna, and chemical oxygen uptake accounts for the
Singh et al. (1989, 1991) found that microbial bio- major portion of total sediment oxygen consump-
mass acts as a source of nutrients for plants. tion. This study shows that subsurface sediments
Scarce nutrients are immobilized and conserved very rich in organic matter play a limited role in
by being tied up in microbial biomass during dry total benthic metabolism. The authors suggested
periods when the growth of plants is low, and that these subsurface deposits act as energy traps.
released during monsoon periods to stimulate Mechanical raking of pond bottoms was recom-
plant growth. Tropical rain forests similarly mended to maintain aerobic degradation and
accumulate high microbial biomass to store nu- inhibit anaerobic metabolites.
trients and show evidence of rapid nitrogen turn- . Anthropogenic and natural disturbances are a
over, suggesting very abundant and productive major factor influencing the species composition
microbial communities (Vitousek & Sanford, and growth of microbial communities in benthic
1986). In comparison with temperate lakes, tropi- deposits (Findlay et al., 1990a, b). The effects de-
cal freshwater bodies are more efficient in recy- pend upon the frequency, duration, real extent
30
and nature (dredging vs sewage) of the disturb- tions (Eh, pH, O2 profiles) in coastal sediments
ance, but some effects are predictable based on in the tropics are the key to restoring damaged
information from other ecosystems (Table 3). ecosystems and conserving healthy habitats. The
Table 3 shows that disruption of tropical sedi- natural sedimentary profile and hydrological con-
ments or soils nearly always leads to changes in ditions maintaining it must be restored in order to
the composition and growth state of the micro- re-initiate microbial biogeochemical cycles which
biota. Such changes lead in tum to loss of nu- drive ecosystem-level processes recycling and
trients or to leaching of metabolic by-products of conserving scarce nutrients. This advice is par-
microbial growth which may be toxic to some ticularly pertinent for tropical aquatic systems as
wildlife. For example, clear-felling and disturb- they appear to be as fragile as (or more so than)
ance of mangrove soils leads to the oxidation of ecosystems of boreal and temperate latitudes
pyrite (Fe S2) usually present in anaerobic sedi- (Clough, 1988; Hatcher et at., 1989).
ments and subsequent release of sulfuric acid
causing acidification.
Other forms of environmental damage usually Acknowledgements
lead to changes in tidal flushing or other hydro-
logical characteristics, or to a build-up of organic I thank the Australian International Development
matter (e.g. sewage, oil spills, animal faeces from Assistance Bureau (AIDAB) for supporting my
aquaculture ponds). Bacteria can decompose vast travel to Malaysia to participate in this seminar
amounts of additional detritus, but processing as a keynote speaker. Dr A. Sasekumar is thanked
efficiency tends to decrease with increasing input. for his assistance. Drs B. Clough. W. J. Wiebe
Such organic loading usually shifts healthy and J. H. Tietjen provided helpful comments on
aerobic-anaerobic systems to a state of complete the manuscript. Kathleen Vincent typed the
anaerobiosis. Anaerobic processes are less effi- manuscript. Further support was provided by the
cient in recycling nutrients, resulting in the Australian Institute of Marine Science. Contri-
build-up and release of toxic sulfides. bution No 665 from the Australian Institute of
Microbial and associated geochemical condi- Marine Science.
Table 3. Probable effects of some types of environmental damage on microbial food webs with subsequent effects on the rest of
the ecosystem.
Type of disturbance Microbial effect Ecosystem consequences
Clear-felling of vegetation Loss of biomass and leaching of Impoverishment of sediment, in-

nutrients ability to restore vegetation with
erosion of clay
Chemical defoliation Death of most microbes; suspen- Loss of most trophic groups
sion of nutrient recycling
Reclamation for agriculture/aquaculture Oxidation of soils and destruction Release of sulphuric acid and sub-
of anaerobic microbial biomass sequent acidification; high alu-
minium concentrations to toxic
levels
Sewerage, change in hydrology, dredge Shift from mixed aerobic/anaerobic Less efficient nutrient recycling,
spoil dumping, increased biomass from microbes to anaerobic microbial buildup of organic matter with
aquaculture ponds, spills of heavy oils processes only release of toxic amounts of HzS
31
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Studies on the production of useful chemicals, especially fa'tty acids in

the marine diatom Nitzschia conspicua Grunow
Wan-Loy Chu, Siew-Moi Phang* & Swee-Hock Goh

Institute of Advanced Studies, University of Malaya, 59100 Kuala Lumpur, Malaysia
(* author for correspondence)
Key words: lipids, fatty acids, carbohydrates, proteins, culture age, Nitzschia conspicua
Abstract
A local marine diatom, Nitzschia conspicua Grunow, was cultured in enriched synthetic seawater using
flasks (agitated by magnetic stirring) and a 1,21 fermenter. Lipids, fatty acids, proteins, carbohydrates
and ash of the flask cultures were determined at various stages of growth (day 3, 5, 7, 10, 13, 15 and
17), The fermenter culture was harvested during the stationary phase for similar chemical analyses.
N. conspicua attained a higher biomass concentration during the stationary phase when cultured in the
fermenter (188 mg dry weight 1- 1) than in flasks (140-151 mg dry weight 1-1). However, both systems
showed similar specific growth rates based on chlorophyll-a concentration. Appreciable amounts of the
essential fatty acids 20:4 (0.6-4.7% total fatty acids) and 20:5 (1,9-4.7% total fatty acids) are present
in this diatom. Maximal amounts of these fatty acids were produced after 7 days' growth (i.e. 2 days
after the end of the exponential phase). Lipids, fatty acids, proteins, carbohydrates and ash varied with
culture age in N. conspicua.
Introduction acids de novo. These fatty acids are accumulated

through the food chain which consists of micro-
Marine diatoms are known to contain significant algae. Biotechnological exploitation of microal-
amounts of polyunsaturated fatty acids (Kyle gae (especially marine diatoms) as a source of
et al., 1988) such as arachidonic acid (20:4), EFA as an alternative to fish oil has generated
eicosapentaenoic acid (20:5) and docosa- great interest (Kyle et al., 1988).
hexaenoic acid (22:6). Recent studies showed that In another line of development, micro algae
these essential fatty acids (EFA) have hypolipi- have also been exploited in the aquaculture in-
demic and antithrombotic properties (W ojenski, dustry as high-value feeds (Sommer, 1990). This
1991; Harris, 1989). These fatty acids are is due to the high nutritional qualities of these
commercially important in the pharmaceutical micro algae which contain essential nutrients (es-
industry as they are precursors of eicosanoids pecially polyunsaturated fatty acids). Marine dia-
(prostaglandin, prostacylins, thromboxane and toms like Skeletonema costatum, Thalassiosira
leucotrienes) which are used in the treatments of pseudonana, Phaeodactylum tricomutum, Chaetoc-
high blood pressure, asthma, menstrual pains and eros calcitrans and Nitzschia sp. have been mass-
induction of labour (Borowitzka, 1988). cultured for aquacultural purposes (DePauw &
The current source of these EFA is from ma- Persoone, 1988).
rine fish but they do not synthesise these fatty In view of the expansion of biotechnology and
34
aquaculture in Malaysia, it is timely to screen for Similar inoculum as the flask culture experiment
useful products like exotic fatty acids from local was used. Approximately 20 ml samples were
algal resources. From a previous screening offour withdrawn aseptically from the vessel every 2-3
local diatoms (two freshwater and two marine days for growth measurement by chlorophyll-a
isolates) for these products, Nitzschia conspicua determination. The cells were filtered for dry
attained the highest biomass, and contained weight determination (100°C, 24 h) and chemi-
appreciable amounts of arachidonic acid (AA) cal analyses during the stationary phase.
and eicosapentaenoic acid (EPA). N. conspicua
was selected for detailed investigation on its
chemical (especially fatty acid) composition at Chemical analyses
different stages of growth.
Lipids were extracted using MeOH-CHClr H 2 0
(2:1:0.8) and determined gravimetrically (Bligh &
Dyer, 1959). Lipids were transesterified in 1 N
Materials and methods
sodium methoxide (60°C, 20 min) and the fatty
acid methyl esters obtained were analyzed by gas
Growth
chromatography (Christie, 1989).
The gas chromatograph (Shimadzu GC 14A)
N. conspicua Grunow (isolate number 111) was
was equipped with a polar capillary column (DB
isolated in axenic state (Hoshaw & Rowowski,
23) and the following temperature program was
1973) from boat scrapings collected from Umbai,
used: 130°C (2 min) increased to 200°C (1 min)
Melaka. The culture is deposited at the Algal Cul-
at a rate of 3 ° C min - 1 then further increased to
ture Collection at the Institute of Advanced
230 ° C at 2 ° C min - 1 and held for 5 min. Injec-
Studies, University of Malaya. The diatom was
tor and detector temperatures were set at 300 ° C
grown in synthetic seawater (Marine Environ-
and the carrier gas (N 2 ) was at a flow rate of
ment) enriched with NaN0 3 (2.9 mM), K 2 HP0 4
0.6 ml min - 1. Identification of fatty acids was
(0.043 mM), Na2 Si0 3 (0.088 mM) and vitamin
based on retention times of authentic fatty acid
mixture containing cyanocobalamin (2.95 x
standards. Quantitation of the fatty acids was
10- 5 mM), thiamine-HCl (1.19 x 10- 4 mM) and
based on integrated areas of the peaks of the
biotin (1.64 x 10- 4 mM). The cultures were
chromatograms.
grown in conical flasks (cap. 21) and agitated by
Proteins were extracted in 0.5 N NaOH
magnetic stirring to prevent cell clumping and
(80 °C, 20 min) and assayed according to the
settlement. These cultures were placed under cool
dye-binding method (Bradford, 1976). Carbo-
fluorescent lamps (31.9 J.LEm - 2 s - 1) with a
hydrates were extracted in 2 N HCI (80°C,
12: 12 h light-dark cycle in a controlled tempera-
20 min) before determination by the phenol-
ture room (26-28 0e). A 10% inoculum from an
sulphuric acid method (Kochert, 1978). Chloro-
exponential phase culture (OD 7oo at 0.016) was
phyll-a was extracted in acetone and left in the
used for the total culture volume of 11.
dark (-20°C, 24 h) before being determined
The cultures were harvested during the light
spectrophotometric ally (Strickland & Parsons,
phase at various stages of growth. Cells from 2-4
1968).
flasks were pooled and filtered for chlorophyll-a
determination and chemical analyses after 3, 5, 7,
10, 13, 15, 17 and 20 days' growth.
Results and discussion
The diatom was also grown in a 1.2 I fermenter
which was agitated by an impeller (150 rpm) and
Growth
aerated by an air-sparger (1500 ml min - 1). Light
was provided from two fluorescent lamps Growth based on chlorophyll-a concentration
(31.9J.LEm- 2 S-I, 12:12h light-clark cycle). (Fig. 1) showed that the exponential phase of
35
Chlorophyll-a (mg/l) Table 1. Chemical composition (% dry weight) of N. con-

1.---~~--~---------------------, spicua and another species of Nitzschia.
Chemical Flask 1 Fermenter 2 Nitzschia"

o sp.
o
Ash 33.3-46.7 26.1 20.4
Lipids 2.7-14.1 14.2 17.6
Proteins 3.0-13.8 16.8 13.4
Carbohydrates 2.5-4.8 3.5 7.3
0.1
1 Range of values obtained from chemical analyses of cultures
harvested at various stages of growth.
2 Stationary phase culture.
a Ben-Amotz et al. (1985).
of the fermenter culture was lower than the cul-

0.01 "-----'----'---"--------'------'-----'------'----'-----'---------'
tures grown in flasks. The high ash content of this
o 2 4 6 8 10 12 14 16 18 20 diatom is due to its highly silicified frustules. Car-
Days bohydrates form only a minor component of this
diatom.
-e- 1.2 L Fermenter --8- 2 L Flasks
Variation in chemical composition of N. con-
spicua with age is shown in Fig. 2. The actively
Fig. 1. Growth of Nitzschia conspicua based on chlorophyll-a growing culture (day 3) had the lowest lipid con-
concentration. tent (2.7% dry weight) and maximal lipids (14.1 %
N. conspicua flask cultures ended by day 5. Mter

day 5, a transition period followed before the %Dry weight
50.----------- ---------------------.
onset of the stationary phase on day 15. The cul-
tures grown in the 1.2 I fermenter had an expo-
nential phase lasting for 7 days and the station- 40
ary phase was attained from day 9 onwards.
Specific growth rates of the flask and fermenter
cultures based on chlorophyll-a concentration 30
were 0.51 and 0.48 d 1- 1 respectively. N. con-
spicua grown in the 1.2 I fermenter attained a
20
slightly higher biomass concentration (188 mg dry
weight 1-1) than the flask cultures (140-151 mg
dry weight 1- 1) during the stationary phase. 10
Chemical composition OL-----'----L---'----'------'----L---'----'-------'---·

o 2 4 6 8 10 12 14 16 18 20
Ash was the major chemical component of Days

N. conspicua in both culture systems (Table 1).
The fermenter culture of N. conspicua had similar --+- LipIds -&- Proteins
--H-- Carbohydrates --* Ash
chemical composition (except its lower carbohy-
drate content) than a species of Nitzschia (Table 1) Fig. 2. Chemical composition of Nitzschia conspicua at differ-
reported by Ben-Amotz et al. (1985). Ash content ent stages of growth.
36
dry weight) were produced on day 15 (stationary a big proportion (55.8%) of the dry mass was not
phase). Lipid accumulation during the stationary identified. Unknown fraction of the chemicals in
phase has been reported in other diatoms like N. conspicua at other stages of growth ranged from
Nitzschia palea (Opute, 1974), Biddulphia sinensis 21.4-35.4 % dry weight. This may be attributed to
(Volkman et al., 1980), Chaetoceros gracilis, incomplete extraction of the chemicals with the
Hantzschia sp. and Cyclotella sp. (Taguchi et al., methods employed. A major unidentified fraction
1987). amounted to 51.8 % dry weight was demonstrated
The cultures on day 5 had the highest ash con- in Nitzschia sp. (Ben-Amotz et al., 1985 and
tent. Ash content decreased after that and re- Table 1).
mained at a consistent level from day 13 until the
stationary phase.
Carbohydrate content remained low (2.5-4.8 % Fatty acid composition
dry weight) throughout the growth cycle of this
diatom. During the exponential phase, proteins Two major fatty acids of N. conspicua were 16:0
increased markedly from 3.0% dry weight on and 16:1 whilst other fatty acids detected were
day 3 to 11.0% dry weight on day 5 and did not 14:0, 18:1, 18:2, 18:3,20:4 and 20:5 (Table 2). In
differ much after this stage. general, almost equal proportions (1:1) of un sat-
The total chemical content of the 5 day-old urated and saturated fatty acids were exhibited.
cultures was only 44.2 % dry weight which means This ratio was much lower than values reported
Table 2. Fatty acid composition (in percentage) of N. conspicua and other species of Nitzschia.
Fatty acid Present study N. palea" N. alba b Reported values
N. conspicua * N. conspicua ** Nitzschia c N. angularis d Nitzschia d

(flask) (fermenter) sp. sp.
12:0 1.5
14:0 2.9-5.9 3.9 6.2 30.0 9.0 6.0 3.0
16:0 38.2-44.0 38.2 22.8 20.9 10.4 22.0 17.0
16:1 25.5-43.9 37.3 44.7 8.5 20.7 31.0 46.0
16:2 3.6 13.4 7.9 2.0 6.0
16:3 1.6 5.9 4.0 5.0
16:4 2.0
18:0 tr. 1.3 2.0
18: 1 0.7-2.7 1.1 2.5 23.6 3.6
18:2 0.5-3.3 1.3 tr. 4.7 1.7 2.0
18:3 1.7-7.5 5.6 tr. tr. 0.9 3.0
18:4 1.1 1.0
20:3 4.4
20:4 0.6-4.7 4.1 6.3 tr. 1.0 1.0
20:5 1.9-8.9 6.1 12.0 10.8 18.2 18.0 15.0
Total fatty acids 0.4-3.6 2.2
% dry weight
Unsaturated: saturated 0.9-1.2 1.2 2.4 1.6 1.8 1.8 2.7
Abbreviation: Tr. - Traces; -: not present.

* Cultures at different stages of growth were analysed and range of values are presented.
** Stationary phase culture.
REFERENCES: a) Opute (1974); b) Ackman et al. (1968).
c) Ben-Amotz et al. (1985); d) Kyle et al. (1988).
37
elsewhere (Table 2). EFA like 20:4 (AA) and 20:5 Variation in fatty acid composition with culture age
(EPA) constituted less than 10% of the total fatty
acids. Similar fatty acid constituents were shown Figure 3 shows the variation of total fatty acid
by the flask and fermenter cultures. content with culture age. Total fatty acid content
Table 2 clearly shows that fatty acid profiles of was lowest during the exponential phase and
various species of Nitzschia are different. How- maximal during the stationary phase.
ever, most of the species exhibit the typical dia- Effect of culture age on each of the fatty acid
tom fatty acid profiles (Borowitzka, 1988) with constituent is shown in Figs 4a-h. Higher pro-
the dominance of 16:0 and 16:1 and C18 fatty portions of 14:0 and 16:1 (Figs 4a and c) were
acids as the minor constituents. The non- produced by N. conspicua at the later stages of
photosynthetic diatom N. alba has rather unusual growth (from day 10 onwards) than earlier stages.
fatty acid composition with high amounts of 14:0 By contrast, 18:3 was at its maximum during the
and 18:1 (Ackman et al., 1968). exponential phase (day 3 and 5) and decreased
N. conspicua had a higher percentage of the fatty after this stage (Fig. 4f). More 18:1 was also syn-
acids as 20:4 compared with other reported spe- thesized during the exponential phase but de-
cies except N. palea (Table 2). But this content creased at the later stages (Fig. 4d). However,
was much lower compared to Porphyridium cruen- this fatty acid increased again on day 17 (station-
tum (the current commercial source of AA) which ary phase). On the other hand, 18:2 was maximal
has 36 % of its total fatty acids as 20:4 (Ahem on day 5 but decreased after this and was not
etal., 1983). EPA (20:5) content of N. conspicua detected on day 15 (Fig. 4e). Low quantity of this
was lower than other reported species. However, fatty acid was detected on day 17. No definite
manipulation of the culture conditions may im- trend was shown by 16:0 (varied between 38.2 to
prove production of these EFA.
Factors like salinity (Lee & Tan, 1988), culture
age (Opute, 1974), nitrogen and silicate levels
(Sriharan et al., 1990) and irradiance (Thompson 4
% Dry weight
- - - - --- - - ---
et al., 1990) are known to control the biosynthe-
sis of fatty acids in microalgae. For example,
Thompson et al. (1990) demonstrated that higher
proportion of 20:5 was produced at low levels of
3
irradiance in the marine diatoms Chaetoceros cal-
citrans, C. simplex, C. gracilis and Thalassiosira
pseudonana. Cycotella cryptica was reported to
produce more 20:5 under silicate-sufficiency than
silicate-deficiency (Roessler, 1988). 2
In general, autotrophic micro algae contain
about 3-20% dry weight as fatty acids (Ahem,
1984) which constitute 20-40 % of the total lipids
(Cohen, 1986). Total fatty acid contents ranged
from 5.8 to 8.2% dry weight (Hirano et al., 1990)
in some marine Chlorella. Biddulphia sinensis had
exceptionally low fatty acid content (1.4-2.1 %
dry weight) (Volkman et al., 1980). In the present o - -----r------.- r - , - , --r- ~'( -------r----
study, maximal fatty acid content was only 3.6% o 2 4 6 8 10 12 14 16 18 20

dry height (Table 2). Further manipulative stud- Days
ies are deemed worthwhile to be carried out in Fig. 3. Total fatty acid content of Nitzschia conspicua at dif-
order to increase fatty acid production. ferent stages of growth.
38
a)14:0 b)16:0
6 %Total fatty acids 50 %Total fatty acids
5
45
4
3 40
2
35
1
O~~~--~-r~r-'--.--~~~ 30~~-.--r-~~--~~~--~~
o 2 4 6 8 10 12 14 16 18 20 024 6 8 ro n M re ~ ~
Days Days
c)16:1 d)18:1
50 %Total fatty acids
~
40
30
20
10
O~'--.--r-~~--r-~~--~~ O~~~--~-r--r-~~--~-r~
o 2 4 6 8 10 12 14 16 18 20 o 2 4 6 8 ro n M re ~ ~
Days Days
e)18:2 f)18:3
3 6
2 4
O~~~--~-r--r-~~~~-r~ O~~~--~-r--r-~~--~-r~
o 246 8 ro n M re ~ ~ o 2 4 6 8 ro n M re ~ ~
Days Days
g)20:4 (AA) h)20:5 (EPA)

4 8
3 6
2 4
2
O~~~--~-r--r-~~--~-r~ O~'--.--r-~~--~~~--~~
o 2 4 6 8 10 12 14 16 18 20 o 246 8 ro n M re ~ ~
Days Days
Fig. 4. Variation in each fatty acid constituent of Nitzschia conspicua with culture age.
39
44.0% total fatty acids) in relation to culture age References

(Fig.4b).
Both 20:4 and 20:5 (Fig. 4g and h) were at their Ackman, R. G., C. S. Tocher & J. McLachlan, 1968. Marine
maximum on day 7, 2 days after the end of the phytoplankter fatty acids. J. Fish. Res. Bd. Can. 25: 1603-
1620.
exponential phase. Yields of 20:4 and 20:5 at this Ahern, T. J., 1984. Plant-derived catalysts and precursors for
stage of growth were 68.2 and 127.4 Ilg 1- I cul- use in prostaglandin synthesis. J.A.O.C.S. 61: 1754-1757.
ture respectively. Lower quantities of these two Ahern, T. A. J., S. Katoh & E. Sada, 1983. Arachidonic acid
fatty acids were produced during the stationary production by the red alga Porphyridium cruentum. Biotech.
Bioeng. 25: 157-170.
phase. EPA (20:5) content was reported to be
Ben-Amotz, A., T. G. Tornabene & W. H. Thomas, 1985.
higher during late exponential than the stationary Chemical profile of selected species of microalgae with
phase in Phaeodactylum tricornutum (Cooper emphasis on lipids. J. Phycol. 21: 72-81.
et a!., 1985). Bligh, E. G. & W. J. Dyer, 1959. A rapid method of total lipid
Effects of culture age on fatty acid composition extraction and purification. Can. J. Biochem. Physio\. 37:
911-917.
were also reported in other diatoms. For example,
Borowitzka, M. A., 1988. Fats, oils and hydrocarbons. In
C16 fatty acids increased while C18 fatty acids M. A. Borowitzka & L. J. Borowitzka (eds), Microalgal
decreased when cultures of Coscinodiscus eccen- Biotechnology. Cambridge University Press: 257-287.
tricus aged (Pugh, 1971). Apart from that, C20 Bradford, M. M., 1976. A rapid and sensitive method for the
fatty acids increased at the beginning of the sta- quantitation of microgram quantities of protein utilizing the
principle of protein-dye binding. Analyt. Biochem. 72: 248-
tionary phase after which they declined in C.
254.
eccentric us (Pugh, 1971). Increasing polyunsatu- Christie, W. W., 1989. Gas chromatography and lipids. The
rated fatty acids and decreasing monosaturates Oily Press of Scotland: 69-70.
and saturates due to aging was demonstrated in Cohen, Z., 1986. Products from microalgae. In A. Richmond
Thalassiosira pseudonana (Fisher & Schwarzen- (ed.), CRC Handbook of Microalgal Culture. CRC Press:
bach, 1978). 421-454.
Cooper, S. F., A. Battat, P. Marscot, M. Sylvestre & c.
Lalibert, 1985. Identification of antibacterial fatty acids
from Phaeodactylum tricornutum grown in dialysis culture.
Conclusion Microbios 42: 27-36.
DePauw, N. & G. Persoone, 1988. Microalgae for aqua-
culture. In M. A. Borowitzka & L. J. Borowitzka (eds),
Appreciable amounts of the EFA 20:4 and 20:5
Microalgal Biotechnology. Cambridge University Press:
were present in N. conspicua. Maximal contents 197-221.
of these fatty acids were detected on day 7. The Fisher, N. S. & R. P. Schwarzenbach, 1978. Fatty acid
fatty acids 16:0 and 16: 1 predominated in both dynamics in Thalassiosira pseudonana (Bacillariophyceae):
flask and fermenter cultures. Fatty acid profiles implications for physiological ecology. J. Phyco\. 14: 143-
150.
varied at different stages of growth. Lipids were
Harris, W. S., 1989. Fish oil and plasma lipid and lipoprotein
minimal during the exponential phase and maxi- metabolism in humans: a critical review. J. Lipid Res. 30:
mal during the stationary phase (day 15). Varia- 785-807.
tion in other chemicals like proteins, carbohy- Hirano, M., H. Mori, Y. Miura, N. Matsunaga, N. Nakamura
drates and ash with culture age was also & T. Matsunaga, 1991. gamma-Linolenic acid production
demonstrated. by micro algae. App\. Biochem. Biotech. 24/25: 183-191.
Hoshaw, R. W. & J. R. Rowoski, 1973. Methods for micro-
scopic algae. In J. R. Stein (ed.), Handbook of Phycologi-
cal Methods: Culture Methods and Growth Measurements.
Acknowledgements Cambridge University Press: 53-67.
Kochert, A. G., 1978. Carbohydrate determination by the
The above study was sponsored by a research phenol sulphuric acid method. In J. A. Hellbust and
J. S. Craigie (eds), Handbook of Phycological Methods -
grant (R & D 1/026/02) from the Malaysian Physiological and Biochemical Methods. Cambridge Uni-
government. The authors wish to thank Dr E. Y. versity Press: 95-97.
Haworth for identifying the diatom. Kyle, D., P. Behrens, S. Bingham, K. Arneti & D. Liberman,
40
1988. Microalgae as sources of EPA-containing oils. In Sriharan, S., D. Bagga & T. P. Sriharan, 1989. Environmen-
T. H. Applewhite (ed.), Proceedings of the World Confer- tal control of lipids and fatty acid production in the diatom
ence on Biotechnology for the Fats and Oils Industry. Navicula saprophila. Appl. Biochem. Biotech. 20/21: 281-
American Oil Chemists' Society, Champaign, Illinois: 117- 291.
121. Strickland, J. D. H. & T. R. Parsons, 1968. A practical hand-
Lee, Y. K. & H. M. Tan, 1988. Effect of temperature, light book of seawater analysis. Bull. Fish. Res. Bd. Canada 167,
intensity and dilution rate on the cellular composition of the 311 pp.
red alga Porphyridium cruentum in light-limited chemostat Taguchi, S., J. A. Hirata & E. A. Laws, 1987. Silicate defi-
culture. MIRCEN J. 4: 231-237. ciency and lipid synthesis of marine diatoms. J. Phycol. 23:
Opute, F. I., 1974. Studies on fat accumulation in Nitzschia 260-267.
palea Kutz. Ann. Bot. 38: 889-902. Thompson, P. A., P. J. Harrison & J. N. C. Whyte, 1990.
Pugh, P. R., 1971. Changes in the fatty acid composition of Influence of irradiance on the fatty acid composition of
Coscinodiscus eccentricus with culture age and salinity. Mar. phytoplankton. J. Phycol. 26: 278-288.
BioI. 11: 118-124. Volkman, J. K, G. Eglinton & E. D. S. Corner, 1980. Sterols
Roessler, P. G., 1988. Effects of silicon deficiency on lipid and fatty acids of the marine diatom Biddulphia sinensis.
composition and metabolism in the diatom Cyclotella cryp- Phytochemistry 19: 1809-1813.
tica. 24: 394-400. Wojenski, C. M., M. J. Silver & J. Walker, 1991. Eicosapen-
Sommer, T. R., W. T. Potts & N. M. Morrissy, 1990. Recent taenoic acid ethyl ester as an antithrombotic agent: com-
progress in the use of processed micro algae in aquaculture. parison to an extract of fish oil. Biochim. Biophys. Acta
Hydrobiologia 204/205 (Dev. Hydrobiol. 58): 435-443. 1081: 33-38.
Zoogeography and biodiversity of the freshwater fishes of

Southeast Asia
Mohd. Zakaria-Ismail
Department of Zoology, University of Malaya, 59100 Kuala Lumpur, Malaysia
Key words: zoogeographic regions, freshwater fishes, local extinction
Abstract
The ichthyofauna of the freshwater system of Southeast Asia is extremely diverse. A recent estimate of
about lOOO species is probably an understatement. More than lO new species are being added to the
list annually. The distribution pattern of the Southeast Asian freshwater fishes can be divided into five
zoogeographic regions. The first one is the Salween basin in Burma, with fishes mainly of the Indian
subcontinent origin such as Amblypharyngodon atkinsoni, Bangana almorae and Brachydanio jayarami.
The second zoogeographic area is the Mekong, Chao Phraya and Mae Khlong drainages which harbour
fishes typical of the mainland of Southeast Asia such as Acanthorhodeus deignani, Barbichthys nitidus and
Cirrhinus siamensis. The Malay Peninsula is the third region whose species composition is heavily in-
fluenced by the Siamese (such as Homaloptera smithi, Tuberoschistura baenzigeri and Botia beauforti) and
Indonesian (such as Botia hymenophysa, Luciocephalus pulcher and Parosphromenus deissneri) elements.
The islands of Sumatra, Borneo and Java are the fourth zoogeographic area of fish distribution. These
islands show a high degree of endemism, especially in fishes of the family Belontiidae. Finally, the
freshwater system of the Philippines is the last zoogeographic region of Southeast Asia. The area is
characterized by the presence of closely related species of the cyprinids especially in Lake Lanao.
Currently, high diversity of these freshwater fishes is being threatened by land development, such as
deforestation, road construction and land expansion for plantation. Recent studies in the Gombak River
basin show the extermination of 41 per cent of native fish species from 1969 to 1990. This is probably
due to the construction of highways, logging, as well as land clearing for agriculture.
Introduction graphic regions, with the publication of his clas-

sic work, The Geographical Distribution of Ani-
Southeast Asia, which includes Burma, the Indo- mals, in 1876.
Chinese and Malayan peninsulas, the Indo- Wallace (1860) proposed a hypothetical bound-
Malayan archipelago, and the Philippines, has ary between the Oriental and Australian faunas
been a focus of zoogeographic interest. Although when he said 'we may consider it established that
the foundations of biogeography can be traced the Strait of Lombok marks the limit and abruptly
to the writing of Georges Louis Leclerc, comte de separates two of the great zoological regions of
Buffon and Augustin Pyramus de Candolle the globe'. Later, Huxley (1868) named the
(Nelson, 1978), it was Alfred Russel Wallace, a boundary Wallace's Line and modified its limits
British naturalist and codiscover of the theory of on the basis of bird studies placing the Philippines
evolution, who elaborated the concept of zoogeo- east of the line within the Australian realm. As the
42
fauna of the region became better known, many of changes offish diversity and its probable cause
examples of animal groups crossing Wallace's line in Peninsular Malaysia is also presented.
were noted. For this reason, most zoogeographers
today do not take Wallace's line literally as an
exact boundary between the Oriental and Aus- Materials and methods
tralian fauna but rather a boundary of major fau-
nal break separating the relatively rich Oriental Observations and speculations on zoogeography
continental fauna from the depauperate Austra- and diversity of the freshwater fishes of Southeast
lian island fauna. This demarcation is apparent Asia are primarily based on the evidence pre-
with respect to the distribution of freshwater sented by various workers such as Anderson &
fishes. Collette (1991), Inger & Chin (1962), Kottelat
Myers (1949,1951) developed an ecological (1985, 1989, 1990), Ng & Lim (1990), Parenti
classification of freshwater fishes based on their (1989), Roberts (1980, 1982a, 1982b, 1986, 1989),
tolerance to salt water. The primary division fish Smith (1945), Taki (1974), Uwa & Magtoon
families are those whose members have little salt (1986) Weber & de Beaufort (1913, 1916), and
tolerance and are confined to freshwaters, al- Zakaria-Ismail (1987, 1989). Studies on changes
though salt tolerance of individuals within a fam- in the species composition in the Gombak River,
ily does vary. Secondary freshwater fishes are Peninsular Malaysia were carried out in 1985 and
salt-tolerant families, whose origin can be traced 1990 to provide evidence of the impact of land
to marine relatives, that live chiefly in freshwaters development on fish communities. Data of these
but sometimes enter the sea and can survive there studies were compared with the baseline infor-
for a limited time. The peripheral division fresh- mation of Bishop (1973). Fish diversity in the
water fishes are those that occur in freshwaters Kerling River, Peninsular Malaysia was deter-
but are actually semi-marine, or migratory, or de- mined by exhaustive sampling with an electro-
rived from marine families. Although these divi- shocker to observe some pattern of dominance of
sions are not absolute and do not apply strictly to the various groups of fish in a forest that was
whole taxonomic units, their usefulness in zoo- logged forty years ago.
geography cannot be denied.
Freshwater fishes of Southeast Asia are di-
verse. Unfortunately, up-to-date references on Results and discussion
these fishes are scarce, especially at the generic
revision. Recent generic revision works include There are many ways to classify the pattern of fish
Sontirat (1976) on Cyclocheilichthys, Karnasuta distribution of Southeast Asia. Kottelat (1989)
(1981) on Osteochilus, Roberts (1982a, 1982b, for example identified seven zoogeographic re-
1983, 1986) on Wallago, Chaca, Bagarius and gions for the freshwater fishes of Southeast Asian
Macrognathus, Roberts & Vidthayanon (1991) on mainland on the basis of the main river systems.
Pangasius, Parenti (1989) on Phallostetus, Siebert They are the Salween, Mae Khlong, Chao Phraya
(1991) on Acanthopsoides, and Burridge (1992) on and Mekong river drainages and Annam, south-
Acanthophthalmus. More revisional works are re- east Thailand and the Malay Peninsula. For the
quired, such as on Lobocheilos, Mystacoleucus, purpose of this paper, only five main zoogeo-
Neolissochilus and 'Puntius', before we can esti- graphic regions are used (Fig. 1). They are the
mate the number of species present in the area Salween basin in Burma, the Indo-Chinese pen-
with some degree of confidence. insula which includes the Mekong, Chao Phraya
The objective of this paper is to summarize and Mae Khlong river drainages in Thailand and
patterns of distribution and diversity of the fresh- Kampuchea, the Malay Peninsula, the Indo-
water fishes of Southeast Asia based on current Malayan archipelago, and the Philippines.
and historical data offish taxonomy. An example The Salween River, with a length and drainage
43
lissochilus, and genera whose systematics are

poorly known such as Puntius are found on both
sides of the border.
The Indo-Chinese zoogeographic region har-
bours fishes of typical of the mainland of South-
east Asia such as Acanthorhodeus deignani, Bar-
bichthys nitidus and Cirrhinus siamensis. This area
also shows a sizeable gap in the distribution of
some genera of catfishes. The angler catfish of the
genus Chaca comprises two species. Chaca chaca
occurs in Burma and also India, while Chaca ban-
kanensis is present only in the southern part of the
Malay Peninsula, Sumatra and Borneo. A simi-
lar pattern of distribution can also be observed in
Amblyceps mangois which is present in India and
the Malay Peninsula but not in the Indo-Chinese
peninsula.
The Malay Peninsula is the third zoogeographic
region. Species composition is heavily influenced
by Siamese as well as Indonesian elements. Many
100 110 120 Bornean and Sumatran species have their north-
ernmost distribution limit in the peninsula. As an
Fig. 1. Major zoogeographic regions of the freshwater fishes example, Luciocephalus pulcher is not known to
of Southeast Asia. ICP, Indo-Chinese peninsula; IMA, Indo- exist in the Indo-Chinese peninsula, but is com-
Malayan archipelago; M, Mindanao; MP, Malay Peninsula; monly found in the Malay Peninsula especially in
S, Salween basin.
blackwaters. Other species with a similar pattern
of distribution are Osteochilus spilurus, Parosph-
romenus deissneri and Sphaerichthys osph-
area of about 2800 km and 330 km 2 respectively, romenoides. Several Indonesian species are also
is the second largest river system in Southeast found only in the Malay Peninsula but not par-
Asia. This area has little in common in terms of ticularly restricted to the blackwaters; they are
the cyprinid fish composition with other zoogeo- Botia hymenophysa, Chaca bankanensis, Chela
graphic regions in Southeast Asia. The majority maassi, Crossocheilus langei, Doryichthys deokha-
of species in the area are of Indian Sub-continent toides, Mystus bimaculatus, Osteochilus enneaporos,
origin such as Amblypharyngodon atkinsoni, Ban- Oxygaster anomalura, and Vaillantella maassi.
gana almorae and Brachydanio jayarami. Several Siamese as well as Indian species have
In his recent study of the zoogeography of their southernmost distribution limit in the pen-
Southeast Asian mainland freshwater fishes, insula. These species include Amblyceps mangois,
Kottelat (1989) indicated that the Salween basin Botia beauforti, Garra cambodgiensis, Glyptothorax
is the eastern limit of the range of several genera callopterus, Homaloptera smithi, Mystacoleucus
widely distributed in India (e.g. Amblypharyn- chilopterus, Nemacheilus masyae, Paralaubuca
godon, Aspidoparia and Chagunius) as well as the typus, Puntioplites proctozysron, Rasbora borape-
western limit of the range of many Southeast tensis and Tuberoschistura baenzigeri. Thus, the
Asian genera (e.g. Albulichthys, Amblyrhynchich- Malay Peninsula has become an area of overlap
thys and Barbichthys). Only those genera that have for the Indian, Indo-Chinese and Indonesian
wide distribution is Asia such as Notopterus and fishes.
Mystus, montane forms such as Garra and Neo- Based on available literature, I estimate more
44
than 1000 species of fishes present in the fresh- exchange with other systems for a long period of
water system of Southeast Asia. The Salween time.
basin has some 150 species representing 77 gen- Great diversity of the ostariophysan fishes is
era, of which 55 % are shared with the lower found in the Indo-Chinese peninsula. Many stud-
Mekong, 60 % with the middle Mekong, 60 % with ies have shown that the area has the highest con-
the Chao Phraya, 87 % with the Irrawaddy, 77 % centration of this group of fishes especially the
with the Brahmaputra, and 71 % with the Ganges. Balitoridae (Kottelat, 1989), Sisoridae (Roberts,
Thus, the fish diversity is comparable to that of 1983) and Cobitidae (Sawada, 1982). Although
the Indian sub-continent. Two genera (2.5 %) are the Indo-Chinese peninsula is probably not the
endemic to the basin, and both are known only centre of origin of ostariophysan fishes (Regan,
in Inle Lake (Kottelat, 1989). 1922; Myers, 1967; Roberts, 1975), it is probably
The Indo-Chinese peninsula is probably the a place of rapid speciation of these fishes .
most diverse zoogeographic area in Southeast Although the Malay Peninsula is only slightly
Asia. The Mekong is the longest and has the larger than the Salween basin, its diversity is much
largest drainage area of all rivers in Southeast higher (260 vs. 150). According to Kottelat (1989)
Asia. High diversity of fishes in the Mekong is 15 % of the species are shared with the Salween
expected. More than 500 species (Smith, 1945; basin, 47 % with Chao Phraya, 44 % with the
Taki, 1975; Rainboth et al. , 1976; Kottelat, 1985; Mekong and 66% with the Indo-Malayan archi-
Mai & Nguyen, 1988) are present in the area. pelago. Peninsular Malaysia, the southern half of
Kottelat (1989) estimated some 300 species of the Malay Peninsula, harbours more than 200
freshwater fishes from the Indo-Chinese parts of species (Zakaria-Ismail, 1989) four of which
the Mekong and Chao Phraya basins alone. Of are endemic. They are Neolissochilus hendersoni,
the 103 genera found in the region, 67 % are in Poropuntius birtwistlei, P.smedleyi, Parosph-
common with the Sundaic forms, whereas only romenus nagyi and P. paludicola. These endemic
47 % of the genera have a strong affinity with the species are confined either to montane streams or
Salween basin. Forty-eight percent of the species blackwater swamps.
are endemic to both rivers. A high degree of en- The Indo-Malayan archipelago is the fourth
demism suggests that the basin has not had water zoogeographic region. The area, which includes
Poecilidae 9.6%
Hemiramphidae 9.6%
Clariidae 9.6%
Channidae 18.0%
9.6%
Belontiidae 9.6%
FIg. 2. Ichthyofauna composItIOn of the Pusu RJVer, Penmsular Malaysia.
45
the islands of Sumatra, Borneo and Java, has are more apparent throughout its middle and
more than 400 species. Some 290 species lower reaches (Rainboth, 1991).
(Roberts, 1989) inhabit the Kapuas River and it U sing the freshwater fishes of Thailand as an
is perhaps the most diverse fish community in the example (Smith, 1945), the cyprinoids consist of
region. High fish diversity is also seen in the more than 40 % of the total fauna. In the study of
Baram River, in which more than 115 species of the ichthyofauna of the Mekong River, Taki
fish were found (E. J. Crossman, pers. com.). The (1978) discovered that 49% of the fauna are cyp-
presence of endemic species of the perciforms rinoid fishes, and the percentage of cyprinoids
such as Belontiidae, Helostomatidae, Osphrone- increased progressively upstream, reaching 60%
midae, and Luciocephalidae suggest this area is a in the upper middle Mekong. In a study of the two
centre of specIation of this perciform group. small streams in Peninsular Malaysia, I observed
The freshwater habitat of Southeast Asia is a similar pattern of cyprinid fish diversity. In the
dominated by the cyprinid fish. Some 70 genera Pusu River, a tributary of the Gombak River that
of cyprinids are endemic to the area. Many other runs through old rubber estates, the fish commu-
genera found here have a few representatives nity is dominated by the cyprinid fishes followed
living outside the region. Some regions of South- by the catfishes, snakeheads and some other
east Asia, especially rivers of the Indo-Chinese groups (Fig. 2). An increase in the percentage of
peninsula that traverse different faunal regions the cyprinid fish was observed in the Kerling
have components of different cyprinid fauna. For River, a second order stream that runs through a
instance, the head waters of upper Mekong on the secondary forest which was logged 40 years ago
Tibetan Plateau has greater numbers of high but has since been left for forest regeneration.
Asian fauna. As it passes through Yunnan Pro- Here, more than half of the fish community are
VInce, components of the East Asian fauna domi- the cyprmid fish (Fig. 3).
nate, while components of Southeast Asian fauna The diversity of cyprinids on Mindanao, Phil-
Cyprinidae 53.8%
Cobitidae 7.7%
Hemiramphidae 7.7%
Channidae 7.7%
Bagridae
Clariidae 7.7%
Belontiidae 7.7%
Fig 3 lchthyofauna composItion of the Kerhng RIver, Penmsular Malaysia

46
Table 1. Fish species composltlon in the Gombak River has often been cited as an example of explosive
Selangor in 1969, 1985 and 1990. Data for 1969 is based on speciation in the last 10000 years (Myers, 1960).
Bishop (1973), and his sampling techniques as well as sam-
pling locations were repeated for the 1985 and 1990 studies.
Although there has been some doubt regarding
Presence and absence of each species is indicated by + and this pattern of speciation (Reid, 1980), there is no
- respectively. question that the fauna is endemic, but the age of
the lake, the actual size of the fauna, the exact
Species of fish 1969 1985 1990
distribution of the members, and the length of
Family Poeciliidae isolation are certainly suspect (Rainboth, 1991).
Poeci/ia reticulata + + + Land developments exert a tremendous pres-
Family Cyprinidae sure on biodiversity of aquatic organisms, par-
Hampala macrolepidota + ticularly fishes. Reduction in the number of
Mystacoleucus marginatus + + + species has always been the result of land devel-
Neolissochilus soroides + + +
Osteochilus hasseltii +
opments. As an example, Singapore has lost over
Poropuntius smedleyi + + + half of its freshwater fishes by 1960 (Alfred, 1961)
Puntius binotatus + + + due to urbanization. The best example in Malay-
Rasbora sumatrana + + + sia is observed in the Gombak River, a tributary
Family Bagridae
of the Klang river drainage. Ichthyofauna diver-
Mystus planiceps + +
Family Sisoridae sity in the river has been well documented
Glyptothorax major + + + (Bishop, 1973) in which 27 species were recorded
Glyptothorax platypogonoides + + in the survey. I resampled the river seven times in
Family Clariidae 1985 and five times 1990, repeating exactly
Clarias batrachus + + + Bishop's sites and method, and observed sub-
Clarias teijsmanni + + +
Family Siluridae stantial changes in fish structure of the river
Silurichthys hasseltii + + + (Table 1).
Family Hemiramphidae Data for 1985 indicate that seven species of
Dermogenys pussilus + fish have completely vanished from the river
Hemirhamphodon pogonognathus + + + which include two cyprinid fish (Hampala mac-
Family Synbranchidae
Monopterus albus + + +
rolepidota), a pipefish (Doryichthys deokhatoides),
Family Syngnathidae an anabantid (Anabas testudineus), a belontiid
Doryichthys deokhatoides + (Trichogaster trichopterus), a halfbeak (Dermog-
Family Anabantidae enys pussilus) and a mastacembelid (Mastacembe-
Anabas testudineus + Ius favus). These species represent about 25.9%
Family Belontiidae
Trichogaster trichopterus +
of the total fish species in the river. The 1990
Family Channidae survey indicates the disappearance of four more
Channa gachua + + + species (Glyptothorax platypogonoides, Mystus
Channa lucius + + + planiceps, Macrognathus maculatus and Channa
Channa melasoma + melasoma) from the river system. Thus far, about
Channa striata + + +
Family Mastacembelidae 40.7% of the total fish fauna of the Gombak River
Macrognathus maculatus + + have completely disappeared since 1969. It is
Mastacembelus favus + speculated that land developments around the
Gombak watershed for logging, agriculture, in-
dustry and housing, and highway construction
ippines is worth mentioning. The island is the contributed to the substantial reduction of fish
easternmost boundary of primary freshwater fish diversity of the Gombak River.
in Southeast Asia. Of the 23 cyprinid species The use of illegal methods to catch fish can also
known from the island, 19 species are endemic. become a major factor in reducing the diversity of
Fifteen of them are found in Lake Lanao, and this the freshwater fishes of Southeast Asia (Zakaria-
47
Ismail, 1991). Various kinds of insecticides and Mai, D. Y. & V. T. Nguyen, 1988. Species composition
and distribution of the freshwater fish fauna of southern
cyanide have been illegally used in many rivers
Vietnam. Hydrobiologia 160: 45-51.
that flow through remote, logged forested areas. Myers, G. S., 1949. Salt-tolerance of fresh-water fish groups
Recently, there is some evidence indicating the in relation to zoogeographical problems. Bijdr. Dierk. 28:
use of explosives in some of these rivers. Beyond 315-322.
any doubt, these indiscriminate methods coupled Myers, G. S., 1951. Fresh-water fishes and East Indian ZoO-
geography. Stanford Ichthyol. Bull. 4: 11-21.
with habitat destructions will have a negative im-
Myers, G. S., 1960. The endemic fish fauna of Lake Lanao
pact on freshwater fish communities in Southeast and the evolution of higher taxonomic categories. Evolutio~
Asia. 14: 323-333.
Myers, G. S., 1967. Zoogeographical evidence of the age of
the South Atlantic Ocean. In: Studies in tropical oceano-
Acknowledgements graphy, Miami, no. 5: 614-621.
Nelson, G., 1978. From Candolle to Croizat: comments on
I thanks R. J. Behnke, T. R. Roberts and L. R. the history of biogeography. J. Hist. Bio. 2: 269-305.
Parenti for valuable discussion and encourage- Ng, P. K. L. & K. K. P. Lim, 1990. Snakeheads (Pisces:
ment during the course of the study. Research Channidae): natural history, biology and economic impor-
tance. Essays in Zoology (special publication of the Bulletin
leading to this paper was supported by Ministry of the Raffles Museum): 127-152.
of Science and Environment grants R&D 04-07- Parenti, L. R., 1989. A phylogenetic revision of the phal-
04-048 to me at the University of Malaya. lostethid fishes (Atherinomorpha, Phallostethidae). Proc.
Calif. Acad. Sci. 46: 243-277.
Rainboth, W. J., 1991. Cyprinids of South East Asia. In
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Rainboth, W. J., K. F. Lagler & S. Sontirat, 1976. Maps of
Anderson, W. D. III & B. B. Collette, 1991. Revision of the
freshwater fish distribution in the lower Mekong basin.
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Sungei Gombak. Dr W. Junk Publishers, The Hague, 316.
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Huxley, T. H., 1868. On the classification and distribution of
the Alectoromorphae and Hereromorphae. Proc. zool. Soc. Roberts, T. R., 1980. A revision of the Asian mastacembelid
Lond.: 294-319. fish genus Macrognathus. Copeia 1980: 385-391.
Inger, R. F. & P. K. Chin, 1962. The freshwater fishes of Roberts, T. R., 1982a. Systematics and geographical distri-
bution of the Asian silurid genus Wallago, with a key to the
North Borneo. Fieldiana Zool. 45: 1-268.
Karnasuta, J., 1981. Revision of the southeastern Asiatic cyp- species. Copeia 1982: 890-894.
rinid genus Osteochilus GUnther. Unpublished Ph.D. dis- Roberts, T. R., 1982b. A revision of the South and Southeast
sertation, University of Alberta, 374 pp. Asian angler-catfishes (Chacidae). Copeia 1982: 895-901.
Roberts, T. R., 1983. Revision of the South and Southeast
Kottelat, M., 1985. Fresh-water fishes of Kampuchea - A
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Roberts, T. R., 1986. Systematic review of the Mastacembe-
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Dr Friedrich Pfeil, Pub., Munich, 262 pp.
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Acad. nat. Sci. Philad. 143: 97-144. Malay Archipelago. J. Proc. Linn. Soc. London 4: 172-184.
Sawada, Y., 1982. Phylogeny and zoogeography of the Wallace, A. R., 1876. The geographical distribution of ani-
superfamily Cobitoidea (Cyprinoidei, Cypriniformes). mals. Macmillan, London, 2 vols.
Mem. Fac. Fish. Hokkaido Univ. 28: 65-223. Weber, M. & de Beaufort, 1913. The fishes of the
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(Cypriniformes: Cobitidae), with the description of new phoidea, Ostariophysi: Siluroidea. Brill, Leiden,
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land. Bull. U.S. Natn. Mus. 188: xi + 622 pp. Indo-Australian archipelago. III. Ostariophysi: II Cyp-
Sontirat, S., 1976. Revision of the Southeastern Asiatic cyp- rinoidea, Apodes, Synbranchi. Brill, Leiden, xv + 455 pp.
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sertation, University of Michigan, Ann Arbor, 133 pp. freshwater fishes in South and Southeast Asia. Wallaceana
Taki, Y., 1974. Fishes of the Lao Mekong basin. U.S.A.I.D. 47: 6-9.
Mission to Laos, Agri. div., Vientiane vi + 232. Zakaria-Ismail, M., 1989. Systematics, zoogeography, and
Taki, Y., 1975. Geographic distribution of primary fresh- conservation of the freshwater fishes of Peninsular Malay-
water fishes in four principal areas of Southeast Asia. South sia. Unpublished Ph. D. dissertation, Colorado State Uni-
East Asian Stud. (Kyoto Univ.) 13: 200-214. versity, USA 473 pp.
Taki, Y., 1978. An analytical study of the fish fauna of the Zakaria-Ismail, M., 1991. Freshwater fishes in Peninsular
Mekong Basin as a biological production system in nature. Malaysia. In R. Kiew (ed.), The state of nature conserva-
Res. Inst. Evo!. Bio!. Tokyo Spec. Pub!. 1: 1-77. tion in Malaysia. Malayan Nature Society, Kuala Lumpur:
Uwa, H. & W. Magtoon, 1986. Description and karyotype of 115-119.
The damselfishes (Pisces: Osteichthyes: Pomacentridae) of Peninsular

Malaysia and Singapore: systematics, ecology and conservation
T. M. Sin, M. M. Teo, Peter K. L. Ng, L. M. Chou & H. W. Khoo

Department of Zoology, National University of Singapore, Kent Ridge, Singapore 0511,
Republic of Singapore
Key words: Pomacentridae, Malaysia, Singapore, taxonomy, ecology, conservation
Abstract
Damselfishes (Pomacentridae) are among the most common of reef fishes in tropical seas, comprising
320 species in all, with 268 species alone in the Indo-West Pacific regions. 15 genera (AbudefduJ,
Amblyglyphidodon, Amphiprion, Chelioprion, Chromis, Chrysiptera, Dascyllus, Pristotis, Dischistodus, Hemig-
lyphidodon, Neoglyphidodon, Neopomacentrus, Plectroglyphidodon, Pomacentrus, Stegastes) and at least 39
species inhabit the waters of Peninsular Malaysia and Singapore. They are important ecologically be-
cause many species are extremely abundant in reefs, and also economically, as several are collected in
large numbers for the lucrative aquarium trade. This demand has led to some fishermen using destructive
methods in obtaining the fishes, to the point that original standing populations may not be recoverable.
This threat poses a need for conservation of the reefs. The present study based primarily on fishes
collected from the waters of Singapore and specimens from the Zoological Reference Collection at the
National University of Singapore, will serve the dual purpose of establishing a current species list for
Malaysia and Singapore, and to update and clarify the local taxonomy of the group. An annotated list
(with localities) is provided of the known or reported species to aid studies on reefs.
Introduction Abudefduf Forsskal, 1775, and Chromis Cuvier,

1815. Tay (unpublished data) and Khoo & Tay
Damselfishes of the family Pomacentridae are one (1990) listed a total of 44 species in their study of
of the most abundant coral reef fishes found in the pomacentrids of Pulau Salu in Singapore. The
tropical seas, both in terms of actual numbers, as majority of these belong to five genera Amphip-
well as in diversity. Allen (1975a, 1991) estimated rion, Dascyllus, Chromis, Abudefduf and Pomacen-
that there are at least 320 species inhabiting the trus.
world's tropical seas, with a few in temperate and Studies on local pomacentrids have been on-
fresh waters. Of this number, 123 were reported going for many years in Singapore. The studies by
from Indonesia and 118 from Philippines, ac- S. W. Tay and one of the authors (KHW) began
counting for 38% and 36%, respectively, of the some 20 years ago, and some of the results remain
known species in the world (Allen, 1975a, 1991). unpublished. Over the past three years, studies of
De Beaufort (1940) recorded 25 species of Singapore reefs and its ichthyofauna have also
damselfishes in Singapore waters alone. These been initiated as part of the ASEAN-Australian
fishes are from four genera - Amphiprion Bloch & project (see Chou, 1989).
Schneider, 1801, Pomacentrus Lacepede, 1803, Recent changes in the classification of the
50
family (Allen, 1975a, 1991), together with rapid tion (ZRC) of the Department of Zoology, Na-
changes in the environment in which these fish tional University of Singapore. Fresh specimens
occupy (see Chuang, 1973), require an update were collected using antillean traps (mesh size
and reappraisal of the pomacentrid fauna for 2.54 em, dimensions 87 x 75 x 30 em). A total of
Peninsular Malaysia and Singapore. The ecology 210 specimens representing 15 genera and 39 spe-
and conservation of the local pomacentrid fauna cies were examined. New records since de Beau-
is discussed with regards to various threats. In- fort (1940) were collated for Singapore and Pen-
formation has been drawn freely from our insular Malaysia. Measurements were made as
colleagues' unpublished sources to provide as outlined diagrammatically in Fig. 1.
complete a picture as possible.
Taxonomic notes
Bleeker (1877) revised the classification of the
Specimens examined which were collected from subfamily Pomacentrinae, which at that time
the waters of Singapore and Peninsular Malaysia comprised mainly the genera Pomacentrus, Chro-
are deposited at the Zoological Reference Collec- mis, Dascyllus and Abudefduf His classification
length
soft dorsal fin spinous dorsal fin
ca peduncle
,,-----l- ---
.- ,. .- laterch. line
","
caudal f1']_---->.,~-
pectoral f i n ' - - + - - - preorbital
suborbital
anal f i n - - - + - - - f - - -
preopercle
pelvic fin
~--------------s~l~~--------------------
Fig. 1. External features of a typical pomacentrid (Chromis).
51
separated the genera Glyphisodon Lacepede, 1803, Allen (1975a) elevated two subgenera to genera
and Paraglyphidodon Bleeker, 1877, from Abu- (Amblyglyphidodon and Hemiglyphidodon), and in-
defduf. He also established several new subgenera troduced a new genus, Neopomacentrus. As the
in Glyphisodon, including Amblyglyphidodon, and names of some of the pomacentrid genera and
Hemiglyphidodon which occur in this region. He species have undergone substantial changes over
recognised Stegastes Jenyns, 1842, as a subgenus the years, the names might create some confu-
of Glyphisodon Fowler & Bean (1928), in their sion. These are reviewed below. The system of
review of Philippine fish, recognised Hemiglyphi- Allen (1975a, 1991) is quite effective, and has
dodon as a separate genus on the basis of the proved invaluable in our studies of the local
length and large number of gill rakers as com- pomacentrid fauna. The shapes of the fishes of
pared to Abudefduf The other subgenera were the various genera, although difficult to quantify,
synonymised with Abudefduf De Beaufort (1940) are nevertheless very useful, especially in the field.
agreed with most of the actions of Fowler & Bean
(1928), but synonymised Hemiglyphidodon with
Abudefduf He stated that the basis for his deci- Genera
sion was that the number of gill rakers was known
to vary greatly within genera. However, he ad- Amblyglyphidodon Bleeker, 1877
mitted that he had not examined the specimens of
the fishes. Type species
Bleeker (1877) also resurrected the genus Dis- Glyphisodon aureus Cuvier, 1830. Bleeker (1877)
chistodus Gill, 1863, previously synonymised with described the genus as a subgenus of Glyphisodon.
Pomacentrus (see later). This was generally disre- Allen (1975a) agreed with his basis for separating
garded by later workers until Allen (1975a) re- them, i.e. using the relatively orbicular shape of
vived it. Allen (1975a, 1991) revised the taxonomy constituent species, as well as the following mer-
of the pomacentrids and proposed a provisional istic features - Abudefduf pectoral rays (P): 18-20
generic framework based on, but modified from vs Amblyglyphidodon P: 15-18; Abudefduf gill
Bleeker's (1877) classification. rakers: 21-27 vs Amblyglyphidodon gill rakers:
25-30. For the local species, we have also found
Family: Pomacentridae that the shape of the dorsal and anal fins of Abu-
Subfamily: Amphiprioninae defduf and Amblyglyphidodon differ. The shape of
Genus: Amphiprion Bloch & Schneider, 1801 the soft anal fin of Abudefduf is more triangular,
Subfamily: Chrominae the tip being sharper, while that of Amblyglyphi-
Genus: Chromis Cuvier, 1814 dodon is more lobiform. The dorsal fin spines in
Genus: Dascyllus Cuvier, 1829 Abudefduf also do not project as far from the
Subfamily: Pomacentrinae membrane as compared to Amblyglyphidodon.
Genus: Abudefduf ForsskaI, 1775
Genus: Amblyglphidodon Bleeker, 1877
Genus: Cheiloprion Weber, 1913 Hemiglyphidodon Bleeker, 1877
Genus: Chrysiptera Swainson, 1839
Genus: Dischistodus Gill, 1863 Type species
Genus: Hemiglyphidodon Bleeker, 1877 Glyphisodon plagiometopon Bleeker, 1852. Bleeker
Genus: Neopomacentrus Allen, 1975 (1877) established Hemiglyphidodon (as a mono-
Genus: Neoglyphidodon Allen, 1991 typic subgenus) of Glyphisodon for only one spe-
Genus: Plectroglyphidodon Fowler & Ball, cies. Allen (1975a) essentially followed Bleeker's
1924 rationale, but elevated it to a full genus. The char-
Genus: Pomacentrus Lacepede, 1802 acters used ar.e a unique body shape and the fol-
Genus: Stegastes Jenyns, 1842 lowing meristic features: Abudefduf gill rakers:
52
21-27 vs Hemiglyphidodon: 65-85; Abudefduf Stegastes Jenyns, 1842

lateral line scales: 20-22 vs Hemiglyphidodon:
14-18. Type species
Stegastes imbricatus Jenyns, 1842. Emery & Allen
(1980) reinstated Stegastes as a senior synonym
Neoglyphidodon Allen, 1991 for Eupomacentrus Bleeker, 1877. They noted that
key characters of Stegastes correspond to those
Type species used by Bleeker (1877) for Eupomacentrus. Their
Glyphisodon melas Cuvier, 1830. Bleeker (1877) common characters are - 1. maxillary teeth unis-
established Paraglyphidodon as a separate genus erial, 2. rostrum scaled, 3. suborbital bones and
from Abudefduf Allen (1991) determined that the preoperculum denticulated, 4. subopercle den-
type species for Paraglyphidodon, P. oxycephalus ticulated, 5. opercle unarmed, 6. dorsal fin XII 16,
(Bleeker, 1877) actually belongs to the genus 7. anal rays 12. Stegastes is distinct from Abu-
Chrysiptera Swainson, 1839, and as such, Parag- defduf as the latter has no serrations on the sub-
lyphidodon becomes its junior synonym. Neo- orbitals, preopercle, or subopercle, has notched
glyphidodon was proposed by Allen (1991) as a teeth on the upper jaw, and the colour pattern is
substitute name for the other species Bleeker had composed of dark bands on a paler background.
placed in his Paraglyphidodon. Stegastes can be distinguished from Pomacentrus
by the presence of a biserial row of teeth on the
upperjawand 12(vs 13-14) dorsal spines (Emery
Neopomacentrus Allen, 1975 & Allen, 1980).
Type species Chrysiptera Swainson, 1839

Glyphisodon anabatoides Bleeker, 1847. Species
listed in this genus by Allen (1975a) were mostly Type species
transferred from Pomacentrus and Abudefduf His Glyphisodon cyaneus Quoy & Gaimard, 1824. This
separating it from Abudefduf was based on the genus was previously better known as Glyphido-
condition of the preopercle - the hind margin for dontops Bleeker, 1877. The type species of Glyphi-
Abudefduf is entire whilst that for Neopomacen- dodontops is Glyphisodon azureus Cuvier, 1830,
trus is either crenulate or serrate. For Pomacen- whereas that of Chrysiptera is Glyphisodon cyaneus
trus, the suborbital was at least naked anteriorly QUoy & Gaimard, 1824. It is however well known
whereas it is scaly in Neopomacentrus. that Glyphisodon azureus Cuvier, 1830, and Gly-
phisodon cyaneus Quoy & Gaimard, 1824, are
subjective synonyms of each other (see Allen,
Dischistodus Gill, 1863 1975). Glyphidodontops Bleeker, 1877, thus also
becomes a junior synonym of Chrysiptera Swain-
Type species son, 1839. All species previously referred to
Pomacentrus fasciatus Cuvier & Valenciennes, Glyphidodontops have been referred to Chrysiptera
1830. The separation of the genus Dischistodus by (see Allen, 1987, 1991).
Allen (1975a) is based on the prominent notch
observed between the preorbital and suborbital in
Species
Pomacentrus but absent in Dischistodus. Also, the
snout is scaled in Pomacentrus up to the level of Abudefduf sexfasciatus Lacepede, 1801
the nostrils and beyond, but mostly naked in Dis-
chistodus, extending at most to just beyond the This species was formerly better known as
orbits. Abudefduf coelestinus (Cuvier, 1830). Allen et al.
(1978), on examining the syntypes of the original
53
Table 1. Distribution of the species of the family Pomacen- regard them as subspecies of each other (see
tridae in Singapore and Malaysia. Allen, 1991). The two taxa can be separated on
Species name Malaysia Singapore the basis of the position of the fourth black bar.
This bar is positioned directly behind the last
Abudefduf bengalensis
* * dorsal spine in the Atlantic species but well be-
Abudefduf notatus hind the last dorsal spine in the Indo-Pacific spe-
*
Abudefduf sexfasciatus * * cies (Allen, 1991). In A. vaigiensis, this black bar
Abudefduf sordidus *
Abudefduf vaigiensis is below the middle part of the soft dorsal fin.
Amblyglyphidodon curacao
* *
* *
Amphiprion clarkii
*
Amphiprion frenatus Plectroglyphidodon lacrymatus (Quoy & Gaimard,
* *
Amphiprion ocellaris *
* 1825)
Amphiprion perideraion
* *
Amphiprion polymnus
* This species was originally known as Glyphisodon
Chelioprion labiatus
*
Chromis atripectoralis
* * lacrymatus. Bleeker (1877) placed this species in
Chromis cinerascens * Stegastes, a subgenus of Glyphisodon. Whitley
Chromis viridis
* * (1929) recognised the species lacrymatus as dif-
Chromis weberi * *
Chrysiptera unimaculatus fering from typical Stegastes in having thick,
* plaited lips and a relatively naked snout. In
Dascyllus reticulatus
*
Dascyllus trimaculatus
* * Stegastes s. str. (fide Whitley, 1929), scales extend
Dischistodus chrysopoecilus up to the level of the nostrils whereas in Plectro-
*
Dischistodus fasciatus
* * glyphidodon, the snout is scaled up to the anterior
Dischistodus melanotus
* edge of the orbits. Whitley consequently proposed
Dischistodus perspecillatus
* * a new genus, Neostegastes for lacrymatus. As both
Dischistodus prosopotaenia
* *
Hemiglyphidodon plagiometopon
* Neostegastes and Plectroglyphidodon have the same
Neoglyphidodon melas generic characters, Neostegastes Whitley, 1929,
* *
Neoglyphidodon nigroris
* * becomes a subjective junior synonym of Plectro-
Neopomacentrus anabatoides *
Neopomacentrus jilamentosus glyphidodon Fowler & Ball, 1924 (see Allen,
* * 1975a).
Plectroglyphidodon lacrymatus
Pomacentrus alexanderae
*
* *
Pomacentrus chrysurus * *
Pomacentrus grammorhynchus
* Results
Pomacentrus littoralis * *
Pomacentrus moluccensis
*
Pomacentrus philippinus
* The species from Peninsula Malaysia and
Pomacentrus tripunctatus
* * Singapore of which specimens have been exam-
Pristotis jerdoni
* ined are listed in Table 1. In addition to these, de
Stegastes obreptus
* *
Beaufort (1940) has listed 10 other species as
being present here. These are:
Glyphisodon coelestinus Cuvier, 1830, found that
they corresponded extremely well with the de- Amblyglyphidodon leucogaster Bleeker, 1847
scription for A. sexfasciatus. Abudefduf sexfascia- Amphiprion ephippium Bloch, 1790
tus is thus the senior synonym. Amphiprion melanopus Bleeker, 1852
Chromis xanthurus Bleeker, 1854
Abudefduf vaigiensis (Quoy & Gaimard, 1825) Dischistodus melanotus Bleeker, 1858
Stegastes lividus Bloch & Schneider, 1801
This taxon closely resembles the Atlantic species, Neopomacentrus violascens Bleeker, 1848
A. saxatilis Linnaeus, 1758, and some workers Pomacentrus amboinensis Bleeker, 1868
54
Pomacentrus breviceps Schlegel & MUller, The 'rock pool zone' is at the high water tide
1839 level. It is totally submerged during high spring
Pomacentrus melanopterus Bleeker, 1852 tides. The characteristic algae found are Caulerpa
sp., Chaetophora sp. and Padina sp.
Although no specimens of the above species have
The 'rocky reef flat zone' making up of loose
been collected, various unconfirmed sightings of
rocks, lying at the upper inter-tidal region. The
some of them have been made by several of our
sandy reef flat consisting of hard and soft corals,
colleagues. These are Amblyglyphidodon leuco-
lying between the low water neap tide to the mar-
gaster, Amphiprion ephippium, Dischistodus melano-
gin of the reef edge. These first two zones may be
tus, and Neoglyphidodon melas (Chou, 1989). Tay,
absent in some fringing reefs in Singapore due to
in his unpublished M.Sc. thesis and Khoo & Tay
land reclamation.
(1990), listed five species not in de Beaufort's list
The 'reef edge zone' which lies at the lower
or the ZRC. These (nomenclature corrected by
inter-tidal. This zone is completely exposed once
G. Allen, in litt.) are:
every fortnight at the lowest water spring tides. At
Dascyllus aruanus Linnaeus, 1758 low tides it is covered by a shallow layer of water.
Dascyllus melanurus Bleeker, 1854 A wide variety of corals inhabit this area and its
Neopomacentrus anabatoides Bleeker, 1847 growth therefore slightly elevates this zone as
(as Neopomacentrus taeniurus, nec Bleeker, compared to the sandy reef zone.
1856) The 'reef slope zone' is the subtidal portion of
Pomacentrus chrysurus Cuvier, 1830 (syn- the reef. Even at low tides, the zone is never ex-
onym of Neopomacentrus rhodonotus posed. Characteristic of this zone are the scler-
Bleeker, 1877) actinian (hard) and alcyonarian (soft) corals, the
Pomacentrus coelestis Jordan & Starks, 1901 sea-fans (gorgonians), the sea-urchins (Diadema
setosum) and the fire-ferns (Aglaophenia cupres-
There are some minor problems with Tay's spe-
sina). The majority of pomacentrid species stud-
cies lists. He did not state where his specimens
ied are solitary and territorial in behaviour, with
were collected from, although it was implicit from
three species inhabiting the rock pool zone, eight
the title of his thesis that all were supposedly from
occupying the rock patch, seven in the sandy
Singapore. In the ZRC however, are specimens
patch, and 17 each in both the reef edge and reef
of some of these species (e.g. Dischistodus mel-
slope areas.
anotus) which were all collected from Malaysia.
Abudefduf spp. range from the rock pool to the
No Singapore specimens are extant. Tay was
reef slope, with A. bengalensis inhabiting the en-
known to have made many collections in Malay-
tire range. The other two species, A. vaigiensis
sia, and it seems possible that some of the Sin-
and A. sexfasciatus were restricted to the reef edge
gapore records have been incorrectly based on
and reef slope only (Khoo & Tay, 1990). These
Malaysian specimens.
fish were often observed to be schooling above
coral heads, especially Acropora, and used the
crevices for shelter from larger organisms when
Habitats of Damselfishes found in Peninsular threatened.
Malaysia and Singapore Dischistodus spp. also occupied a similar range,
with D.fasciatus occupying the rock pool and
Zonation of the reefs in Singapore and Malaysia rock patch region, D. prosopotaenia inhabiting the
were defined based on descriptions given by Khoo stretch from the rock patch to the reef slope, while
& Tay (1990), with modifications. This system the territory covered by D. chrysopoecilus ex-
differs from Allen's (1975a, 1991) categorisation tended from the rock pool to the slope. These fish
in that it is more specific and useful for local were commonly observed to be aggressive, de-
waters. fending individual territory against other fish.
55
Pomacentrus spp. occupied a lesser expanse of Conservation and discussion

habitat, ranging form the rock patch to the reef
slope. Only P. littoralis was observed to inhabit The pomacentrid species in Singapore range
the entire range. Most of the species were en- along the reef profile from the sublittoral zone to
countered in deeper water, at the reef edge and the reef slope. Land reclamation in Singapore
the slope. These include P. alexanderae, P. gram- has, between 1965-1985, added on 40 km 2 to
morhynchus, and P. moluccensis. Juvenile speci- the total land area (Chia et al., 1988). Much of
mens of these were observed to aggregate in this was carried out on the southern islands of
schools near structure. This was commonly ob- Singapore, thereby affecting the total area of reef
served at Pulau Semakau in the southern waters available to these fish. Proposals have been made
of Singapore, where schools ofjuveniles were seen that space, rather than food, is the primary lim-
congregating around disused sewage pipes. iting factor of reef fish abundance (Sale, 1978).
Amphiprion ocellaris and A. melanopus were ob- Reclamation would therefore directly quantita-
served to inhabit mainly the reef edge and the reef tively reduce the amount of ecological niches
slope in symbiotic relationship with large anemo- available to all reef fish, including pomacentrids.
nes (Stoichactis sp. and Actinia sp.). The relation- Other destruction of the reef arises from collec-
ship between the anemone fishes and the host tion, either by recreational divers, or by fisher-
anemones has been the subject of many studies men, who have found a demand for corals from
by several researchers. The general consensus is local aquariums (e.g. see Straits Times, Sin-
that the relationship is one of mutual protection gapore, 28 Sept. 1991).
(Allen, 1975b, 1991, 1980; Uchida, 1975). Generally, reclamation processes in Singapore
Dascyllus trimaculatus similarly occupied the have resulted in the destruction of most, if not all,
above range, developing a commensal relation- of the reef fiats. This could have led to the elimi-
ship with stoichactid sea-anemones in the young nation of certain species of coral found almost
stage. No such observations were made for the exclusively on reef fiats, such as Acropora spp.,
adult stages (Uchida, 1975). These fishes oc- which aside from being a major constituent of
curred both in small and large aggregations. Ju- territories for adult damselfishes such as Dischis-
veniles of the fish were also commonly observed todus spp., are a refuge for juveniles of many other
to seek refuge amongst the spines of the common pomacentrids (see Sano et al., 1984). These
sea urchin, Diadema spp. authors found that there were substantially fewer
Neoglyphidodon nigroris inhabits a similar range species and lower populations of pomacentrids
of reef edge and reef slope, browsing on algae on coral reefs with dead coral compared to the
encrusted on dead coral heads and rocks. It is number in live coral colonies. Chelioprion labiatus
sympatric with other dark-coloured species of was found to be completely absent from colonies
damselfishes (Dischistodus chrysopoecilus, Poma- with dead coral while being abundant in colonies
centf'Us littoralis, P. albimaculus), appearing to with live coral. A few exceptional species such as
share the same resources in terms of physical Pomacentrus littoralis, an omnivore, were encoun-
space and food (Khoo & Tay, 1985). tered in greater numbers over dead coral colonies.
Amblyglyphidodon curacao occupied similar Non-associated species such as Chromis spp.,
ranges of reef edge and reef slope. It is frequently also aggregated in larger numbers over live coral
observed to aggregate in large numbers over the colonies.
staghorn coral (Acropora). The substrate eliminated as a result of recla-
Hemiglyphidodon plagiometopon are solitary, re- mation may also have caused a change in the
stricted only to the deeper reef slope zone. They relative abundance of the respective species
were not observed to form associations with present, as well as eliminating others as a large
corals as adults. proportion of the pomacentrids in Singapore
often require crevices for refuge. The apparent
56
loss of species may also have been due to disrup- important role as components in the coral reef
tions in the food chain, and loss of host organ- ecosystem.
isms such as the sand-dwelling anemones Actinia To bring the present situation into perspective,
spp., which are symbiotic with many Amphiprion de Beaufort (1940) listed 24 species of po mac en-
species (Allen, 1972). trids as being present in Singapore, including two
Many of the pomacentrids found in regional species for which Singapore is the type locality.
waters are economically valuable in the aquarium Ten of the species for which we have not found
trade, especially the anemone-fishes, Amphiprion. specimens are part of de Beaufort's list. The
Some other brightly coloured pomacentrids are present study also records several species which
also popular and commonly available to aquar- are new for Peninsular Malaysia and Singapore
ists. These include the genera Dascyllus, Chromis, (see Table 2). There has been a surprising dearth
Pomacentrus and Abudefduf While the exact ex- of literature on pomacentrid systematics in this
tent and range of the collection of these fishes region since de Beaufort's work. The unpublished
from local waters is not known, it is clear that work of Tay and that of Khoo & Tay (1990)
these fishes are being depleted rapidly in the re- are the only local reports on the Malaysian and
gion. The marine aquarium market in the Philip- Singapore fauna. A search through the Zoologi-
pines for example, has expanded from exporting cal Records for the last 50 years have not uncov-
33391 kg of fish in 1968 to export in excess of ered any literature specifically referring to the
1800000 kg of fish in 1979 to meet the expand- Indo-Malayan pomacentrid fauna. The studies
ing markets abroad, especially in America and by Gerald Allen, John Randall and their associ-
Europe (Albadejo & Corpuz, 1981). Harvest of ates, mainly on the fauna of other parts still form
this valuable resource cannot go on unchecked the basis of all taxonomic knowledge about the
especially if destructive fishing methods are em- group for this area.
ployed as the recovery rate of the fish may not be The species diversity of the region is certainly
able to compensate for the rate of removal. Also, not depauperate. This is already evident by the
size limits should be imposed on collection, so large number of species found in the region by
that only juveniles are taken, allowing established past workers, even without the sophisticated
adults to continue reproducing. apparatus and equipment available to researchers
Legislation of collection and harvesting of the at present. That the area is very species-rich is
fish should be made a priority amongst tropical demonstrated by the findings of Allen (1991), who
countries. Steps should be taken to check the lists a total of 130 pomacentrids known from
over-harvesting of these fish, or control the har- Southeast Asia, particularly from Indonesia/
vest by setting size and catch limits, implement Malaysia, Philippines, Singapore and Thailand.
closed seasons, as well as clamp down on de- These countries are also the type localities for 70
structive collecting methods. The collection of of the species present, some of which are type
coral, anemones and other such sea organisms species for their respective genera. Of these, five
which provide refuge for, or form close associa- Indonesian and one Malaysian species were only
tions with damselfishes should also be closely very recently described by Allen (1991).
monitored. Perhaps more marine parks should be That studies on Malaysian pomacentrids are
set up, functioning as a nature reserve where these lacking is an unfortunate fact. A valuable resource
fish and other reef organism are protected. The is being overlooked and underestimated. Further-
establishment of a marine reserve as in Pulau more, pomacentrid resources are being rapidly
Redang, Trengganu, Malaysia, is very timely, and endangered due to illegal harvesting and habitat
more such established, especially in the coral reef- destruction. The potential for research in this field
rich islands off Pahang and Johor. Fishes like is evident. There have been many sightings of
pomacentrids are not only ornamental on the unidentifiable species of pomacentrids by trained
many reefs in the region, they serve a vastly more divers that frequent local waters. There are sight
57
Table 2. New records of species of the family Pomacentridae long to Neopomacentrus, while the other appears
in Singapore and Malaysia.
to be related to Pomacentrus alexanderae. At
Species name Malaysia Singapore present, the rapid urbanisation of Peninsular
Malaysia and Singapore is posing a threat to coral
Abudefduf bengalensis * reefs here. There is therefore a very urgent need
Abudefduf notatus not only for taxonomic and ecological studies of
*
Abudefduf sexfasciatus * * these fish, but also increased emphasis on their
Abudefduf sordidus
Abudefduf vaigiensis
* conservation. The conservation of the natural reef
Amblyglyphidodon curacao
* habitat is an obvious priority.
* *
Amphiprion clarkii
Amphiprion frenatus
*
*
Amphiprion ocellaris * * Acknowledgements
Amphiprion perideraion
Amphiprion polymnus
* *
Chelioprion labiatus
* The authors are grateful to Dr S. W. Tay, Ms
*
Chromis atripectoralis * * Grace S. Y. Lim, Mr C. B. Leng and Mr Kelvin
Chromis cinerascens Lim for permission to use their unpublished data
*
Chromis viridis * in formulating the present paper. Dr Gerald Allen
Chromis weberi
Chrysiptera unimaculatus
* (Western Australian Museum) and Dr John
Dascyllus reticulatus
* Randall (Bernice P. Bishop Museum) have been
*
Dascyllus trimaculatus
* * most encouraging with their support, obtaining
Dischistodus chrysopoecilus some literature and quick answers to many ques-
Dischistodus fasciatus
*
* tions. They also read an early version of this paper
Dischistodus melanotus
Dischistodus perspecillatus
* and provided many very useful suggestions which
* * we have incorporated into the present paper to
Dischistodus prosopotaenia *
Hemiglyphidodon plagiometopon * the best of our abilities. Mrs C. M. Yang was
Neoglyphidodon melas * most forthcoming in permitting us free access to
Neoglyphidodon nigroris * the collections in the ZRC. This study forms part
Neopomacentrus anabatoides
Neopomacentrus jilamentosus
* of the Asean-Australia Economic Co-operation
Plectroglyphidodon lacrymatus
* * Program: Living Resources in Coastal Areas
*
Pomacentrus alexanderae * * Project (Phase I). This study has also been par-
Pomacentrus chrysurus * * tially supported by a Research Grant RP 910410
Pomacentrus grammorhynchus
* to PKLN from the National University of Sin-
Pomacentrus littoralis *
Pomacentrus moluccensis gapore.
Pomacentrus philippinus
*
*
Pomacentrus tripunctatus * * References
Pristotis jerdoni
*
Stegastes obreptus
* * Albaladejo, v. D & v. T. Corpuz, 1981. A market study of
aquarium fish industry of the Philippines: an assessment of
the growth and the mechanics of the trade. Proc. Fourth
Intern. Coral Reef Symp., Manila 1: 75-81.
records of at least one undescribed species which Allen, G. R., 1975a. Damselfishes of the South Seas. T.F.H
was first obtained by Tay (unpublished data) and Pub!. Inc., Neptune City, New Jersey, USA, 237 pp.
subsequently referred to G. Allen in 1977 for Allen, G. R., 1975b. Anemonefishes and their Amazing
study (pers. comm.). This species, to the authors' Partnership. Aust. Nat. Hist. 18: 274-278.
knowledge, remains undescribed. Fresh speci- Allen, G. R., M. L. Bauchot& M. Desoutter, 1978. The status
of Abudefduf sexfasciatus (Lacepede), a pomacentrid fish
mens however, are still wanting. There are at least, from the Indo-west Pacific. Copeia 1978: 328-330.
two other species of pomacentrids in local waters Allen, G. R., 1980. Anemonefishes of the World. Aquarium
that are, as yet, not recorded. One appears to be- Systems, Mentor, Ohio, USA, 103 pp.
58
Allen, G. R., 1987. Chrysiptera sinclaM, a new species of Fowler, H. W. & B. A. Bean, 1928. The fishes of the families
damselfish from the tropical western Pacific Ocean. Revue Pomacentridae, Labridae, and Callyodontidae, collected be
Aquario!. 13: 107-110. the United States Bureau of Fisheries Steamer 'Albatross',
Allen, G. R., 1991. Damselfishes of the World. Aquarium chiefly in the Philippine seas and adjacent waters. Bull.
Systems, Mentor, Ohio, USA, 271 pp. U.S. Nat. Mus. 100: 1-525.
Bleeker, P., 1877. Atlas Ichthyologique des Indes Orientales Lim, G. S. Y., L. M. Chou & L. S. Chia, 1990. The Biologi-
Neerlandaises, publie sous les auspices du Government cal Communities of the Coral Reefs of Singapore With
Colonial Neerlandais. Amsterdam. MUller. 1862-77, 9 Emphasis on Reef Fishes and Hard Corals. The Second
Vols., 420 pp. Asian Fisheries Forum. Pub!. Asian Fisheries Society,
Carpenter, K. E., 1981. The influence of substrate structure Manila: 381-384. Khoo, H. W. & S. W. Tay, 1990. Coral
on the local abundance and diversity of Philippine reef Reef Fishes of Singapore. In L. M. Chou and P. K. L. Ng
fishes. Proc. Fourth Intern. Coral Reef Symp., Manila 2: (eds), Essays in Zoology. Dept. Zoo!., Natn. Univ.
497-502. Singapore, Singapore: 127-152.
Chia L. S., K. Habibullah & L. M. Chou, 1988. The coastal Randall, J. E., 1985. Chromis viridis (Cuvier, 1830), the correct
environment profile of Singapore. International Center for name for the Indo-Pacific damselfish previously known as
Living Aquatic Resources Management (ICLARM) Tech- C. caerulea (Cuvier, 1830) (Pomacentridae). Cybium 9:
nical Reports 21. Pub!. International Center for Living 411-413.
Aquatic Resources Management, Manila, Philippines, Sale, P. F., 1978. Coexistence of coral reef fishes - a lottery
92pp. for living space. Envir. Bio!. Fish. 3: 85-102.
Chou, L. M. (ed.), 1989. Final report of the Singapore Com- Sano, M., M. Shimizu & Y. Nose, 1984. Changes in structure
ponent of the Asean-Australia Project on Living Resources of coral reef fish communities by destruction of hermatypic
in Coastal Areas Phase One: July 1985-June 1989, corals: Observation and experimental views. Pacific Sci. 38:
Singapore: 15-21. 51-79.
De Beaufort, L. F., 1940. The Fishes of the Indo-Australian Uchida, H., 1975. Symbiosis between sea-anemones and
Archipelago. E. J. Brill, Leiden: 326-472. fishes. Marine Flowers, 1975: 57-60.
Emery, A. R. & G. R. Allen, 1980. Stegastes; A senior syn- Whitley, G. P., 1929. Some fishes of the order Amphriprioni-
onym for the damselfish genus Eupomacentrus; osteological formes. Mem. Qld. Mus. 9: 207-246.
and other evidence, with comments on other genera. Rec.
West. Aust. Mus. 8: 1-199.
<£) 1994 Kluwer Academic Publishers.
The systematics and ecology of snakeheads (Pisces: Channidae) in

Peninsular Malaysia and Singapore
P. G. Lee & Peter K. L. Ng

Key words: Malaysian Channidae, systematics, ecology, identification key
Abstract
Seven species of snakeheads (Channidae) are known from Peninsular Malaysia and Singapore, viz.
Channa bankanensis, C. gachua, C. lucius, C. marulioides, C. melasoma, C. micropeltes and C. striata. Up-
to-date distribution maps of each species are presented, including new records. Their systematics is
reviewed and partially revised. The taxonomic status of C. marulioides and C. melanoptera is clarified.
Specimens from Peninsular Malaysia identified as C. melanoptera sensu Weber & de Beaufort, 1922,
proved to be the adult form of C. marulioides s.str. The real C. melanoptera appears to be restricted to
Borneo and possibly Sumatra. The life history of a blackwater species C. bankanensis is also documented,
with regards to the morphological and colour-pattern changes associated with growth. An updated key
to the seven species based on morphometric measurements and meristic counts is presented.
Introduction records in the peninsula. Records or taxonomic

notes that include information on the snakeheads
Literature on the systematics, ecology and con- of the region through the years are numerous. Not
servation of snakeheads (Channidae) in Peninsu- unexpectedly, in the course of these studies, vari-
lar Malaysia and Singapore is very scattered. In ous incorrect identifications have occurred, in-
terms of systematics studies, two subspecies were cluding C. bistriatus for C. lucius (see Tweedie,
described from this region. Peters (1868) de- 1936, 1950; Herre & Myers, 1937; Fowler, 1938;
scribed Ophiocephalus guachua var. malaccensis Alfred, 1964), C. melanopterus for C. marulioides
from Kranji River, Singapore, but this is now (see Tweedie, 1940, 1950; Zakaria-Ismail, 1984;
regarded as a junior synonym of Channa gachua Ng & Lim, 1990b), C. orientalis for C. gachua (see
Hamilton-Buchanan, 1822 (see Weber & de Fowler, 1938; Kottelat, 1989), C. pleurophthalmus
Beaufort, 1922; Ng & Lim, 1990b). Gianferrari for C. lucius (see Tweedie, 1940), C. punctatus for
(1930) described Ophiocephalus striatus var. quala- C. melasoma (see Fowler, 1938), C. striata for
mudensis from Malacca, but the status of this C. bankanensis (see Davis & Abdullah, 1989) and
subspecies is unclear. Besides these works, the a doubtful record of C. afJinis/ = punctatus (see
study of channid systematics, has in most cases, Karoli, 1882 in Weber & de Beaufort, 1922).
been limited to establishing the correct identities In this paper, the information from these and
of described species, and their distribution other sources are collated with new data so as to
60
present a useful summary of the status of the and origin of anal fin; postorbital head length -
family in Peninsular Malaysia and Singapore to distance from posterior edge of eye to posterior
facilitate future studies. Although Ng & Lim edge of operculum; predorsal scales - number of
(1990b) have a useful key to the species, it is scales on top of head from origin of dorsal fin to
based mainly on body patterns and colours. This tip of snout including cephalic shields; preoper-
is useful for identification of live or freshly-killed cular length - distance from posterior edge of eye
specimens, but might prove difficult to use for to posterior edge of preoperculum; preopercular
some museum specimens which are extremely scales - number of scales from posterior edge of
bleached or faded. A key that relies on morpho- eye to posterior edge of preoperculum.
metric and meristic characters is thus constructed Distribution records were determined from lit-
to augment that by Ng & Lim (1990b) for the erature as well as museum specimens we have
identification of such specimens. examined. Other than those already cited in the
Specific references to the ecology of the various text, distribution records were also obtained from
snakehead species in Malaysia are few. Except Alfred (1961a, 1961b, 1963, 1966), Ali & Kather-
for Soong (1949) which gives a brief life-history gany (1987), Duncker (1904), Johnson (1960),
description of C. striata (Bloch, 1793), most stud- Lim et at. (1990) and Zakaria-Ismail (1987).
ies conducted in the region address some ecologi- Definite localities cited were plotted on maps as
cal problem and bring in snakeheads only as part accurately as possible with reference to maps and
of the data. Some discuss the distribution of gazetteers. Specimens examined are contained
freshwater fishes in the region and mention in the Zoological Reference Collection (ZRC),
snakeheads. Others place emphasis on the aqua- Department of Zoology, National University of
culture and fisheries aspects. Alfred (1968) con- Singapore. Localities which were ambiguous or
sidered one snakehead species (c. gachua as cannot be determined precisely were excluded.
C. orientalis) in the conservation of freshwater These were relatively few and did not detract from
fishes in Malaysia. A brief review of channid the usefulness of the maps as they were from
biology was done by Ng & Lim (1990b). The states which were already heavily marked rather
information from these as well as the taxonomic than from outlying regions.
studies are combined into distribution maps for Four specimens labelled Channa marulioides
each species. and five labelled C. melanoptera deposited in the
The present paper also seeks to present data in ZRC were examined. Morphometric measure-
an attempt to clarify the identity of Channa ments and meristic counts were taken as de-
marulioides (Bleeker, 1851), which has been con- scribed in Holcik et al. (1989). Aquarium speci-
fused with C. melanoptera (Bleeker, 1855). A de- mens that refer to the above preserved specimens
scription of morphological growth changes in the were also maintained and observed. Thirty-six
poorly known acid blackwater species, C. ban- specimens of C. bankanensis in the ZRC (SL 2.7-
kanensis (Bleeker, 1852) is also presented. 23.2 cm) were examined in the same way.
Detailed description of the body coloration and
patterns of each specimen was recorded and aug-
Materials and methods mented with observation of live specimens. The
morphometric data were fitted to straight lines
Abbreviations used are SL = standard length; against standard length using the linear regres-
TL = total length; asl = above sea level. Explana- sion procedure in SPSSjPC + 4.0 (Norusis, 1986)
tion of some terms used in the tables and key are and the slopes and intercepts reported. This does
as follows: anal depth - length of longest (third not imply that the measurements are statistically
from last) anal fin ray; dorsal depth - length of regressed to standard length. The procedure was
longest (third from last) dorsal fin ray; pelvic - used simply as a line-fitting program. In addition,
anal distance - distance between tip of pelvic fin 27 specimens of C. melasoma, 38 specimens of
61
C. striata, and 20 specimens each of C. gachua, It can be generally seen that the distribution
C. lucius and C. micropeltes in the ZRC were ex- of C. gachua tends to centre on hilly and
amined. Data for all seven species were com- mountainous regions. Seven ZRC specimens
pared and a key constructed. (ZRC.735a-g) were collected at 610 m asl at Jor,
Perak; another (ZRC.916) at 1430 m asl at the
tributary of Parit waterfall, Cameron Highlands
Results and discussion (4 0 27' N 101 0 22' E), Pahang. Channa gachua
can also be found at foothills and lowlands (in-
Distribution cluding swamps) as well. Channa lucius is quite
widespread and has been recorded from acidic
Seven species of snakeheads are now known to blackwaters (Davies & Abdullah, 1989; Ng &
be present in Peninsular Malaysia, five of which Lim, 1991) but is not exclusive to it (Johnson,
also occur in Singapore: 1967). The distribution of C. melasoma seems to
be more southerly, with less records from the
Channa bankanensis (Bleeker, 1852)
north. Channa micropeltes is found where there
Channa gachua Hamilton-Buchanan, 1822
are great river and lake systems like Chenderoh
Channa lucius (Cuvier & Valenciennes, 1831)
Dam (4 58' N 100 50' E), Perak, or Tasek Bera
0 0
Channa marulioides (Bleeker, 1851)
(3 0' N 102 40' E) and Tasek Chini (3 27' N
0 0 0
Channa melasoma (Bleeker, 1851)

102 0 54' E), Pahang. It is also found in Tasek
Channa micropeltes (Cuvier & Valenciennes, 1831)
Kenyir (ca. 5 05' N 102 40' E), Trengganu
0 0
Channa striata (Bloch, 1793)

where they are angled for sport (N. Sivasothi,
Specimens from Peninsular Malaysia identified pers. comm.). Channa striata is concentrated
as C. melanoptera (Bleeker, 1855) by Tweedie among more populated areas on both coasts and
(1940, 1950) and Ng & Lim (1990b) are here absent from the central highlands, in contrast to
shown to belong to C. marulioides instead, with C. marulioides which can be found more towards
C. melanoptera being a separate species known the centre of the peninsula in Pahang. Channa
thus far only from Borneo (see later). bankanensis is an acidic blackwater species and is
The distribution of each species is shown in limited to Johor, Pahang, Trengganu and the
Fig. 1. Other than localities listed in the literature North Selangor Peat Swamp Forest. No records
and from the ZRC specimens, new data include of Channa are available from Kedah.
the record of C. marulioides 50 km northeast from
Kluang (1 0 59' N 103 0 20' E), J ohor (J. Vierke,
in litt.) and C. bankanensis in Johor, Pahang and
Trengganu. The collection of the specimen by Ecology
Vierke was recorded in Vierke (1979) but incor-
rectly identified as C. marulia. One adult, nine Channa bankanensis
juveniles and 20 fry of C. bankanensis were col- Channa bankanensis has been reported from the
lected on 18 August 1991 by the second author mouth and middle courses of rivers as well as
and M. Kottelat in blackwater ditches 21 km highly acidic blackwaters in freshwater peat
south of Muar (r 25' N 102 0 34' E) along the swamps. The water they inhabitat has been re-
Muar-Parit Sulong Road. The second author also ported to have pH 2.80-2.90 or 3.5-3.8, conduc-
collected C. bankanensis on 20 October 1991 at tivity 75.0-100.3 j.lS cm - 1 and temperature 26-
Sungai Soi, Pahang, ca. 16 km south of Kuantan 30 °C. No records of C. bankanensis are known
(3 0 49' N 103 0 21' E) along the Kuantan-Johor as yet from blackwater areas in the Larut and
Bahru road, and on 18 March 1992 ca. 56 km Krian districts of Perak. Johnson et al. (1969)
south of Kuala Trengganu along the Kuantan- listed these areas as having similar conditions to
Kuala Trengganu road (4 0 54' N 103 0 21' E). the blackwaters ofJohor, Pahang, Trengganu and
62
.........
gachua lucius
C\i:)
.........
melasoma micropeltes
<E..
c. marulioides
striata
£. bankanensis
-l-
o 200 km
Fig. 1. Maps showing the distribution of the seven species of snakeheads in Peninsular Malaysia and Singapore. Dotted lines
indicate state boundaries.
63
Selangor, and it would be interesting to find out Citations: Hamilton-Buchanan, 1822; Day, 1876;
if C. bankanensis occur in these areas as well. Weber & de Beaufort, 1922; Deraniyagala, 1929;
Tweedie, 1950; Johnson et al., 1969; Bishop,
Citations: Bleeker, 1853; Gunther, 1861; Weber & 1973; Wee, 1982; Cramphorn, 1983; Ettrich &
de Beaufort, 1922; Davies & Abdullah, 1989; Ng Schmidt, 1989; Ng& Lim, 1989; Lim & Ng, 1990;
& Lim, 1991; Ng et aI., 1991. Ng & Lim, 1990b; Ng et al., 1991; Pethiyagoda,
1991.
Channa gachua
Channa gachua has been reported from rivers, Channa lucius
lakes, ponds, well-shaded, small forest streams Channa lucius inhabits lakes, ponds, well-shaded
less than 20 cm deep, hillstreams not continuous forest streams, peat swamps and middle zones of
to mountain ranges and in the upper zone of rivers. It prefers faster moving waters and has
rivers. It has been recorded from sea-level up to been recorded in waters of temperature 24-29 °C,
1520 m als in India, but in the Malay Peninsula, pH 5.5-6.0 or less, with anion excess and low
the highest record is from the Cameron Highland calcium. Crepuscular or nocturnal, it is a mid-
specimens in the ZRC at 1430 m asl. The sub- water or surface predator relying on camouflage
strate of its habitat was reported to be leaf-litter to ambush its prey, mainly fish. Both male and
strewn, soft and muddy or consisting of laterite, female guard their young. C. lucius seems to be
pebbles, sand, silt and vegetable detritus. It in- able to survive both in black- and non-black-
habits stationary or slow-flowing water with flow waters, which might account for their wide-spread
rates reported at 22.19-68.63 cm 3 s - 1 or rapids. distribution throughout the peninsula.
Channa gachua has a wide temperature toler-
ance. It has been reported to survive tempera- Citations: Weber & de Beaufort, 1922; Johnson,
tures as low as 13 °C and has also been recorded 1967; Bishop, 1973; Mohsin & Ambak, 1983;
from hot springs in Sri Lanka. It can tolerate a Lim & Ng, 1990; Ng & Lim, 1990b; Ng et al.,
pH range of 3.1-9.6 and may inhabit waters with 1991.
low calcium, low mineral content and pronounced
anion excess. It has also been reported to enter Channa marulioides
brackish water. It is a nocturnal predator dwell- Little is known of the ecology of C. marulioides.
ing near or at the bottom. Prey items include It can be found in rivers and lakes. From the
mouse, rat, frog, tadpole, fish, Ephemerophtera distribution, it seems to be an inland or highland
and other insects, mosquito larvae, prawn (Mac- species inhabiting larger bodies of water.
robrachium sp.), crab (Irmengardia johnson i), and
other crustaceans. It moves with agility over land Citations: Bleeker, 1851; Smith, 1945; Tweedie,
and is a mouth-brooder, the male carrying the 1950; Mizuno & Furtado, 1982.
eggs and fry in its mouth. Between 20 and more
than 200 eggs per reproductive effort has been Channa melasoma
reported. Since C. gachua is well adapted to Channa melasoma has been reported from the
mountain or highland streams, its distribution middle zones of rivers and well-shaded forested
seems to follow the contours of the mountain streams with slow-flowing waters, fibrous roots of
ranges in north and central Malaysia. The adapt- trees and muddy substrate. It was recorded from
ability of the species seems to have also allowed water of pH 5-5.3. It moves over land easily, and
them to colonise lowland areas in the south, hunts at night. Prey recorded include lizard:
though many localities in Johor are near low hills Tropidophorus sp., fish: Lepidocephalichthys san-
in the region. dakanensis (as an Acanthophthalmus, fide Roberts,
1989), Rasbora sumatrana, ants, insect larvae,
crabs and other crustaceans. Only one parent
64
seems to guard the fry. If C. melasoma is more land, highly adaptable species, its wide distribu-
adapted to acidic waters, it would explain partly tion in the densely populated coastal plains can
its more southerly distribution. As one moves be explained. It is a valuable food fish which is
north up the peninsula, less records of the species easily transported, and this has probably also
is known. Smith (1945) states that it is rare in helped to distribute the species over a wide area.
Thailand.
Citations: Willey, 1909; Weber & de Beaufort,
Citations: Bleeker, 1851; Inger & Chin, 1962; 1922; Deraniyagala, 1929; Smith, 1945; Soong,
Bishop, 1973; Ng & Lim, 1990a, 1990b; Ng et al. 1949; Johnson, 1967; Bishop, 1973; Jhingran &
1991. Gopalakrishnan, 1974; Wee, 1982; Mohsin &
Ambak, 1983; Goeltenboth & Kristyanto, 1987;
Channa micropeltes Ali et al., 1988; Ataur Rahman, 1989; Davies &
Channa micropeltes is an open country species Abdullah, 1989; Munro, 1990; Ng & Lim, 1990b.
found in large rivers, lakes, reservoirs, ponds and
disturbed forest streams. It is a pelagic, midwater,
mainly diurnal predator which often aggregate as The identity of Channa marulioides
a pack. Prey include small birds, frogs and fish,
such as carp or tilapia. The parents guard the nest Channa cf. marulioides (Bleeker, 1851) (Fig. 2)
and young viscously. The distribution of C. Ophicephalus iris Cuvier & Valenciennes, 1831:
micropeltes in regions of large river courses and 439-440 [nomen nudum]
lakes fits it habitat requirements. Ophicephalus marulioides Bleeker, 1851: 424-425
[Sambas, in fluviis]
Citations: Smith, 1945; Lim & Ng, 1990; Ng & Ophicephalus marulioides - Bleeker, 1852a:89
Lim, 1990b. [Billiton]
Ophicephalus marulioides - Bleeker, 1852c:411
Channa striata [Borneo]
Channa striata can be found in lakes, reservoirs, Ophicephalus marulioides - Bleeker, 1858:223
ponds, canals, drains, streams in open country, [Billiton]
rice fields, the middle zone of rivers as well as in Ophicephalus marulioides - Bleeker, 1859:368
the more polluted lower zone. It is found from sea [Banka]
level only up to 119.5 m. Highly adaptable, C. Ophicephalus marulioides - Bleeker, 1860a: 12
striata can be found in polluted water, blackwa- [Borneo]
ter, or water with pH> 5. Conductivity of water Ophicephalus marulioides - Bleeker, 1860b: 15
that C. striata was found in was recorded as [Bankayan]
20.8-20.9 J1S cm - I. It is able to tolerate tempera- Ophiocephalus marulioides - Gunther, 1861:479
tures of II-40°C and pH values of 4.25-9.40. It [no new record]
is reported to be able to survive in slightly brack- Ophiocephalus marulioides - Bleeker, 1877:
ish water. Although it is more common in sta- Table 399, Fig. 2 [no new record]
tionary rather than flowing water, it was recorded Ophiocephalus marulioides - Bleeker, 1879:40 [no
in streams with flow rate as high as 0.47 m S-I. new record]
It moves easily over land and is reported to aes- Ophiocephalus marulioides - Weber & de Beau-
tivate during drought in Thailand. It is a crepus- fort, 1922:315 [Sumatra (Deli, Djambi)]
cular predator feeding on snakes, frogs, tadpoles, Ophiocephalus melanopterus - Weber & de Beau-
fish, Trichogaster sp., snails, insects, prawns, e.g. fort, 1922:315-316, fig. 84 [Sumatra (Indragiri,
Macrobrachium, and worms. Nest construction Gunung Sahilan, Djambi, Deli?)]
and breeding is well described and male or female Ophicephalus marulioides - Herre & Meyers,
guard the eggs and fry. Since C. striata is a low- 1937;71 [Singora, Siam]
65
Fig 2 IllustratIOns of Channa maruhO/des and C melanoptera A OphlOcephalus marullOldes, after Bleeker (1877) B 'OphlOcephalus
melanopterus', after Weber & de Beaufort (1922) C OphlOcephalus melanopterus, after Bleeker (1877)
Ophiocephalus melanopterus - Tweedie, 1940:78 Channa marulioides - Zakaria-Ismail, 1984:26

[Chenderoh, Upper Perak] [near Taman Negara, Pahang]
Ophicephalus marulioides - SmIth, 1945:468 [Tale Channa melanopterus - Zakaria-Ismail, 1984:26
Sap?] [near Kuala Tahan, Pahang]
Channa marulioides - Tweedie, 1949:101 [Tem- Channa marulioides Roberts, 1989: 170
beling, Pahang] [Kapuas, Western Borneo]
Channa melanoptera - Tweedle, 1949:101 [Kuala Channa marullOldes - Ng & Lim, 1990b:134,
Tahan, Pahang] figs. 3J, Q, R [no new record]
Channa marulia - Vlerke, 1979:274, fig. 2 [South Channa melanoptera - Ng & Lim, 1990b:134,
Malaysia] figs. 31, 0, P [no new record]
Channa marullOldes - Suvatti, 1981:124 [no new
record] Material examined. - 1 spec. (ZRC.377), George
Channa maruloides - Mizuno & Furtado, 1982: V National Park, Pahang, leg. E.O. Shebbeare,
323 [Tasek Bera, Pahang] 1939. - 1 spec. (ZRC.378), Kuala Tahan, Pahang,
66
leg. C.S. Ogilvie, 1948. - 2 specs. (ZRC.379), ides might be a colour-morph or juvenile of
Kuala Tahan, Pahang, leg. C.S. Ogilvie, 1949. - C. melanoptera. The present studies show that
1 spec. (ZRC.2834), Kuala Tahan, Pahang, leg. specimens from Peninsular Malaysia presently
C.S. Ogilvie, ix.1950. - 1 spec. (ZRC.2835), known as C. melanoptera are in fact the adult
Chenderoh Dam, Perak, leg. A.H. & M.W.T., form of C. marulioides (Bleeker, 1851), and not
iii. 1937. - 1 spec (ZRC.3071), Sungei Tahan, leg. Ophicephalus melanopterus Bleeker, 1855 s.str.
C.S. Ogilvie, xii. 1952. - 1 spec. (ZRC.9253), The morphometric and meristic characters of
Peninsular Malaysia, no data. - 1 spec. C. marulioides and C. melanoptera sensu Bleeker
(ZRC.11986), Peninsular Malaysia, no data. (1851, 1855) and Weber & de Beaufort (1922)
(ZRC.2834, 2835, 3071, 9253 and 11986 deter- were compared with those ofthe ZRC specimens
mined as C. melanoptera). identified as these species (see Table 1). There are
almost no differences in the morphometric mea-
Remarks. - Bleeker (1851) described Ophiceph- surements and meristic counts between the ZRC
alus marulioides on the basis of a TL 270 mm specimens labelled C. marulioides and those
specimen collected from Sambas, Borneo. He labelled C. melanoptera, with the possible excep-
(1855) also described Ophicephalus melanopterus tion of the proportion of eye diameter to head
on the basis of a TL 601 mm specimen from the length. When the data are compared to the de-
river Kapuas, Pontianak, Borneo. Schematic scriptions given in the literature, however, the
colour illustrations of the two species were given character becomes a continuum, and no clear-cut
in Bleeker (1877). Bleeker (1851) considered difference can be used to differentiate them.
O. iris, described by Cuvier & Valenciennes Since the quantitative characters do not shed
(1831), as invalid and a synonym of O. marulio- much light on the matter, the qualitative descrip-
ides because the former is based on a Chinese tion of the various categories had to be compared.
painting and is incompletely described. Subse- When this was done, there was correspondence
quent authors like Gunter (1861) followed amongst the categories in terms of body shape
Bleeker's descriptions of O. marulioides and and the conformation of teeth. The only distin-
O. melanopterus. Volz (1906) recorded O. melan- guishing features seem to be body coloration and
opterus from Palembang, Sumatra. He did not patterning. Basically, the descriptions of body
give a description of the specimen(s). Weber & colour and pattern for C. marulioides in Bleeker
Beaufort (1922) described O. marulioides from a (1851) and Weber & de Beaufort (1922) agree
270 mm specimen (presumably Bleeker's) and with that of the ZRC specimens labelled as such.
O. melanopterus from a 650 mm specimen from The diagnostic feature is the ocellus at the upper
an unspecified locality. The description of the part of the base of the caudal fin (see Fig.2A).
latter and its schematic illustration in Weber & de This is described in Bleeker (1851) as a black
Beaufort (1922) however, differed markedly from round spot girdled with red; and in Weber & de
that in Bleeker (1855; 1877), but no explanation Beaufort (loc. cit.) as a black white-edged ocellus.
or clarification was provided by Weber & de The ZRC specimens had white-rimmed ocelli
Beaufort (1922). Subsequent authors like Tweedie whilst live specimens had orange-rimmed ones.
(1940; 1949), Zakaria-Ismail (1984) and Ng & The ZRC specimens of'C. melanoptera' fitted the
Lim (1990b) followed Weber & de Beaufort in description of Ophiocephalus melanopterus in
identifying their fish to c. melanoptera. Roberts Weber & de Beaufort (loc. cit.) in that '(the body
(1989) recorded a 172 mm specimen of C. is) dark brown or bluish above, lighter below;
marulioides from Kapuas, Western Borneo. He along the sides, just below the lateral line, there
listed C. melanoptera but did not examine any are groups of about six scales which are black,
specimen. Ng & Lim (1990b) discussed the pos- forming a longitudinal series of irregular dark
sibility of C. melanoptera being a synonym of patches.' (see Fig. 2B). Bleeker's original descrip-
C. marulioides. They speculated that C. marulio- tion of Ophicephalus melanopterus in 1855 and his
67
Table I. Comparison of the morphometric and meristic descriptions of Channa marulioides and C. melanoptera in Bleeker (1851;
1855 & 1877) and Weber & de Beaufort (1922) with data from specimens labelled as such in the ZRC. Morphometric data are
presented as mean ± standard deviation; meristic data as ranges. Values marked with asterisks were determined from the illus-
trations in Bleeker (1877). ZRC specimens labelled C. marulioides, n = 4; c. melanoptera, n = 5. SL = standard length; TL = total
length; HL = head length; PO = postorbital head length; L.tr. = number of scale rows above/below lateral line.
Character C. marulioides C. melanoptera
Weber & Bleeker ZRC ZRC Weber & Bleeker

de Beaufort 1851 de Beaufort 1855
1922 1922
Head length in TL 3.7 ::;;4 3.9 ± 0.1 4.3 ± 0.1 4-4.6 5

Head length in SL 3.1 3.2 ± 0.1 3.5 ±O.I 3.5-3.8
Body depth in TL 7.2 7.5 7.4 ± 0.8 7.6 ± 1.4 7-7.2 7
Body depth in SL 6 6.0±0.7 6.3 ± 1.1 5.9-6.3
Head width in HL 1.8 ± 0.0 l.7±0.1 1.5-1.6
Head depth in HL 1.25 2.3 ± 0.1 2.2 ± 0.1 2
Eye diameter in HL 6.5 7 6.6± 0.6 8.9±0.9 7-8.2 7
Lower jaw length in HL 2.5 ± 0.0 2.6 ± 0.1 2.4
Upper jaw length in HL 2.5 2.9±0.0 2.9±0.1
Eye diameter: interorbit 1.6 2 2.0 ± 0.6 2.4 ± 0.4 2.5 >2
Pectoral length: PO 1.1 ± 0.1 1.0 ± 0.1 ca. I
Pelvic in pectoral length 1.7±0.1 1.8 ± 0.2 >0.5
Pelvic length: PO >1 1.8 ± 0.0 1.8±0.1
Caudal length in TL 5 5.2 ± 0.3 6.3 ± 0.5 5.5
Snout length: eye diameter ca. 1 1.2 ± 0.1 1.9 ± 0.3 <2
Pectoral length in TL 6.2± 0.4 6.3 ± 0.7 6.6
Pelvic length in TL 10.4 ± 0.5 11.3 ± 0.6 12.6
Total length 270 270 191.8 ± 35.9 493.4 ± 138.8 650 601
Dorsal fin rays 46-47 46 47 47-48 44-47 45-46
Pectoral fin rays 18 18 15-18 17-19 17-18 17
Pelvic fin rays 6 6 6 6 6 6
Anal fin rays 30-31 31 31-32 31-32 28-31 30-31
Caudal fin rays 14 15 15 14
Anal origin on dorsal ray 16 16* 14-19 14-17 16 14*
Lat. line curved at dorsal ray 16 10-14 11-15 12
Lateral line 55-58 55 55-57 55-57 54-57 55
Lat. line curved at scale 18 18* 15-16 16-18 16 18*
L.tr. 3!/1O 3i/ 1O 3i/ 10- 11 3-3i/10
Preopercular scales 5 6* 6-7 6-8 5
} 8-9
Opercular scales 3-4 4* 3-4 3-4 3-4
Predorsal scales 13 13-15 15 13-15
illustration of 1877 is quite different from this. He condition found in the ZRC specimens. The
described the body colour as violet-olive above length of the specimens in the ZRC possessing
and rosy orange below, with no mention oflateral lateral black scales ranged from TL 311 to
black scales at all. The diagnostic features listed 650 mm. Bleeker's specimen was TL 601 mm. If
by Bleeker include the dark violet vertical fins we assume that the lateral black scales are adult
which are marked by blue bands or spots (see characteristics, then Bleeker's specimen of O. m-
Fig.2C). Weber & de Beaufort (1922) describe elanopterus should have already developed them.
the vertical fins as blackish, with the dorsal and It is not very likely that Bleeker could have over-
anal somewhat freckled with whitish. This is the looked such a prominent character. It thus ap-
68
pears that C. melanoptera as described by Weber anal fins. This, together with the great variability
& de Beaufort (1922) is not con specific with that in expression of the lateral black scales in young
described by Bleeker (1855). specimens and tail ocellus in large specimens,
The specimens ZRC.11986, 2835 and 2834 (TL makes it perhaps prudent to temporarily ascrib-
311, 423 and 472 mm respectively) labelled as ing this little-known species to C. cf. marulioides
'Co melanoptera' possess lateral black scales. On pending examination of the holotype and/or
close inspection, they can be seen to possess a topotypes.
faint tail ocellus partially obscured by the dark
colour of the caudal fin. The ocellus becomes Morphological changes during growth in Channa
clearer if the caudal fin is held against a bright bankanensis
light. The remaining two larger specimens (TL
611 and 650 mm) do not seem to possess the For the Malaysian channid fauna, other than for
ocellus. Hence the ocellus could be a character C. bankanensis and C. marulioides, the morpho-
that is present in young individuals and becomes logical changes during growth ofthe other species
obscured with age as the fins darken. Live speci- have more or less been described: C. gachua by
mens that have lateral black scales edged with Deraniyagala (1929), c. lucius by Alfred (1964),
white also possess a faint caudal ocellus. C. melasoma by Ng & Lim (1990a), C. micropeltes
These observations suggest that the specimens by Tweedie (1949) and C. striata by Willey (1909).
identified from Peninsular Malaysia as C. melan- This description of the morphological changes
optera on the basis of Weber & de Beaufort's during growth in C. bankanensis is to characterise
(1922) report is merely the adult form of this little-known species and to provide future
C. marulioides; and that Ophicephalus melanoptera researchers with a reference to the morphology of
described by Bleeker (1855) and Gonther (1861) this species. It is well known that young and adult
is a separate species found in Borneo (Kapuas) channids often have strikingly different colour
and possibly Sumatra (Palembang). patterns (see Ng & Lim, 1990b) and a better un-
Both Bleeker (1851) and Weber & de Beaufort derstanding of changes associated with growth is
(1922) based their description of C. marulioides particularly relevant.
on specimens TL S 270 mm. We have seen live Data on the variation of morphometric mea-
specimens in aquaria TL 110-117 cm long al- surements with growth are presented in Table 2.
ready possessing lateral black scales fringed pos- It should also be noted that the jaws do not project
teriorly with white scales. None of the specimens behind the eye in young individuals and usually
labelled C. marulioides in the ZRC (total lengths only starts to do so in individuals longer than SL
150-232 mm) have black lateral scales. We have 12.2 cm, but specimens that are longer than this
been keeping eight specimens purchased from an may still not have developed this jaw projection.
aquarium shop alive since March 1991. At TL The following is a description of the changes in
150 mm, they still did not have definite black body coloration and pattern that occur as speci-
scales. At TL 180 mm, they only developed dark mens of Channa bankanensis increase in size. All
patches where the black scales would have been. lengths quoted are standard lengths. Figure 3 rep-
This could mean that the development of the lat- resents some typical stages in the body coloration
eral black scales is variable and may depend on of the species.
internal and/or external factors affecting the fish. Specimens of Channa bankanensis of length
The live specimens of C. marulioides possess ca. 1.5 cm are grey on the dorsal and cream on
the diagnostic orange-rimmed tail ocellus. Their the ventral part of the body. There are minute
body pattern, however, does not quite resemble faint black spots on the dorsal but the most
that shown in Bleeker (1877) in that the live speci- prominent markings are two longitudinal black
mens bear broad striations on the dorsal part of lines along the lateral sides of the body with a
the body and a mottled pattern on the dorsal and median orange band between them. The two black
69
Table 2 Changes III morphometnc measurements of Channa bankanensls WIth Illcrease III standard length (SL) expressed as a
linear functIon wIth slope (a) and Illtercept (b) Mean ± standard devIatIon (range) values of the measurements as percentage of
standard length (% SL) also gIven n = 36
Character Slope a Intercept b %SL
Body depth 016 - 015 13 7± 18 (112-210)

Head length 028 019 32 2 ± 1 9 (28 5-36 1)
Head wIdth 015 004 164±12 (128-199)
Head depth 015 -014 13 3 ± 1 4 (11 4-18 1)
Snout length 006 006 6 6± 18 (4 8-16 5)
Eye dIameter 003 007 4 5±0 7 (3 7-6 0)
InterorbItal wIdth 007 002 7 5±0 5 (6 5-8 7)
Preopercular length 013 - 0 01 124±09 (103-145)
PostorbItal head length 022 -002 21 2 ± 2 7 (66-23 3)
Upper Jaw length 009 006 10 2 ± 11 (68-12 1)
Lower Jaw length 011 004 11 8 ± 11 (10 2-16 2)
Pectoral fin length 015 012 172±15 (142-207)
PelVIC fin length 011 -006 105±09 (77-121)
Caudal fin length 015 016 18 1 ± 22 (15 1-22 1)
Dorsal depth 012 -009 10 3 ± 1 3 (69-125)
Flfst dorsal ray 005 001 52±15 (2 8-8 8)
Anal depth 011 -004 103± 10 (70-121)
PelvIc-anal dIstance 002 005 30±08 (13-48)
lines continue onto the head where the upper line two longitudinal rows runnmg the length of the
ends at a pomt at the top of the eye whilst the fin. The base of the pectorals darken and the
lower line passes through the eyes to end on the fins develop spots. On the caudal fin two to four
Jaws. The fins at thIS stage are transparent and spotted concentnc rings radIate from the base of
the two black hnes on the side of the body con- the caudal peduncle.
tinue to the tip of the caudal fin whilst the orange From 3.1-3.3 cm onwards, the lower black line
band trruls off from the base to the distal part of begins to break up as well from the posterior end
the caudal fin. The dorsal and anal fins are fruntly forwards mto a senes of mne to twelve dashes.
spotted. ThIs process may not even be complete 10 speci-
ThIs pattern of the lower black line and the mens 6.4 cm in length, though by 7.1 cm almost
median orange line continues till the fish is around all specimens possess the series of dashes. The
3.2 cm. During this period, the dorsal half of the effect of this breaking up of the two original lon-
body changes to a brown colour and gradually gitudmal black hnes IS to make the medIan light
darkens whIlst the orange median hne fades band assume a zig-zag pattern along the SIde of
qUIckly to just a light-coloured line. From 2.7 cm the body. This effect becomes obscured by the
onwards, the upper black longitudinal line starts darkening body from 6.2 cm onwards. From
to break up and gradually becomes a series of 4.3 cm onwards, the lower black line on the side
eIght to twelve transverse stnatIons. The hne dIS- of the head starts to break up as well mto an
appears on the head and the dorsal part of the enlarged black mark on each of the opercula and
head develops dIstinct spots. Two to three black an eye streak that curves down to the ventral part
eye streaks radIating downwards to the jaws de- of the head. This process may not be complete
velop. The dIstal parts of the dorsal and anal fins until up to 6.8 cm standard length. From 3.3 cm
become darker whIlst the nm of the anal fin be- onwards the ventral SIde of head and body de-
comes white. More spots appear on the dorsal velops spots. On the body, the spots eventually
and anal fins and in the former, the spots form form around 16 stnations at 5.0 cm standard
70
o E
1cm
F,g. 3. Channa bankanen.ws Specimens showing characteristic patternmgs. A. standard length (SL) = 7.2 cm, B. SL = 5.9 em;
C. SL = 3.2 cm; D. SL = 1.5 cm, lateral View; E. SL = 1.5 em, dorsal view
length; whilst on the head, they culminate in a mens would have the typical coloration and
mottled or marbled pattern from 6.1 cm on. From pattern of the species: Body brown dorsally and
4.3 cm on, the spots on the dorsal and anal fins cream ventrally; body pattern: 10-13 striations
develop into oblique streaks pointing forwards on dorsal part of the body, nine to 14 dashes on
and distally. At this time three to seven prominent lateral, and around 16 fine spotted striations on
rings can be seen on the caudal fin. The pectoral ventral; dorsal of head spotted, lateral with dis-
fins develop four to five semicircular rings from tinct opercular mark and one to three eye streaks,
6.2 cm; the pelvic fins may become spotted after ventral spotted or mottled; dorsal and anal fins
6.4 cm. with oblique dark streaks and numerous black
By around 7.2 cm, all these changes would have spots; pectoral fins with three to six rings; caudal
been completed and the vast majority of speci- fin with five to nine rings.
71
As the fish increase in length, many of these 4-14 ....................................... 2

markings start to fade. At around 10.8 cm, the 2a. Dorsal 47-48 (mode = 47); anal 31-32
striations on the dorsal part of the body may not (mode = 31); head length 29.8 ± 2.0 (27.5-
be discerned; the lateral dashes on the body may 32.6); snout length 6.0 ± 0.4 (5.5-6.5); eye
have faded to a faint impression of only four to diameter 3.9 ± 0.9 (2.8-5.3) ................ .
five marks; the oblique streaks on dorsal and anal ............................... C. marulioides
fins may become obscured; the rings on pectoral b. Dorsal 31-45; anal 20-31; head length mean
and caudal fins may have faded. By 23.2 cm, even 31.3-33.2 (range 14.9-40.0); snout length
the opercular mark and eye streak may be ob- mean 6.5-7.1 (range 4.8-16.5); eye diameter
scured. Observations oflive specimens, however, mean 4.1-5.1 (3.0-10.7) .................. 3
show that this fading of patterns is not perma- 3a. Lateral line curves down 1 scale; number of
nent, for during agonistic encounters with con- scale rows between lateral line and dorsal
specifics, the patterns have been seen to intensify origin 3.5; dorsal 31-36 (mode = 34); lateral
again. line 39-47 (mode = 44); pelvic length
8.9 ± 0.8 (7.8-10.3); preopercular length
15.5 ± 1.2 (12.5-17.2) ........... C. gachua
Key to the species of channids in Peninsular b. Lateral line curves down 2 scales; number of
Malaysia scale rows between lateral line and dorsal
origin 4.5 or 5.5; dorsal mode 38-42 (range
The present key is intended to supplement that of 33-45); lateral line mode 51-63 (43-69); pel-
Ng & Lim (1990b). Characters printed in bold are vic length mean 10.5-12.8 (range 7.7-15.3);
more diagnostic and would prove more useful in preopercular length mean 12.4-14.4 (range
differentiating the species. As far as possible, 6.2-26.4) .................................. 4
meristic characters are chosen over morphomet- 4a. Number of scale rows between lateral line and
ric ones as there is less variation in these char- dorsal origin 5.5; number of scale rows be-
acters amongst individuals of the same species. tween lateral line and anal origin 9-12
Morphometric measurements are expressed as (mode =10); preopercular scales 10-14
percentage of SL and if written as X ± Y (Z), then (mode = 10); pelvic fin in 50% of specimens
X represents the mean, Y the standard deviation extend behind origin of anal fin; lateral line
and Z the range. The key is suitable for older curves at 17-25th scale, mode = 21; body
juveniles and adults longer than SL 9-10 cm. See depth 18.6 ± 2.4; pectoral length
Materials and Methods for explanation of some 15.8 ± 1.7 .......................... C. lucius
of the terms used. b. Number of scale rows between lateral line and
dorsal origin 4.5; number of scale rows be-
la. Lateral line 83-94 (mode = 91); predorsal tween lateral line and anal origin 7-9
scales 23-30 (mode = 26); number of scale (mode =8); preopercular scales 5-10
rows between lateral line and anal origin 13- (mode = 6-8); pelvic fin does not extend be-
16, mode = 14; snout length 8.1 ± 0.8 (7.2- hind origin of anal fin; lateral line curves at
10.4); dorsal depth 7.9± 1.8 (5.1-14.8); anal 16-17th scale (modes; range 13-22); body
depth 6.9 ± 0.9 (4.5-8.1); preopercular scales depth mean 13.7-17.1 (range 11.2-21.5);
14-21, mode = 14 ............ C. micropeltes pectoral length mean 17.2-19.9 (range 14.2-
b. Lateral line 39-69; predorsal scales 12-22; 24.2) ....................................... 5
number of scale rows between lateral line and 5a. Lateral line 55-68 (mode = 63); body depth
anal origin 5-12; snout length mean 6.0-7.1 13.7 ± 1.8 (11.2-21.0); head width 16.4 ± 1.2
(range 4.8-16.5); dorsal depth mean 10.2- (12.8-19.9); upper jaw length 10.2 ± 1.1
12.8 (range 6.9-15.2); anal depth mean 9.8- (6.8-12.1); lower jaw length 11.8 ± 1.1 (10.2-
12.0 (range 6.7-14.1); preopercular scales 16.2); pelvic length 1O.5±0.9 (7.7-12.1);
72
pelvic-anal distance 3.0 ± 0.8 (1.3- This is of interest as it is still unsure if C. gachua
4.8) .......................... C. bankanensis is a variable, very widely distributed species, or
b. Lateral line mode 51-54 (range 43-59); body if there is more than one species. In Peninsular
depth mean 16.2-17.1 (11.6-21.5); head width Malaysia, we have found that certain populations
mean 18.0-19.0 (15.8-21.7); upper jaw of C. gachua, e.g. those from Panti, have distinct
length mean 12.3-12.8 (10.0-15.2); lowerjaw black spots on the head and body whilst those
length mean 13.8-14.0 (12.2-16.0); pelvic from near Kuala Lumpur and Singapore do not.
length mean 12.6-12.8 (9.0-15.3); pelvic- As for C. micropeltes and C. striata, being popu-
anal distance mean 5.9-9.6 (3.5-14.8) ... 6 lar game and/or food fish means all the more that
6a. Pelvic-anal distance 5.9 ±1.4 (3.5-8.8); inter- their conservation must be looked into, either by
orbital width 7.3 ±0.7 (5.1-8.9); postorbital maintaining viable populations in the wild to meet
head length 23.1 ± 2.6 (20.2-34.3); caudal demands, or to culture them for the market so
length 19.8 ± 2.3 (11.9-23.3); head length that wild stocks are not depleted. Looking further
33.2 ± 1.9 (28.6-37.2); preopercular length afield, C. melanoptera as originally described by
14.4 ± 2.7 (6.2-26.4) .............. C. striata Bleeker (1855) waits to be rediscovered from
b. Pelvic-anal distance 9.6 ±2.0 (4.6-14.8); in- Borneo or Sumatra. On top of this, other Indo-
terorbital width 9.1 ± 1.9 (7.6-17.8); postor- nesian species that require more study include
bital head length 20.2 ± 1.0 (17.9-22.0); cau- C. pleurophthalma and C. cyanospilos. The status
dallength 21.7 ± 2.4 (16.8-26.1); head length of Sundaic species like C. mystax, C. polylepis and
31.5 ± 1.9 (27.8-34.8); preopercular length C. baramensis remains unclear.
12.6 ± 1.0 (10.3-14.4) ......... C. melasoma
Acknowledgements
General discussion
The authors would like to thank the following
The present paper has attempted to summarise people: Mrs C. M. Yang and the staff of the Zoo-
what is known about the systematics and ecology logical Reference Collection, Dr N. Kang, Dr
of the snakeheads in Peninsular Malaysia and J. Vierke, K. K. P. Lim and others who have
Singapore. It is perhaps surprising that a family helped in one way or other. Dr M. Zakaria-Ismail
often thought of as being well-known should offer was most kind in letting us refer to his unpub-
so much scope for research. The rarer species, lished thesis. We thank Dr G. G. Teugels and Dr
C. marulioides and C. bankanensis, require further M. Kottelat for their useful comments. Work de-
study. Little is known of the ecology of the former, scribed in this paper was funded by National
and the distribution of both species requires more University of Singapore Research Grant RP
investigation. This information is needed if we are 910410.
to conserve the two species. At the moment, con-
servation of blackwater habitats like the North
Selangor Peat Swamp Forest is pressing for
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Malaysia - with emphasis on the ichthyofauna of the North
An annotated checklist of mangrove brachyuran crabs from Malaysia

and Singapore
Cheryl G. S. Tan & Peter K. L. Ng

Key words: mangrove crabs, Peninsular Malaysia, Singapore, checklist
Abstract
The mangrove brachyuran fauna of Singapore and Malaysia is generally regarded as well studied. This
is not the case. Over a hundred brachyuran species are now known from mangroves in Peninsular and
East Malaysia, a substantial part of the known fauna in these waters. In Singapore, for example, of the
approximately 350 brachyurans known, 76 (i.e. ca 22 %) are mangal species, The systematics of several
groups remain very unsatisfactory. The taxonomy of the Sesarminae, the largest subfamily of the
Grapsidae and the dominant group in most mangroves, is still very unstable, with the identities of many
ecologically important genera and species still unclear. A revision is still unavailable. Until recently,
representatives of the families Hymenosomatidae and Leucosiidae were unknown from mangroves.
Detailed collections and studies have resulted in the discovery of new genera and species from areas as
purportedly well explored as Singapore. The Sarawak mangrove brachyuran crab fauna is based almost
entirely on one paper written 40 years ago whilst that of Sabah is almost unknown. Although the state
of mangal brachyuran systematics in Malaysia and Singapore is still in an exploratory phase, the
mangroves in these two countries are relatively well-known compared to those of the surrounding re-
gions. The present annotated checklist of mangal brachyuran species is intended to guide future sys-
tematic work in these countries, as well as to assist ecologists and other mangrove biologists. This is
especially in view of the important role of crabs in mangroves.
Introduction there, and others which affect the overall man-

grove ecology (see Davie, 1982; Jones, 1984). In
Mangroves are one of the main brachyuran crab this present paper, the systematic problems asso-
habitats in Southeast Asia, with a species diver- ciated with mangrove brachyura are reviewed and
sity as high as, if not higher than coral reefs. This the state of their overall taxonomy is discussed.
fact, however, is usually not apparent or well The present checklist of brachyurans associ-
known as the mangrove brachyuran fauna is less ated with the mangrove forests in Singapore and
well studied compared to coral reefs. The defini- Malaysia (East and West) expands and updates
tion of mangrove used in this paper follows that a preliminary list prepared by D. H. Murphy (see
in Macnae (1968). The role of crabs in mangroves Chou et al., 1980) of the Singapore fauna. The
has been emphasised in recent years, especially in compilation of this list is intended to help focus
view of the commercially valuable species found future taxonomic work on the group, as well as
76
assist mangrove biologists and ecologists. Many and Sarawak were also kindly presented by
of the species listed here were originally described Dr A. Sasekumar (University of Malaya) and
from Singapore and Malaysia (here marked with Dr Charles Leh (Sarawak Museum) respectively.
an asterisk, '*', and a' + 'sign respectively). Given
the ecological importance of mangrove crabs, the
maintenance of high crab species diversity is Taxonomy
integral to the health of the mangroves.
The annotated list of taxonomic comments pro-
vided here is preliminary. Many aspects remain
Materials and methods unresolved. A good example of this is the un-
stable state of sesarmine systematics, with the
The present revised checklist is based on the separation of the various genera and subgenera,
available literature, as well as the authors' collec- as proposed by Serene & Soh (1970), being often
tions over the years. Opportunity is taken here difficult and subjective. The information from the
not only to update records and nomenclature, but unpublished theses in the Department of Zool-
also to correct some errors present in the earlier ogy, National University of Singapore by S.
report. As far as possible, the most recently pub- Harminto and N. Sivasothi has been included
lished and/or accepted nomenclature for the vari- here to make the present report complete.
ous families in question is used, although the
1. The sesarmine genus Episesarma de Man,
status and validity of many of these taxa remain
1895, is synonymous with Neoepisesarma Serene
unclear.
& Soh, 1970, with the former having priority. Se-
One of the difficulties of compiling the check-
rene & Soh (1970) established their genus as a
list was that of defining the limits of the mangrove
replacement for de Man's (1895) taxon, but
habitat. Several of the taxa listed here cannot be
Holthuis (1978) provides details of why their ac-
regarded as true mangal species, being normally
tion is unjustified. All species referred by Serene
found in other habitats. In many cases, however,
& Soh (1970) to Neoepisesarma must now be
since there is some overlap of their habitats, these
transferred to Episesarma instead. The gender of
species might be encountered. These will be re-
the genus Episesarma is neuter.
ferred to here as peripheral mangal species
(PMS). On the other hand, there are several spe- 2. The sesarmine genus Chiromantes Gistel, 1848,
cies that are commonly found not only in man- should be spelt without the second 'h'. Most lit-
groves but also in many other similar habitats. erature, including Serene & Soh's (1970) revision
These have been classified under the broad cat- incorrectly spells it as 'Chiromanthes'. The name
egory of non-obligate mangal species (NMS). A was originally proposed as a replacement name
third category, associated mangal species (AMS), for Pachysoma de Haan, 1833.
includes those species which are associated with
3. Serene & Soh (1970) established Selatium as
driftwood, barnacles and clumps of mussels
a subgenus of their new genus Neoepisesarma. The
(Perna viridis). Obligate mangal species (OMS)
subgenus is sufficiently different from Episesarma
form the last category. The classification of the
( = Neoepisesarma, see above) to regard it as a full
species is based on the available literature as well
genus, as was used by Tan & Ng (1988). The two
as the authors' collecting experiences over the
genera are distinct in many respects, not only
years.
morphologically but also ecologically and etho-
The majority of the species studied here are
logically (N. Sivasothi, unpublished data).
deposited in the Zoological Reference Collection
(ZRC), Department of Zoology, National Uni- 4. The genus Nanosesarma Tweedie, 1950, was
versity of Singapore. Specimens and some un- partially revised by Serene & Soh (1970), with
published data concerning Peninsular Malaysia several new subgenera described. The status of
77
this genus is not satisfactory and a thorough re- Macrophthalminae, a system first established by
vision is now in progress (P. Davie, pers. comm.). Alcock (1900). Serene (1974) resurrected the sub-
The various subgenera of Serene & Soh (1970) family Camptandriinae Stimpson, 1858, justify-
are here regarded as distinct genera. ing the taxon because of its peculiar highly re-
curved first male pleopod. The subfamily had
5. The crab described as Varuna yui (Grapsidae,
previously been synonymised under the Mac-
Grapsinae) by Hwang & Takeda (1986) is exter-
rophthalminae. The Camptandriinae is certainly
nally similar to V. litterata (Fabricius, 1793).
a valid taxon, sharing several apparently synapo-
Males of V. litterata however, differ markedly in
morphic features, and there are good grounds for
the form of the first male pleopod from those of
recognising it as a separate family (see Harminto
V. yui. The second author has seen the type speci-
& Ng, 1991).
mens of Cancer litteratus Fabricius, 1793. Studies
of the specimens from the region in the ZRC by 9. The two ocypodid subfamilies, Dotillinae
Peter Davie (Queensland Museum) and the sec- Stimpson, 1858, and Scopimerinae Alcock, 1900,
ond author have shown that all the Sundaic consist of the same genera and are thus synony-
Varuna are V. yui and not V. litterata (unpublished mous (Manning & Holthuis, 1981). Dotillinae,
data). It would appear that V. yui is more com- being the senior name, has priority and is thus
mon in the continental shelf waters while V. lit- used instead of Scopimerinae in this checklist.
terata prefers habitats facing the open ocean.
10. Shenius anomalum (Shen, 1935) occupies a
6. The family Xanthidae MacLeay, 1838, has very peculiar position in the Dotillinae. Its exter-
been thoroughly revised in recent years, and the nal features and gonopod structures, as well as its
modern arrangement differs from that adopted by larvae are very unusual (S. Harminto, unpub-
Balss (1957), which is followed by many authors. lished data). It should probably be separated into
There is, however, sufficient evidence (see Guinot, a new subfamily, or even family.
1978, 1979; Ng, 1987) that the family should be
11. The fiddler crabs of the genus Uca sensu lato
split into several separate families (Xanthidae
have undergone extensive revision in the past few
s. str., Carpiliidae, Panopeidae, Pilumnidae,
decades, the most recent, and certainly the most
Menippidae etc.) instead, and tentatively grouped
comprehensive was by Crane (1975). There are,
in one superfamily. Guinot's (1978) classification
however, still many practical problems with the
is followed in this checklist, although super-
use of many of the subgenera proposed. There are
family rank is not used here.
also serious taxonomic and nomenclatural prob-
7. The Rhizopinae Stimpson, 1858, is here re- lems in this system especially since Bott (1973a, b)
garded as a subfamily of the family Pilumnidae had earlier split Uca into ten genera based solely
(see Ng, 1987). In the older system ofBalss (1957) on the gonopod tip structure. In this paper, the
and other authors, the Rhizopinae are classified authors prefer to adopt Von Hagen's (1976) sug-
under the Goneplacidae. The weight of existing gestion that only one broad genus, without sub-
evidence however, suggests that the Goneplacidae genera, be recognised for the time being.
sensu Balss (1957) (and aut.) is a heterogeneous
12. The camptandriine genus Leipocten Kemp,
assemblage and must be divided into several
1915, is a junior synonym of Baruna Stebbing,
smaller groups (see Guinot, 1978; Ng, 1987).
1904. Four distinct species are now known (Dai
8. The higher classification of the Ocypodidae is & Song, 1986; Harminto & Ng, 1991). The man-
still not completely clear. Carcinologists at grove species known locally as Leipocten sordidu-
present divide the Ocypodidae into three, four or lum Kemp, 1915, by most authors after Kemp
five subfamilies. Most of the modern workers (1915) is in fact, a separate species, Baruna man-
(e.g. Lewinsohn, 1977; Serene, 1968) recognise gromurphia Harminto & Ng, 1991. The true
three subfamilies - Ocypodinae, Dotillinae and L. sordidulum is a junior synonym of B. socialis
78
Stebbing, 1904, and occurs in India and northern genus Nursia Leach, 1817, by Tweedie in 1937
Peninsular Malaysia in non-mangrove habitats. (unpublished data, M. W. F. Tweedie, pers.
Dai & Song (1986) described a new species, comm.). Other Malaysian specimens were also
L. trigranulum which possessed first male pleo- determined as N. malefactrix by Serene in 1970
pods very similar to that of B. mangromurphia. (unpublished data). As the taxonomy of the genus
The carapace features of the males seem to dif- Ebalia is still very unclear, and to avoid confu-
fer and until direct comparison can be made of sion, the species malefactrix is retained within the
the two species, both taxa are regarded as valid. genus Ebalia.
13. The camptandriine genera Cleistostoma de
Haan, 1833, and Paracleistostoma de Man, 1895,
Discussion
have been revised by Manning & Holthuis (1981),
and have been separated into several smaller gen-
Although the contributions by Tweedie, Serene
era. The most significant change for a local spe-
and Johnson to mangrove brachyuran taxonomy
cies is for Paracleistostoma microcheirum Tweedie,
of Singapore, West and East Malaysia have been
1937, which has been transferred to a monotypic
significant, mangrove brachyuran crab taxonomy
genus Ilyogynis Manning & Holthuis, 1981. Un-
in Singapore and Malaysia remains unstable as is
published studies by S. Harminto, have also re-
true for mangrove fauna in general. Compared to
vealed the presence of a Paracleistostoma species
other regions in Southeast Asia however, the
not previously known from Peninsular Malaysia
mangroves of Malaysia and Singapore are rela-
- P. wardi (Rathbun, 1910). This species was,
tively well-explored. New species are still being
until then, only known from Australia.
discovered, and regional studies have shown that
14. One of the more serious and pressing prob- much of the fauna remains poorly known.
lems of mangrove brachyuran taxonomy is the Tweedie (1950) listed only 46 species of mangal
identity of the commercially important mud crab, brachyuran fauna from Labuan and Sarawak, the
Scylla serrata (ForskaI, 1755) (type locality Jidda, majority being sesarmines. This is the only paper
Red Sea). The number of species within the genus on Sarawak mangrove crabs published, although
Scylla has been the subject of much discussion Tai & Manning (1984) described a new Pota-
and study and it is not known whether the man- mocypoda from streams near a mangrove in
grove species is the true Scylla serrata. Pending a Sarawak. The mangrove brachyura of Sabah is
definite revision of the genus, the specific name almost unknown, with no known detailed study
serrata is used for the mangrove species. A de- ever made. The shortage of recent taxonomic lit-
tailed study is clearly necessary, especially if their erature for the mangrove brachyuran fauna of
fisheries is to be properly managed. Singapore and Malaysia is not a reflection of the
taxonomic stability for species inhabiting man-
15. A new genus and species of obligate man-
grove areas. It is, however, an indication of the
grove leucosiid, Praosia punctata, apparently allied
need for more taxonomic work than is now being
to the genera Nursia and Ebalia was recently de-
done in the two countries.
scribed. Due to the existing confusion concerning
The keys to the various taxa of crabs from the
the definition, size and composition of the various
mangrove are few and often fall into either of two
leucosiid subfamilies, with the possible exception
categories: those that add to the complexity of the
of the Ebaliinae and the Cryptocneminae (see
problem rather than serve to ease the situation,
Manning & Holthuis, 1981), Praosia punctata has
and those that may be helpful but are outdated.
not been classified under any subfamily (Tan &
An example of the former would be the compila-
Ng, 1992).
tion by Lovett (1981). Portions of keys were
16. Specimens of Ebalia malefactrix, Kemp 1915, compiled apparently 'piecemeal' from different
from Peninsular Malaysia, were placed in the authors, including taxa not found here and
79
obvious errors, and many important keys in sup- largest family found in Singaporean and Malay-
posedly 'less well known' publications were omit- sian mangroves. The Ocypodidae, the next largest
ted. Such superficial keys more often than not family, makes up about 30%.
prove to be misleading rather than useful and Many of these crabs are economically and eco-
usually hamper the work of identification of the logically important. Primary food species include
fauna concerned. Some provisional keys to the the mud crab, Scylla serrata and the larger
brachyuran fauna of the mangroves of the Indo- sesarmines of the genus Episesarma. A species
Pacific were written by the late R. Serene in 1972. with commercial potential is the mangrove stone
These, however, remain unpublished. crab, Myomenippe hardwicki, a relative of the
Although the mangrove habitat has a great commercially valuable American stone crab,
abundance of molluscs, such as Anadara granosa Menippe mercenaria (see Williams, 1984). Myo-
and Glauconome rugosus, no pinnotherids, often menippe hardwicki is common in Malaysian and
closely associated with bivalves, have been re- Singapore mangroves and their large chelae
ported thus far from mangroves in Singapore and (claws) in particular are meaty and tasty but as
Malaysia. As many species of pinnotherids are yet, this species is not harvested.
symbionts of mangrove bivalves, they are almost Many species, in particular the sesarmines and
certainly present in these waters. Unidentified ocypodids, are important in mangrove energetics,
species (Pinnotheres spp.) have however been ob- being involved in nutrient cycling within the eco-
tained by the second author from Anadara gra- system. Members of the different families exhibit
nosa and Anomia sp. Special efforts to obtain such varied feeding habits as scavenging, detrital-
mangrove pinnotherids will have to be made. feeding, plankton-feeding and herbivory. Previ-
In recent years, a number of new species have ously, the role of sesarmine crabs as 'trophic
been discovered, particularly from the families vectors' was only considered from the aspect of
Hymenosomatidae (false spider crabs) and Leu- litter consumption (see Macnae, 1968; Robert-
cosiidae (pebble crabs) from Singapore man- son, 1986). Watson (1928) identified crabs as one
groves. The known mangrove leucosiids of of the most serious enemies to Malayan man-
Malaysia are based almost entirely on recent col- grove plantations, with particular reference to
lections (unpublished data) from Johor and Episesarma feeding on mangrove propagules. The
Penang (Ebalia malefactrix). One hymenosoma- degradation of leaf litter mainly by grapsid crabs
tid (Elamenopsis mangalis) is known from Johor, and other organisms provides detritus which may
Malaysia (C. T. N. Chuang, pers. comm.). Many be utilised by detritivores or decomposed by bac-
members of these families are cryptic, being of teria and fungi (Robertson, 1986). Recently, it
similar coloration to their surroundings and usu- was found that active predation by crabs on in-
ally partially buried in mud or vegetation, making tact mangrove foliage occurs, by the tree-climbing
their detection and collection difficult. Thus, in- sesarmine crabs such as Episesarma versicolor,
tensive collections must be made for members of E. chengtongense and E. singaporense (Sivasothi
such cryptic families previously undetected so as et al., 1993). Prior to this, it was thought that crab
to facilitate a complete documentation of the herbivory was restricted to leaf litter, propagules
mangrove species. and flowers. This discovery has important impli-
The significance of the mangrove crab fauna cations with regards to the contribution of crabs
cannot be overstated as Davie (1984) succinctly to the overall energetics of the mangrove ecosys-
puts it, ' ... these animals are by far the most con- tem. Secondary production in the mangrove for-
spicuous and appealing component of the inver- est is beneficial to the surrounding habitats as it
tebrate fauna'. In Singapore and Malaysia, the has already been established that the elements of
mangrove grapsids, of which some 90% are animal production in the forest are not only
sesarmines, comprise about 53 % of the total exported to the adjoining sea, but also consumed
mangrove crab fauna, making the Grapsidae the by marine fish visiting the forest at high tide
80
(Sasekumar, 1980). In the light of all the above, Balss, H., 1957. Decapoda. VII. Systematik. In H. G. Bronns,
serious consideration must be given to the con- Klassen & Ordnungen des Tierreichs 5: 1505-1672.
Bott, R., 1973a. Die typus-art der gattung Uca Leach, 1814
servation of the mangrove habitat and its diverse (Decapoda: Ocypodidae). Senckenbergiana bio!. 54: 315-
fauna. 325.
Bott, R., 1973b. Die verwandtschaftlichen Beziehungen der
Uca-Arten (Decapoda: Ocypodidae). Senckenbergiana
Conclusion bio!. 54: 315-325.
Chou, L. M., S. H. Ho, H. W. Khoo, T. J. Lam, D. H.
Any plan for the conservation of the mangrove Murphy & W. R. Tan, 1980. The present state of mangrove
habitats in Singapore and Malaysia must incor- ecosystems in Singapore and the impact of pollution. South
porate taxonomic studies on the brachyuran China Sea Fisheries Development and Coordinating
fauna. Little can be done in preparing usable keys Programme, Food and Agriculture Organization of the
United Nations, United Nations Environment Program,
for the mangal brachyuran fauna and habitats Singapore, 76 pp.
until adequate attempts are made at stabilising Crane, J., 1975. Ocypodidae: genus Uca. Fiddler crabs of the
the alpha-taxonomy of these taxa. world, Princeton University Press, New Jersey, 736 pp.
Dai, A-Y. & Y-Z. Song, 1986. Intertidal crabs from the Beibu
Gulf of Guangxi. Trans. Chin. Crust. Soc., Beijing: 54-62.
Acknowledgements Davie, P. J. F., 1982. A preliminary checklist of brachyura
(Crustacea: Decapoda) associated with Australian man-
The authors wish to thank Professors L. B. grove forests. Operculum 5: 204-207.
Holthuis, Y. Nakasone and D. A. Jones for their Davie, P. J. F., 1984. The biogeography of littoral crabs
(Crustacea: Decapoda; Brachyura) associated with tidal
helpful comments on the manuscript, Mr S. wetlands in tropical and sub-tropical Australia. In K. N.
Harminto and Mr N. Sivasothi for kindly con- Bardsley, J. D. S. Davie & c. D. Woodroffe (eds), Coasts
senting to the use of materials from their theses, and tidal wetlands of the Australian monsoon region,
Mr Michael Tweedie, Mr Kelvin Lim, Ms Eunice Australian National University North Australia Research
Low, Mr Dennis Ng, Miss C. T. N. Chuang and Unit mangrove Monograph 1, Darwin: 259-275.
Davie, P. J. F., 1992. Revision of Sarmatium Dana (Crusta-
friends at the Reef Ecology laboratory for their cea: Brachyura: Sesarminae) with descriptions of three new
encouragement and support in the course of writ- species. Mem. Qld. Mus. 32: 79-97.
ing this paper, Mr Peter Davie from the Queens- Guinot, D., 1978. Principes deune classification evolutive des
land Museum for most interesting conversations Crustaces Decapodes Bracyuores. Bull. Bio!. Fr. Belg. 112:
on some aspects of grapsid taxonomy, Mrs 211-292.
Guinot, D., 1979. Donnees nouvelles sur la morphologie, la
C. M. Yang for her kind permission to examine phylogenese et la taxonomie des Crustaces Decapodes
the specimens in the ZRC, and Dr A. Sasekumar Brachyoures. Mem. Mus. natn. Rist. nat., Paris, Zoo!. 112:
for kindly referring many of the mangrove speci- 1-354.
mens to the authors for study. The authors would Harminto, S. & P. K. L. Ng, 1991. A revision of the camptan-
also like to acknowledge Dr Chou Loke Ming driin genus Baruna Stebbing, 1904 (Crustacea: Brachyura:
Decapoda: Ocypodidae), with descriptions of two new spe-
and the Reef Ecology Study Team, Department cies from the Indo-West Pacific. Raffles Bull. Zoo!. 39:
of Zoology, National University of Singapore, 187-207.
which collected specimens under the 'ASEAN- Holthuis, L. B., 1978. A collection of decapod crustacea from
Australia Marine Science Project: Living Coastal Sumba, Lesser Sunda Islands, Indonesia. Zoo!. Verh.,
Resources'. Research was made possible by a Leiden 162: 18-687.
Hwang, J. & M. Takeda, 1986. A new freshwater crab of the
grant RP900360 from the National University of
family Grapsidae from Taiwan. Proc. Jap. Soc. Syst. Zoo!.
Singapore. 33: 11-18.
Jones, D. A., 1984. Crabs of the mangal ecosystem. In
F. D. Por & I. Dor (eds), Hydrobiology of the Mangal. The
References
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India, The Brachyura Catometopa or Grapsoidea. J. Asiat. (Crustacea, Decapoda, Brachyura). Zoo!. Verh., Leiden
152: 45-84.
Soc. Beng. 69: 279-486.
81
Lovett, D. L., 1981. A guide to the shrimps, prawns, lobsters, to Sesarma Say 1817 (Brachyura, Decapoda, Crustacea).
and crabs of Malaysia and Singapore. Faculty of Fisheries Treubia 27: 387-416.
and Marine Science, Universiti Pertanian Malaysia, Sivasothi, N., D. H. Murphy & P. K. L. Ng, 1993. Tree-
Malaysia, 156 pp. climbing and herbivory of crabs in the Singapore man-
Macnae, W., 1968. A general account of the flora and fauna groves. In A. Sasekumar (ed.), Mangrove Fisheries and
of mangrove swamps and forests in the Indo-West Pacific Connections. Asean-Australia Marine Science Project.
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Manning, R. B. & L. B. Holthuis, 1981. West African Tai, A-Y. & R. B. Manning, 1984. A new species of Pota-
Brachyuran crabs (Crustacea: Decapoda). Smithson. mocypoda (Crustacea: Brachyura: Ocypodidae) from
Contr. Zool. 306: 1-379. Malaysia. Proc. bioI. Soc. Wash. 97: 615-617.
Ng, P. K. L., 1987. The Indo-Pacific Pilumnidae II. A revi- Tan, L. W. H. & P. K. L. Ng, 1988. A guide to seashore life.
sion of the genus Rhizopa Stimpson, 1858, and the status Singapore Science Centre, Singapore, 159 pp.
of the Rhizopinae Stimpson, 1858 (Crustacea, Decapoda, Tan, C. G. S. & P. K. L. Ng, 1993. Praosia punctata, a new
Brachyura). Indo-Malay. Zoo!. 4: 69-111. genus and species of mangrove leucosiid crab (Crustacea:
Robertson, A. 1., 1986. Leaf-burying crabs: their influence on Decapoda: Brachyura) from Singapore, Crustacean a 64(1):
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Serene, R., 1968. The brachyura of the Indo-West Pacific Von Hagen, H. 0., 1976. Review of 'Fiddler crabs of the
Region. Prodromus for a checklist of the non-planktonic world. Ocypodidae: genus Uca' by J. Crane, Princeton Uni-
marine fauna of Southeast Asia. Singapore Natn. Acad. versity Press. Crustacean a 31: 221-224.
Sci., Special Publication. 1: 33-112. Watson, J. G., 1928. Mangrove forests of the Malay Penin-
Serene, R., 1974. Note on the genera and species of the sula. Malay. Forestry Records 6: 1-275.
Camptandriinae Stimpson, 1858 (Decapoda, Brachyura, Williams, A. B., 1984. Shrimps, lobsters and crabs of the
Ocypodidae). Treubia 28: 59-68. Atlantic Coast of the eastern United States, Maine to
Serene, R. & C. L. Soh, 1970. New Indo-Pacific genera allied Florida. Washington, 550 pp.
82
Checklist
BRACHYURA
Status S'pore E. M'sia W. M'sia
Grapsidae
Grapsinae
Metopograpsus frontalis Miers, 1880 NMS X
Metopograpsus gracilipes de Man, 1891 NMS X X
Metopograpsus latifrons (White, 1847) NMS X X X
Metopograpsus quadridentatus Stimpson, 1858 NMS X
Sesanninae
Beanium andersoni (de Man, 1887) AMS X X
Beanium batavicum (Moreira, 1903) AMS X X
Beanium edamense (de Man, 1887) AMS X
Beanium nunongi (Tweedie, 1950) + AMS X X
Bresidium sediliensis (Tweedie, 1940) + OMS X X
Chiromantes dussumieri (H. Milne Edwards, 1853) OMS X
Chiromantes eumolpe (de Man, 1895) OMS X X X
Chiromantes fasciatus (Lanchester, 1900)* OMS X X
Chiromantes indiarum (Tweedie, 1940) OMS X X X
Chiromantes onychophorus (de Man, 1895) OMS X X
Chiromantes semperi BOrger, 1895 OMS X X
Cleistocoeloma lanatum Alcock, 1900 OMS X
Cleistocoeloma merguiense de Man, 1888 OMS X X X
Cleistocoeloma suvaense Edmondson, 1951 OMS X
Episesarma chengtongense Serene & Soh, 1967 * OMS X X
Episesarma mederi H. Milne Edwards 1853 OMS X X X
Episesarma palawanense (Rathbun, 1914) OMS X X X
Episesarma singaporense (Tweedie, 1936)* OMS X X
Episesarma versicolor (Tweedie 1940) * OMS X X X
Labuanium politum (de Man, 1888) OMS? X
Metaplax crenulata (Gerstaecker, 1856) OMS X X
Metaplax elegans de Man, 1888 OMS X X X
Metaplax sheni Gordon, 1930 OMS X
Nanosesarma minutum (de Man, 1887) AMS X
Nanosesarma pontianacense (de Man, 1893) AMS X
Neosarmatium smithii (H. Milne Edwards, 1853) OMS X X
Neosesama gemmiferum (Tweedie, 1936) + /* OMS X X X
Parasesarma batavianum (de Man, 1890) OMS X X
Parasesarma calypso de Man, 1895 OMS X
Parasesarma lanchesteri (Tweedie, 1936) OMS X X
Parasesarma melissa (de Man, 1887) OMS X X
Parasesarma plicatum (Latreille, 1806) OMS X X
Parasesarma rutilimanum (Tweedie, 1936)* OMS X X X
Pseudosesarma bocourti (A. Milne Edwards, 1868) OMS X
Pseudosesarma crassimanum (de Man, 1887) OMS X
Pseudosesarma edwardsii (de Man, 1887) OMS X
Pseudosesarma johorense (Tweedie, 1940) + OMS X X X
Pseudosesarma laevimanum (Zehntner, 1894) OMS X
Sarmatium striaticarpus Davie, 1992 OMS X X
Sarmatium germaini (A. Milne Edwards, 1867) OMS X
Selatium brocki (de Man, 1887) OMS X X X
Sesamoides borneensis (Tweedie, 1950) + OMS X X
83
(Continued)
Sesarmoides kraussi (de Man, 1887) OMS X

Tiomanium indicum (H. Milne Edwards, 1837) OMS X X X
Varuninae
Ilyograpsus paludicola (Rathbun, 1909) OMS X X X
Utica bomeensis de Man, 1895 PMS X X X
Varuna yui Hwang & Takeda, 1986 PMS X X
Hymenosomatidae
Elamenopsis mangalis Ng, 1988* OMS X X
Menippidae
Epixanthus dentatus (Adams & White, 1848) OMS X X
Myomenippe hardwicki (Gray, 1831) NMS X X X
Ozius guttatus H. Milne Edwards, 1834 PMS X X
Dorippidae
Nobilum histrio (Nobili, 1903) PMS X
Neodorippe callida (Fabricius, 1798) NMS X X X
Leucosiidae
Praosia punctata Tan & Ng, 1993* OMS X
Ebaliinae
Ebalia malefactrix Kemp, 1915 OMS X X
Ocypodidae
Camptandriinae
Baruna mangromurphia Harminto & Ng, 1991 * AMS X X
Camptandrium elongatum Rathbun, 1929 OMS X X
Ilyogynis microcheirum (Tweedie, 1937)* OMS X X
Paracleistostoma depressum (de Man, 1895) OMS X X
Paracleistostoma longimanum Tweedie, 1937 * OMS X
Paracleistostoma wardi (Rathbun, 1926) OMS X X
Tylodiplax tetralyphora de Man, 1895 OMS X
Macrophthalminae
Macrophthalmus brevis (Herbst, 1804) NMS X
Macrophthalmus erato de Man, 1888 NMS X X
Macrophthalmus latreillii (Desmarest, 1822) NMS X
Macrophthalmus tomentosus Souleyet, 1841 NMS X
Macrophthalmus crinitus Rathbun, 1913 NMS X
Ocypodinae
Ocypode ceratophthalmus Pallas, 1772 PMS X X X
Uca coarctata (H. Milne Edwards, 1852) OMS X
Uca dussumieri H. Milne Edwards, 1852 OMS X X
Uca rhizophorae Tweedie, 1950 + OMS X X
Uca annulipes (H. Milne-Edwards, 1837) PMS X X
Uca spinata Crane, 1975 OMS X X
Ucaforcipata (Adams & White, 1848) OMS X X
Uca lactea (de Haan, 1835) OMS X
Uca perplexa (H. Milne Edwards, 1837) OMS X X
84
(Continued)
Uca rosea (Tweedie, 1937)* OMS X X

Uca triangularis (A. Milne Edwards 1873) OMS X
Uca vocans (Linnaeus, 1758) PMS X X
Dotillinae
Dotilla wichmanni de Man, 1892 PMS X X
Dotilla myctiroides (H. Milne Edwards, 1852) PMS X X
Dotillopsis brevitarsis (de Man, 1888) OMS X
Ilyoplax delsmani de Man, 1926 OMS X X X
Ilyoplax lingulata (Rathbun, 1909) OMS X X
Ilyoplax longicarpa Tweedie, 1937 + OMS X
Ilyoplax obliqua Tweedie, 1935* OMS X X
Ilyoplax orientalis (de Man, 1888) OMS X
Ilyoplax punctata Tweedie, 1935 * OMS X X
Scopimera globosa de Haan, 1835 PMS X
Scopimera intermedia Balss, 1934 PMS X X X
Shenius anomalus (Shen, 1935) OMS X X
Pilumnidae
Pilumninae
Heteropanope glabra Stimpson, 1858 NMS X X
Parapilumnus quadridentatus de Man, 1895 AMS X X X
Rhizopinae
Luteocarcinus sordidus Ng, 1990 + OMS X
Xenophthalmus pinnotheroides White, 1846 OMS X
Portunidae
Portunus pelagicus (Linnaeus, 1758) PMS X X X
Scylla serrata (ForsUI, 1755) NMS X X X
Pinnotheridae
Pinnotheres sp. A OMS X
Pinnotheres sp. B NMS? X
Number of species in respective country/region 76 40 76

Total number of species in Singapore and Malaysia = 192 spp.
The ecology and biology of Southeast Asian false spider crabs

(Crustacea: Decapoda: Brachyura: Hymenosomatidae)
Christina T. N. Chuang & Peter K. L. Ng

Key words: Hymenosomatidae, Southeast Asia, ecology, distribution, biology
Abstract
The false spider crabs of the family Hymenosomatidae are one of the most poorly known group of
brachyuran crabs in Southeast Asia. This is largely attributed to their small size and cryptic behaviour.
Despite the many studies of decapods in Southeast Asia, only eleven species of hymenosomatids are
known. Of these, two genera and six species were only described in the last two years. From Singapore
the largest number of species (five) has been reported probably because it has been the best explored,
while from Thailand three species and Peninsular Malaysia two species are known. There are no pub-
lished records of hymenosomatids from Borneo or Java. The hymenosomatid fauna is perhaps best
known in Australia and New Zealand. It is anticipated that when proper collections are made and studies
implemented, the Southeast Asian hymenosomatid fauna will prove to be more diverse than what has
been reported. The ecology of the Southeast Asian species is reviewed, especially in the view that two
species are completely freshwater, one of which is a troglobite. Most species are littoral or sublittoral
in habit and very sensitive to human activities (e.g. pollution).
Introduction ing them strong contenders for being the smallest

crabs in the world (see Lucas & Davie, 1982).
The hymenosomatid crabs are more commonly Their small size, dull and cryptic colouration and
known as crown crabs or false spider crabs. De- habitat have resulted in them being poorly known
spite their general similarity to the true spider and even less well collected in Southeast Asia.
crabs (Majidae), they differ in many respects, in- Consequently, not much is known about the Sun-
cluding zoeal morphology (see Rice, 1980). They daic species.
are also very peculiar in that they are the only What is known about Southeast Asian hy-
brachyurans which have six (not seven) abdomi- menosomatids however, contrasts very strongly
nal segments (see Tesch, 1918; Lucas, 1980). This with what has been reported for India (see Kemp,
is also reflected in their zoeal abdomina (see Rice, 1917; Chopra & Das, 1930), New Zealand (see
1980). Hymenosomatids are among the smallest Melrose, 1968), Australia (see Lucas, 1980; Lucas
brachyuran crabs, their carapace widths ranging & Davie, 1982). Southeast Asia has only ten spe-
from 2 to 26 mm (Lucas, 1980; unpublished data). cies. Up to 1988, only four species were known
In fact, some species (e.g. ovigerous Elamenopsis from Southeast Asia - Halicarcinus coralicola
minima and adult male Halicarcinus orientalis) are (Singapore, Thailand), Trigonoplax unguiformes
only about 1.7 mm in adult carapace width, mak- (Singapore), Elamena sindensis (Singapore), and
86
Elamenopsis exigua (Thailand) (Rathbun, 1909; (Contmued)
Lanchester, 1900; Yang, 1979). In the last two Species DlstnbutlOn Habitat
years, the authors have described two new genera
and five new species. Three species are from Elamenopsls eXlgua Thailand bracklsh/
freshwater
Singapore, one from Thailand and one from Elamenopsls lmeala PhllIppmes bracklsh-
Sulawesi. The present studies strongly suggest marine
that, with more extensive collections, more spe- Elamenopsls mangabs Smgapore, Malay- brackish
sla
cies can be expected to be discovered in the Elamenopsls palawanensls PhllIppmes, Sm- manne
future. The hymenosomatids are not depauperate gapore
in Southeast Asia as previous studies have indi- Hailcarcmus corabcola Smgapore, Thru- marine
land
cated. The present paper serves to review what is Llmnoplios nOlyanetrl Thailand freshwater
known about Southeast Asian hymenosomatid
taxonomy, ecology and zoogeography.
Most of the old hymenosomatid records proved
to be incorrect. The species listed as Elamena
Methods and materials sindensis, in Yang (1979), is Elamenopsis mangalis
(unpublished data). Lanchester's (1900) record of
The terminology used here follows that used by Trigonoplax unguiformis from Singapore, which
Melrose (1968, 1975) and Lucas (1980). Speci- has been cited by almost all subsequent workers
mens studied are deposited in the Zoological (e.g. Tesch, 1918; Sakai, 1938, 1976; Lucas, 1980;
Reference Collection (ZRC), Department of Ng, 1988a) proved to belong to Elamena mendosa
Zoology, National University of Singapore. instead. One of the more widely distributed spe-
cies, Halicarcinus coralicola needs to be further
studied. The species was described very briefly on
Taxonomy the basis of only one female from Singapore (as
a Rhynchoplax, see Rathbun, 1909, 1910) and only
The taxonomy of the Hymenosomatidae has been the cheliped and leg were figured. Considering
relatively well revised by Melrose (1968) and more that this description is so meagre (the type has
recently by Lucas (1980). Hymenosomatid sys- never been reexamined or redescribed since 1910)
tematics is thus relatively stable compared to and Halicarcinus coralicola has a wide reported
many of the better known groups. There are 75 distribution, ranging from Southeast Asia to
species known at present from the world Taiwan and Japan (Naiyanetr, 1980; Sakai, 1976;
(Table 1). Of these, only one native species oc- Lucas, 1980), direct comparisons of materials
curs in the Americas, the majority of the species from different localities must be made. A detailed
being Indo-West Pacific in distribution. taxonomic revision of the Southeast Asian Hy-
menosomatidae is being prepared at present and
The ten species known from Southeast Asia the results will be published at a later date.
(fide Lucas, 1980; Ng, 1988a, 1991; Chuang &
Ng, 1991) are: The taxonomic characters which have proved
useful in hymenosomatid taxonomy are as fol-
lows:
Species DistrIbutIOn Habitat
Amarmus woltereckl PhllIppmes freshwater

1. carapace shape, presence or absence of spines
Cancrocaeca xenomorpha SulaweSI, IndoneSia freshwater on lateral carapace margins and the presence
Elamena cr/Stallpes MalaySia manne or absence of distinct marginal groove,;
Elamena globosa Smgapore manne
Elamena mendosa Smgapore maflne
2. size of the third maxilliped in relation to the
buccal cavity;
87
Table I. List of the World Species of Hymenosomatidae.
Amarinus angelicus (Holthius, 1968) Papua-New Guinea

Amarinus lacustris (Chilton, 1882) S.E. Australasia
Amarinus laevis (Targioni-Tozzetti, 1877) southern Australia
Amarinus latinasus Lucas, 1980 N .E. Australia
Amarinus lutarius Lucas & Davie, 1982 Queensland, Australia
Amarinus paralacustris (Lucas, 1970) eastern Australia
Amarinus pilosus (A. Milne Edwards; 1873) New Caledonia
Amarinus wolterecki (Balss, 1934) Mindanao, Philippines
Cancrocaeca xenomorpha Ng, 1991 Sulawesi, Indonesia
Elamena abrolhensis Gordon, 1940 southern Australia
Elamena cimex Kemp, 1915 Chilka Lake, India; Madagascar
Elamena cristatipes Gravely, 1927 India; Malaysia
Elamena globosa Chuang & Ng, 1991 Singapore
Elamena gordonae Monod, 1956 W. Africa; N.E. Australia
Elamena gracilis Borradaile, 1903 Maldive Archipelago
Elamena longistrostris Filhol, 1885 New Zealand
Elamena mathaei (Demarest, 1825) Red Sea; S.E. Africa
Elamena mendosa Chuang & Ng, 1991 Singapore
Elamena momona Melrose, 1975 New Zealand
Elamena producta Kirk, 1879 New Zealand
Elamena sindensis Alcock, 1900 India; Sri Lanka
Elamena truncata (Stimpson, 1858) Indo-west Pacific
Elamena umerata Lucas, 1980 northern Australia
Elamena xavieri Kemp, 1917 Mandavi R., India
Elamenopsis alcocki (Kemp, 1917) West coast, India
Elamenopsis ariakensis (Sakai, 1969) Nagasaki Prefecture, Japan
Elamenopsis aspinifera Lucas, 1980 N. Queensland, Australia
Elamenopsis bovis (Barnard, 1946) South Africa
Elamenopsis demeloi (Kemp, 1917) Goa, India
Elamenopsis exigua (Kemp, 1917) Thailand
Elamenopsis frontalis Lucas & Davie, 1982 Queensland, Australia
Elamenopsis hirtirostris Lucas & Davie, 1983 Queensland
Elamenopsis inachoides (Alcock, 1900) West Bengal, India
Elamenopsis inermis (Takeda & Miyake, 1971) Palau Islands
Elamenopsis introversa (Kemp, 1917) Kiangsu Prov., China
Elamenopsis kempi (chopra & Das, 1930) Basra, Iraq; Panama canal
Elamenopsis lineata A. Milne Edwards, 1873 New Caledonia; Queensland; Philippines
Elamenopsis mangalis Ng, 1988 Singapore
Elamenopsis minima Lucas & Davie, 1982 Queensland, Australia
Elamenopsis nasalis (Kemp, 1917) West Bengal, India
Elamenopsis octagonalis (Kemp, 1917) Goa, India; Queensland, Australia
Elamenopsis palawanensis (Serene, 1971) Palawan Is., Philippines; Singapore
Elamenopsis sinensis (Shen, 1932) Shantung Peninsular, China
Elamenopsis thorsbornei Lucas & Davie, 1982 Queensland, Australia
Elamenopsis torrensica Lucas, 1980 Torres Straits, Australia
Elamenopsis tuberculata (Chopra & Das, 1930) Cochin, Travancore, India; Sri Lanka
Elamenopsis woodmasoni (Alcock, 1900) Andaman Islands; West Bengal, India; Sri Lanka
Halicarcinus afecundus Lucas, 1980 northern Australia
Halicarcinus bedfordi Monthgomery, 1931 northern Australia
Halicarcinus cookii (Filhol, 1885) New Zealand
Halicarcinus coralicola (Rathbun, 1909) Singapore, Thailand, Japan, Taiwan
Halicarcinus hondai (Takeda & Miyake, 1971) Tyukyu Is., Japan; Great Barrier Reef, Australia
Halicarcinus innominatus Richardson, 1949 New Zealand; Tasmania, Australia
88
Table 1. (Continued)
Halicarcinus keijibabai (Takeda & Miyake, 1971) New Caledonia

Halicarcinus messor (Stimpson, 1858) Japan
Halicarcinus minutus (A. Milne Edwards, 1873) New Caledonia
Halicarcinus orientalis Sakai, 1932 Japan
Halicarcinus ovatus Stimpson, 1858 southern Australia
Halicarcinus planatus (Fabricius, 1775) circum-subantarctic
Halicarcinus rostratus (Haswell, 1882) southern Australia
Halicarcinus setirostris (Stimpson, 1858) Japan; China
Halicarcinus tongi Melrose, 1975 New Zealand
Halicarcinus varius (Dana, 1851) New Zealand
Halicarcinus whitei (Miers, 1876) New Zealand
Halicarcinides nuytsi (Hale, 1927) South Australia
Halimena aotearoa Melrose, 1975 New Zealand
Hymenicoides carteri Kemp, 1917 Gangetic Delta, West Bengal, Bangladesh
Hymenosoma depressum Jacquinot, 1853 New Zealand
Hymenosoma hodgkini Lucas, 1980 eastern Australia
Hymenosoma orbiculare Desmarest, 1825 southern Africa, Zanzibar
Limnopilos naiyanetri Chuang & Ng, 1991 Thailand
Neohymenicus pubescens (Dana, 1851) New Zealand
Trigonoplax longirostris McCulloch, 1908 southern Australia
Trigonoplax spathulifera Lucas, 1980 northern Australia
Trigonoplax unguiformis (de Haan, 1839) Japan; Ternate; Gulf of Martaban; Andaman Is.; Natal
3. structure of the rostrum; & Kannupandi, 1988) in clarifying the uncertain

4. form of the dactyli of the walking legs; relationship of Elamena and Trigonoplax.
5. structure of the Milne Edward's apertures. Sexual dimorphism seems to be a trend for
This feature is not very obvious in some some species. This is especially with regards to
specimens and thus should be used with cau- the chelipeds. This is most extreme in Halicarci-
tion. Lucas (1980) used this character prima- nus coralicola, where the males have much longer
rily to separate the genus Elamena from Trigo- chelipeds (at least twice as long) than the females,
noplax. although the carapaces of the females are up to
50% wider. Amarinus laevis also exhibits extreme
Several other characters which have been used in sexual dimorphism where the cheliped of the male
other brachyuran groups, such as the structure of has a pulvinus between the 'fingers'.
the first and second male pleopods (gonopods)
and male abdomen have not proved very useful,
and their use has not resulted in more coherent Ecology and zoogeography
classifications (see Lucas, 1980; Ng, 1988a, 1991).
Larval morphology has also not proven taxonomi- The biology of the Australian hymenosomatids is
cally useful within the family, with most of the perhaps the best studied (e.g. Lucas, 1971; Lucas
described larvae possessing simple structures & Hodgins, 1970a, 1970b; Melrose, 1975;
which Rice (1980) considers as 'degenerate' fea- Walker, 1969), with that of the Asiatic fauna being
tures. However, the distinctive structure of the almost unknown. Considering the recent discov-
zoe a can be used in sorting plankton samples. eries in Southeast Asia, a brief review of what is
One startling zoea character (possession of three known thus far is prudent.
separate carapace plates) recently described for In general, the habitat preferences of the
Elamena cimex (as a Elamena (Trigonoplax» known Southeast Asian species correspond well
however, seems potentially helpful (see Krishnan with what is generally known for the family, with
89
representatives in fresh, brackish and marine wa- Two of the Southeast Asian species are sublit-
ters. Lucas (1980) noted that the type of substrate toral. Elamena mendosa was collected at about
preferred can usually be deduced from morpho- five metres on sandy/muddy substrates over-
logical features of the crabs (Lucas, 1980). This, grown with the green algae Ulva. Associated
however, may be an oversimplification for some brachyuran species include Doclea gracilipes,
taxa. The only known mangal species, Elamenop- D. canalifera, D. ovis, Phalangipus longipes, Para-
sis mangalis, is known solely from mangroves and panope euagora, Halimede ochtodes, Pilumnopeus
adjacent habitats. The first specimens were col- eucratoides, Myomenippe hardwicki, Menippe
lected from hard substrates in mangroves (Ng, rumphii and Portunus pelagicus. The only speci-
1988a), and some were obtained from under rocks men of Elamena globosa known was dredged from
and drift wood on muddy beaches near man- about 20 metres depth on sandy substrate with
groves. They, however, seem to be most common coral pieces. The rounded and globose carapace
in littoral mud pools, especially in high man- is the most extreme yet known for any hymeno-
groves. They are often in the pools at the base of somatid. Other hymenosomatids have much
mangrove trees, but are very cryptic, being almost flatter carapaces.
undiscernible as they are covered by mud and Three of the Southeast Asian species are fresh-
hide just below the substrate surface. Their long, water taxa - Limnopi/os naiyanetri, Cancrocaeca
curved and hooked ambulatory dactyli are used xenomorpha and Amarinus wolterecki. Limnopilos
not only for grasping onto hard substrates like naiyanetri was found in large numbers among
rocks or roots, but also for moving across soft water hyacinths in a large river in Thailand. It has
mud. Collection of a good series of specimens not yet been found elsewhere. The body and ap-
was usually made by squirting dilute formalin into pendages of this species are very hairy and these
the pools, forcing the crabs to move. On capture, tend to gather mud and detritus. This may serve
some specimens 'play dead' for several minutes to camouflage the animal from would-be preda-
and are extremely difficult to distinguish from the tors. Elamenopsis exigua was described from
mud. Elamenopsis mangalis has remarkable os- brackish water estuarine areas (Kemp, 1917), but
moregulatory faculties - being able to survive for a specimen was recently obtained from among
long periods in concentrations varying from 20- water hyacinth in river waters near the sea from
200% seawater. They are capable of surviving in Thailand. The water was fresh, but still subject to
fresh water for up to a day. tidal influences. The only species truly confined to
The only littoral species is Halicarcinus corali- fresh water is Cancrocaeca xenomorpha from Su-
cola. Thus far, they have been found only in rocky lawesi and Amarinus wolterecki from Philippines.
shores and coral reefs, usually on dead coral rock The former species has the most degenerated eyes
covered with soft mud and scattered clumps of yet known for a cavernicole crab (Ng, 1991). It is
algae. The ambulatory dactyli of this species are the only member of its family known to be a
hooked and help the animal efficiently clamp troglobite (obligate cave dweller). The eggs are
down on the rocks during the often strong tides. large, and the larval development of Cancrocaeca
They are well camouflaged, being often covered is probably suppressed and not associated with
by mud and sometimes small pieces of algae. the sea. Specimens were most common on pieces
Halicarcinus coralicola is the only species in this of driftwood brought into the caves by rivers, and
region known to have varied pigmentations on the species does not seem to have luteophilic ten-
the carapace. This character is undoubtedly im- dencies.
portant in helping the crabs hide among the often Reproduction in the Hymenosomatidae was
richly coloured rock surfaces (sponges, algae, reviewed in detail by Lucas (1980). Egg sizes
etc.). The species has been observed to feed on range from 0.25-0.80 mm, sizes which are rela-
the rich organic detritus accumulations on the tively large in comparison to the small carapaces
rocks. of the adult crabs. The brood size ranges from 20
90
to thousands per egg mass. Such brood sizes, type localities. The species are also habitat spe-
even in the upper range, are small when compared cific and few species occur in more than one
with moderate sized grapsids, like Pachygrapsus biotope. One exception being Halicarcinus plana-
crassipes, which have 48600 eggs (see Hiatt, tus which has a circum-subantarctic distribution
1948). Hymenosomatid species with large egg (Lucas, 1980).
sizes have or are suspected to have suppressed There appears to be a general trend for the
larval developments, and most are freshwater taxa hymenosomatids to enter fresh waters and, in this
(Ng, 1991). Direct development is an extreme case respect, they would be only the fourth group of
of abbreviated larval development, and has been crabs which has done so successfully. The large
reported for three freshwater species, Amarinus number of brackish water species, their ability to
lacustris, Amarinus angelicus and Elamenopsis withstand drastic changes in salinity (euryhalin-
kempi, and one marine species, the South African ity), shortened larval development and lack of
Elamenopsis bovis (fide Barnard, 1950). Another larval structures for a prolonged pelagic phase,
marine species, Halicarcinus afecundus, has been has probably, in a way, 'pre-adapted' them for the
suspected to have direct development (Lucas, fresh water habitat. The freshwater crabs of the
1980). Considering the habits and habitat of Can- family Potamidae, Gecarcinucidae and Parathel-
crocaeca xenomorpha, it is likely that it too has phusidae are dominant in this respect, all the spe-
direct development. Direct development in crabs cies being wholly fresh water or semiterrestrial
is closely but not always associated with wholly (Ng, 1988b). The Grapsidae has only one genus
freshwater habitats, as is well known in the pota- which has completely mastered the fresh water/
moids, gecarcinucoids and some grapsoids (see terrestrial habitat - the sesarmine genus Geo-
Ng, 1988b, 1991). sesarma (see Ng, 1988b). The distribution of the
Hymenosomatid reproduction seems to have freshwater hymenosomatids (summarised from
evolved towards larger eggs with fewer pelagic Holthuis, 1968, 1982; Lucas, 1980; Abele, 1972;
larval stages to limit the dispersal of their off- Ng, 1991), all of which are fully aquatic, is very
spring. Hymenosomatid larvae, for those with disjunct:
planktonic stages, pass through three zoeal
stages. A reduction in the number of stages and Elamenopsis kempi (Chopra & Das, 1930) - Iraq
duration in the planktonic phase is generally be- (introduced to Panama)
lieved to aid the animals restrict the dispersion of Elamenopsis introverta (Kemp, 1917) - China
their populations (Rice, 1980; Lim et al., 1986; Elamenopsis inermis (Takeda & Miyake, 1971) -
Tan et al., 1986). The relatively large and well Palau Islands
developed hymenosomatid zoeae have smooth Amarinus pilosus (A. Milne Edwards, 1873) -
carapaces. In some, the rostral and lateral spines New Caledonia
are poorly developed or even absent. These spines Amarinus lacustris (Chilton, 1882) - Australia
are believed to help the larvae float and aid dis- Amarinus angelic us (Holthius, 1968) - Papua New
persion (Broekhuysen, 1955; Rice, 1980). More- Guinea
over, hymenosomatids differ significantly from Amarinus wolterecki (Balss, 1934) - Philippines
other brachyuran crabs in that the megalopal Limnopilos naiyanetri Chuang & Ng, 1991 - Thai-
stage is completely absent, again reducing the pe- land
riod of larval planktonicity. These reproductive Cancrocaeca xenomorpha Ng, 1991 - Sulawesi
tendencies have almost certainly affected their
distribution patterns. Another euryhaline species, Hymenosoma orbicu-
Endemicity seems to be the general distribu- lare Desmarest, 1825, has been reported from a
tional trend in the Hymenosomatidae. Many of lake in South Africa (Allanson et al., 1966; Boltt,
the species, even in the better studied Australa- 1969), but whether the isolated lake-locked popu-
sian region, are not known outside or near their lation is con specific must be checked. Similarly,
91
the authors have examined a specimen of the References

euryhaline Elamenopsis exigua from a Thai river
some distance from the sea. Abele, L. G., 1972. Introductions of two freshwater decapod
crustaceans (Hymenosomatidae and Atyidae) into Central
and North America. Crustaceana 23: 209-218.
Allanson, B. R., B. J. Hill, R. E. Boltt & V. Schultz, 1966. An
Conservation estuarine fauna in a freshwater lake in South Africa. Nature
209: 532-533.
The distribution patterns of the Hymenosoma- Barnard, K. H., 1950. Descriptive catalogue of South African
tidae parallels in many ways that of the true fresh- decapod Crustacea and Pycnogonidae. Ann. s. afr. Mus.
38: 1-837.
water crabs of the families Potamidae, Gecarci- Boltt, R. E., 1969. The benthos of some southern African
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(Grapsidae) (partim). This poses special prob- Lake Sibayi. Trans. r. Soc. S. Afr. 38: 249-269.
lems for their conservation. Broekhuysen, G. J., 1955. The breeding and growth of
Hymenosoma orbiculare Desm. (Crustacea, Brachyura).
Certainly, they are eaten by many fishes and
Ann. s. afr. Mus. 41: 313-343.
probably play a major role in nutrient recycling. Chopra, B. & K. N. Das, 1930. Further notes of Crustacea
Halicarcinus coralicola for example, although a Decapoda in the Indian Museum. I. On two new species
very small crab, can be quite abundant locally. ofhymenosomatid crabs, with notes on some other species.
Habitat specificity (e.g. in Elamenopsis mangalis) Rec. Ind. Mus. 32: 413-429.
also presents challenges, especially in considering Chuang, C. T. N. & P. K. L. Ng, 1991. Preliminary descrip-
tions of one new genus and three new species of hymeno-
the rate of mangrove degradation in Southeast somatid crabs from Southeast Asia (Crustacea: Decapoda:
Asia. Brachyura). Raffles Bull. Zool. 39: 363-368.
The detailed ecology of hymen os om atids is still Hiatt, R. W., 1948. The biology of the lined shore crab,
poorly known, and their role in the marine food Pachygrapsus crassipes Randall. Pacif. Sci. 2: 135-213.
chain not well understood. The state of Southeast Holthuis, L. B., 1968. On Hymenosomatidae (Crustacea De-
capoda Brachyura) from fresh water, with the description
Asian hymenosomatid taxonomy at present is of a new species. Beaufortia 15: 109-121.
such that studies are still in the discovery stage, Holthuis, L. B., 1982. Freshwater Crustacea Decapod of New
as is evident from the large number of new taxa Guinea. In J. L. Gressitt (ed.), Biogeography and Ecology
being discovered. of New Guinea. Monogr. BioI. 42: 603-619.
Kemp, S., 1917. Notes on Crustacea Decapod in the Indian
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Acknowledgements Krishnan, T. & T. Kannupandi, 1988. Larval development of
Elamena(Trigonoplax) cimex Kemp, 1915 in the laboratory:
Some of the specimens obtained in this study The most unusual larvae known in the Brachyura (Crusta-
were the result of the 'A SEAN-Australia Marine cea: Decapoda). Bull. mar. Sci. 43: 215-228.
Lanchester, W. F., 1900. On a collection of crustacea made
Science Project: Living Coastal Resources'. Col- at Singapore and Malacca. Part I. Crustacea Brachyura.
leagues and friends in the Reef Ecology Labora- Proc. zool. Soc. Lond. 1900: 719-770.
tory (N ational University of Singapore) have also Lim, S. S. L., L. w. H. Tan & P. K. L. Ng, 1986. The com-
been generous with their assistance and encour- plete larval development of Pilumnopeus eucratoides Stimp-
agement. Thanks are due to Mrs C. M. Yang son, 1858 (Decapoda, Brachyura, Pilumnidae) in the
laboratory. Crustaceana 50: 265-277.
for permission to examine the hymenosomatid Lucas, J. S., 1971. The larval stages of some Australian spe-
collections in the ZRC. Dr Chou Loke Ming cies of Halicarcinus (Crusatacea, Brachyura, Hymenoso-
kindly provided equipment, laboratory space and matidae). I. Morphology. Bull. mar. Sci. 21: 471-490.
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Singapore Research Grant RP 900360 to the Mus. 33: 148-247.
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55: 274-278. 1-148.
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Malaysia and Singapore. Department of Zoology, National lacustris (Brachyura: Hymenosomatidae) in Australian
University of Singapore, Shinglee Press, Singapore, 156 pp. inland waters. Aust. J. mar. Freshwat. Res. 20: 163-173.
Ng, P. K. L., 1991. Cancrocaeca xenomorpha, new genus and Yang, C. M., 1979. A list of Brachyura in the Zoological
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nesia. Raffles Bull. Zoo!. 39: 59-63. Singapore, 60 pp. (mimeographed).
Male courtship cycles in three species of tropical Ilyoplax crabs

(Decapoda, Brachyura, Ocypodidae)
Takeharu Kosuge 1, Sombat Poovachiranon 2 & Minoru Murai 3

IAmakusa Marine Biological Laboratory, Kyushu University, Tomioka, Amakusa, Kumamoto 863-25,
Japan; 2 Phuket Marine Biological Centre, Phuket 83000, Thailand; 3 Department of Biology, Kyushu
University, Hakozaki, Higashi, Fukuoka 812, Japan
Key words: mating behaviour, Ilyop/ax, Ocypodidae, Malaysia, Thailand, waving display, semilunar cycle
Abstract
The courtship behaviour and cycles of male courtship activity and colouration of Ilyoplax orientalis,
I. delsmani and I. gangetica were studied in the field in Malaysia and Thailand. Each species had a
distinctive chela waving or beckoning display. Depending on species, the chelipeds, carapaces, or both
of waving males blanched to white in contrast to the cryptic colour of nonwaving males and females.
All three of these tropical Ilyoplax exhibited semilunar cycles in male waving activity at the colony level.
It was confirmed for I. orientalis that individual males cycled each semilunar period between waving and
non-waving phases and exhibited different behaviour toward females during these two behavioural
phases.
Introduction 1978; Zucker, 1978, 1983; Greenspan, 1982;

Salmon & Hyatt, 1983).
The genus Ilyoplax includes about 23 species that During our ecological studies of ocypodid crabs
occur throughout the Indo-Western Pacific (Se- in Southeast Asia, we noted that the number of
rene & Lundoer, 1974). In Southeast Asia, these waving males of some Ilyoplax fluctuated periodi-
small ocypodid crabs ( < 10 mm carapace width) cally with the semilunar cycle. Here we describe
typically live in mangrove forests (Macnae, 1968; the waving displays and cycles of waving activity
Sasekumar, 1974). The behaviour and ecology of in Ilyoplax orientalis (De Man, 1888), I. delsmani
tropical Ilyoplax spp. are known primarily from De Man, 1926 and I. gangetica (Kemp, 1919). We
anecdotal notes published in taxonomic reviews also describe the courtship behaviour of I. orien-
of the genus (Kemp, 1919; Tweedie, 1935; 1954). talis.
Males of many species in the family Ocypo-
didae perform conspicuous waving or chela-
beckoning displays. In the fiddler crabs, genus Methods
Uca, this display is a common component of
courtship (Crane, 1957; Salmon, 1983). Daily Study sites and periods
counts of the number of waving males have been
used to quantify lunar and semilunar cycles of This study was done at two locations, 650 km
male fiddler crab courtship activity (e.g. Christy, apart, on the western coast of the Malay Penin-
94
sula. During October and November, 1989, the two plots during December and January. Obser-
waving activity of I. orientalis and I. delsmani and vations of I. delsmani at Ao N am Bor were made
the courtship behaviour of I. orientalis was stud- in a 1.5 x 2 m plot on the mud bank behind the
ied at Sementa, Selangor, near Klang (3 oN, mangroves.
101 0 25' E), Malaysia. The study site was an in- Preliminary observations showed that the
tertidal mud flat and bank at the end of a tidal number of waving males reached a stable daily
channel that was bisected by an artificial earthern maximum during the 2 hour period around the
dam. The mangroves A vicennia alba, Sonneratia time of dead low tide. Hence, each plot was ob-
alba and Rhizophora spp. grew on the upper banks served daily for 5 to 10 minutes during this pe-
of the channel. I. orientalis lived mainly on the riod. Male colour (see results) was used to dint-
mud flat, between MHNT and MLNT at densi- inguish and count the number of waving and non-
ties of 15 to 201m 2 • I. delsmani occurred on both waving I. orientalis. Counts of waving and non-
the mud flat and channel bank. Sasekumar (1974) waving male I. gangetica were based on whether
has described the distributions of Ilyoplax and or not males waved at least once during the obser-
other organisms in this habitat. vation periods. Only waving I. delsmani were
The waving activity of I. gangetica and I. dels- counted because that do not wave are so cryptic
mani was monitored at the Ao N am Bor tidal inlet that they cannot be counted reliably.
(70 51' N, 98 0 25' E) on the southeastern coast Individual cycles of waving activity of I. orien-
of Phuket Island, southern Thailand. Crabs were talis were determined by observing marked males
observed from September 1990 to March 1991, daily for 11 days beginning 26 October. Males
during the dry season, and again from April to were marked by gluing pieces of polyvinyl tape of
August 1991, during hot and rainy seasons. Dur- different colours and shapes to their carapaces
ing the dry season, waving activity was recorded with 'Aron Alpha' cement. On 4 November, the
at 2 to 7 day intervals and daily from 16 January temporal distribution of waving by 7 marked
to 18 February. Observations were made once or males determined by observing each male once
twice per month during the hot and rainy seasons. every 15 minutes during the entire period of tidal
A sandy mud flat extends seaward of the man- emersion. To measure the amount of time males
grove forest which fringes the shore of the inlet. waved and fed, the duration, to the nearest sec-
I. gangetica occurred at densities of 15 to 201m 2 ond, of each activity was recorded during
on the mud flat. I. orientalis also lived on the flat 5 minute intervals. On 19 and 20 October the be-
but a very low densities (about 21m2) and it was haviour of one male was timed for 5 minutes each
not studied at this site. 1. delsmani lived on a mud 30 minutes that the male was active on the sur-
bank at the landward limit of the mangroves. face. Two males were watched on 3 November
and their behaviour was timed for 5 minutes once
each hour.
Male courtship cycles
Courtship cycles within crab colonies were deter- Courtship behaviour of I. orientalis
mined by counting the number of waving males in
fixed plots. All plots were located where crabs Courtship interactions between free-ranging fe-
occurred at high densities. At Sementa, a 1 x 3 m male and waving male 1. orientalis were observed
plot was established on the mud flat where both directly and recorded in the field. To determine
I. orientalis and 1. delsmani lived. On the mud flat whether waving and non-waving males behaved
at Ao N am Bor, two 2 x 3 m plots for 1. gangetica differently toward females near their burrows,
were set up in November and a third plot was adult females were captured and then released 5
established in February. The third plot was nec- to 10 cm away from the burrows of males that
essary because crab density declined in the first had or had not waved previously. The disturb-
95
ance caused males to enter theIr burrows. When hpeds are flexed in front of the buccal region. One
they emerged, theIr behaviour toward the released cheliped is abruptly raised, unflexed vertically and
females was recorded. Cases in which females left then stopped pointing upward. The body is also
before the males returned to the surface were not raised during this jerky motion. Soon after the
analyzed. first cheliped is moved, the other is unflexed lat-
erally (Fig. la), then raised. Both chelipeds are
then lowered and slightly flexed. Either cheliped
Results may lead the display. The body and chelipeds of
waving males are pale white. Non-waving males
Wavmg display and colour change and females are cryptic dark brown.
Ilyoplax delsmanihas smooth symmetrical wave
Ilyoplax orientalis has an asymmetncal wave (Fig. Ie, d). Both flexed chelipeds are raised to
(Fig. la, b). At the start of the display, both che- maximum elevation, unflexed with tips pointing
Fig 1 Change In male body colour assocIated WIth waving In three troPICal specIes of I1yoplax (a) waving and (b) non waving
I orlentalls at Sementa, (c) waving and (d) non-waving I delsrnam at Sementa, (e) I gangetlca (waving) at Ao Nam Bor.
96
up and then lowered without pause. The body is (Table 1 & Fig. 3). During semilunar periods of
also raised and lowered along with the chelipeds. waving activity, most males then began waving
Waving males blanch to conspicuous pure white, and reduced the time they spent feeding. Waving
while non-waving males and females are coloured continued as the predominant activity until about
dark like the substrate. 1 to 0.5 hours before the tide covered their habitat
Light coloured waving males of both of the when some males again fed briefly before they
above species rapidly become dark and cryptic plugged their burrows.
when they loose their burrows and wander on the Daily observations of marked males showed
surface and when they are captured. that most waved on the same days (Table 2). One
Ilyop/ax gangetica is morphologically similar to male was observed long enough to detect two
I. orientalis but its wave display is very different waving periods separated by 7 days, as expected
(Fig. le). Both flexed chelipeds are unflexed for- if males have semilunar cycles of waving activity
ward, raised until their tips point just above the that last 6 to 8 days.
horizontal and then promptly lowered in nearly
the reverse trajectory that they were raised. The
body is also raised and lowered with the motion Courtship behaviour of Ilyoplax orientalis
of the chelipeds. Unlike the above species, only
the chelipeds of waving males are white. Non- Male I. orientalis oriented toward and directed
waving males and females are cryptically co- repeated waves at wandering females that came
loured. near their burrows. When females approached,
males alternated waves with chela quivering, a
Temporal patterns of waving activity motion that lasted about one second, then
stopped displaying suddenly and entered their
All three species of Ilyoplax exhibited semilunar burrows. Some approaching females then fol-
cycles of waving activity (Fig. 2). At Sementa, lowed males into their burrows. Successful pair
I. orienta/is began waving 2 or 3 days after the full formation occurred when the males, no longer
and new moons and continued for 7 to 8 days showing the white display colour, subsequently
with maximum waving around the quarter moons. reappeared on the surface and plugged their bur-
Males fed on the days they did not wave. At the rows. Mating presumably occurred underground
same location, I. delsmani showed a very similar in males' burrows. Waving always preceded pair
cycle of waving activity with maximum waving formation and, hence, appears to be an essential
just after the quarter moons and about one day element of successful courtship.
later compared to I. orientalis. At Ao Nam Bor, Twenty-seven females were released in front of
waving periods of I. delsmani lasted 6 to 8 days the burrows of waving males. Five females left
with waving peaks just before the quarter moons. before the males reemerged from their burrows.
At this site, waving by this species was seen every In 20 of the remaining 22 cases, the males emerged
month except June. At Ao Nam Bor waving pe- and courted the females as described above. In 2
riods by I. gangetica also lasted 6 to 8 days. Maxi- cases the males emerged and waved but the fe-
mum numbers of waving males usually were re- males entered the burrows first. The males fol-
corded after the full and new moons but before lowed these females and plugged their burrows 8
the quarter moons. This species waved during all and 9 minutes after entry.
months of the year. Males that had not waved before the females
were released responded to them very differently
Waving activity by individual Ilyoplax orientalis (Table 3). In 7 of 20 trials, the males emerged, fed
an did not respond to the females. Males in 6 of
Male I. orientalis usually fed for about 1.5 to the remaining 13 cases directed chela quivering
2 hours after their habitat was emersed by the tide toward the females. In 7 cases males made short,
97
I/)
CJ)
a()-{}O- -
100 E
«I
20
¥,
C)
c
>
«I
~
0 0
i i i ~
20
(t
OCT \0 4) NWV O ()
b
20
V,
0
tn () \0 4) to 0 20
()
-
E 10r,
CJ) OCT NOV
to ...
..4
\
\
\
\
. o
\
-1 -
~ A
i i i i
0
JAN
20 4) to ()
10
FEB
204)
Z
5Od ~ I
100
i I/)
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CJ)
I" \ II
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/ ... I
E
0 i
0 C)
C
0 10 () >
() JAN- 4)
«I
roo;
0 i
0
10 204)
() FEB -
Date
Fig. 2. Daily fluctuation in the number of male Ilyoplax active on the surface (filled cicles), the number of waving males (filled
triangles) and the percentage waving (open circles). At Sementa: (a) I. orientalis, (b) I. delsmani. At Ao Nam Bor: (c) I. delsmani,
(d) and (e) I. gangetica in plots 1 and 2. Lunar phases: . , new moon; (), first quarter; 0, full moon; (), last quarter.
98
Table 1. Temporal distribution of surface activity of individual male [lyoplax orientalis on 4 November, 1990. Abbreviations: f,
feeding; w, waving; bl, wandered without burrow; -, in burrow; ?, unknown.
Crab Time of day

no.
11:45 12:00 12:15 12:30 12:45 13:00 13:15 13:30 13:45 14:00 14:15
1 f f w w w w w w w
2 f f f f w w w w w w
3 f f f f f w w w w
4 f f f f f f w w w w w
5 f f f f bl f w w w w w
6 f f f f f f f f bl f
7 f bl f f w w w w bl w
Crab Time of day

no.
14:30 14:45 15:00 15:15 15:30 15:45 16:00 16:15 16:30 16:45 17:00
1 w w w w
2
3 ? ? ? ? ? ? ? ?
4 w w w w w w w f
5 w w w w f
6 f w w w w w w
7 w w w w w w
Table 2. Daily activity of individual male Ilyoplax orientalis at

Sementa, 26 October-5 November, 1989. Activity was clas-
sified each day as waving (w), feeding (f) or unknown (-).
10:L-+[
a t~I,I----.i,-----,~, . -. ~-. .~,-. . .-~. ,~.n
. . . .~~I~
.. Crab
no.
Date
,~----...I -----,~,. .-. -. ~ ~ ~I~ ..,.-

26 27 28 29 30 31 2 3 4 5
rOJ,--+b ---,.1 ....-..-, 1 w f f f f f w w

2 w f f
r]L-...,.-C~I----..~----,~Ir- '-'-~,. . - -L~.,.- -,

3 f f w w w
4 f f f f f f f
..,.-
5 f f f f f f w
6 f f f f f w
* 10:~[
d t ~I--,-....I,II.......,.......II~II........---II~.~
7
8
f
f
f
f
f
f
f
w
w w
1200 1700 9 f f f w w w w
Time of day 10 f f f f
11 f f f w
Fig. 3. Percent of time spent waving (open bars) and feeding 12 f f f f f f
(filled bars) by male [lyoplax orientalis during sequential
13 f f f f w w
5 minute observation periods on four days at Sementa. Left w f w w w
14
arrows and right arrows show, respectively, the times the
15 f f w
observed area was exposed and covered by the tide. (a), (b) 16 f f w f
and (c): data on three males observed during a peak in waving. f w w w
17
(d): data on one male on a day that males did not wave.
99
Table 3. Responses by waving and non-waving male Ilyoplax Discussion

orientalis to females placed near their burrows. Chi-square
tests were used to test the independence of male response and
behavioural phase. Semilunar cycles of waving activity
(a)
All three species of tropical Ilyoplax showed
Male's phase Wave marked semilunar cycles of male waving activity
and associated colour change. As for fiddler crabs
+
(Crane, 1975; Zucker, 1984), display whitening in
Waving 27 o males probably functions as a visual sexual sig-
Non-waving 2 18 nal. Our studies with marked male I. orientalis
x2 = 10.88, P<O.OOI revealed that these cycles are the result of syn-
chrony among males in their individual cycles of
(b) waving activity. Individual males of the other two
species may also alternate between waving and
Male's phase Retreat into burrow non-waving 'phases'. During the waving phase,
individual males fed at the beginning and,' to a
+
lesser extent, at the end of the activity period.
Waving 20 5* During the middle ofthe activity period they spent
Non-waving o 20 most of their time waving. On these days, males
x2 = 28.99, P<O.OOI waved vigorously at females that were close to
* Two cases which female entered burrow precedent to male their burrows. In contrast, males in the non-
were omitted waving phase spent nearly all their time on the
surface feeding and they responded to females
(c) with aggressive behaviour. Waving in Ilyoplax
clearly functions as a courtship display.
Male's phase Threatening dash
Hitherto, clear phase alternation in male wav-
+ ing activity is known only from tropical forms in
Ilyoplax, or in the subfamily Dotillinae. Warm
Waving o 27 temperate species distributed in Okinawa Island,
Non-waving 7 13 southern Japan, e.g. Ilyoplax pusilla and Tmethy-
P = 0.0012, Fisher's exact probability test pocoelis sp. wave throughout their breeding sea-
sons whenever the weather permits surface activ-
(d) ity (Kosuge, pers. obs.). Perhaps, as has been
implied for fiddler crabs (Crane, 1975; Zucker,
Male's phase Chela-quivering
1978), the limited breeding season of warm tem-
+ perate species may allow the expression of repro-
ductive cycles that are closely timed to particular
Waving 13 14 semilunar or lunar tidal conditions. However, at
Non-waving 6 14 least for Uca, precise timing of hatching relative
x2 = 1.58, not significant. to lunar-related tidal amplitude cycles occurs
(Wheeler, 1978; Christy, 1982; Greenspan, 1982).
Hence, selection favouring female reproductive
threatening dashes toward the females; 2 of these cycles leading to larval courtship cycles, does not
males also waved at the females. No males re- appear to vary in a simple way with latitude. Fur-
treated into their burrows. Except for chela quiv- ther research is needed to understand why tropi-
ering, these differences between waving and non- cal but not warm temperate Ilyoplax exhibit male
waving males are significant (Table 3). courtship cycles.
100
Courtship behaviour of Ilyoplax orientalis research and providing the space and facilities for preparing
the manuscript. This research was supported by Grant-in-Aid
from the Japan Ministry of Education, Science and Culture
The courtship of /. orientalis corresponded to the (Overseas Research No. 01041069) to M. Murai.
'male-first' enticing pattern of I. pusilla (Kosuge,
1989). In I. pusilla, courtship in which the female
enters the male's burrow before the male also is References
common. This 'female-first' pattern was rare in
I. orientalis. The causes and significance of this Christy, J. H., 1978. Adaptive significance of reproductive
variation in the sequence of burrow entry by the cycles in the fiddler crab Uca pugilator: a hypothesis. Sci-
ence 199: 453-455.
sexes are unknown at present. Christy, J. H., 1982. Adaptive significance of semilunar cycles
While waving appears to be a courtship signal of larval release in fiddler crabs (genus Uca): test of an
that attracts females to males and their burrow, hypothesis. BioI. Bull. 163: 251-263.
Crane, J., 1957. Basic patterns of display in fiddler crabs
the function of chela quivering, also a usual com- (Ocypodidae, Genus Uca). Zoologica 42: 69-83.
ponent of courtship, is less clear. Females elicited Crane, J., 1975. Fiddler crabs of the world. Princeton Univ.
chela quivering in male I. orientalis in both the Press, New Jersey, 736 pp.
Greenspan, B. N., 1982. Semi-monthly reproductive cycles in
waving and in the non-waving phases, indicating male and female fiddler crabs, Uca pugnax. Anim. Behav.
that the display is not used only during courtship. 30: 1084-1092.
Waving /. delsmani males also gave this display Kemp, S., 1919. Notes on Crustacea Decapoda in the Indian
Museum. XII. Scopimerinae. Rec. Indian Mus. 16: 305-
when females approached them. Chela quivering 353.
has not been observed in I. gangetica, I. pusilla, or Kosuge, T., 1989. Mating behaviour of I1yoplax pusillus (De-
I. integra (Kosuge, pers. obs.). However, chela capoda: Ocypodidae) (in Japanese with English summary).
Nankiseibutu 31: 25-30.
quivering appears to be the predominant form of Macnae, W., 1968. A general account of the flora and fauna
courtship display in I. obliqua at both Sementa of mangrove swamps in the Indo West Pacific region. Adv.
and Ao Nam Bor. Males would quiver one of mar. BioI. 6: 73-720.
Salmon, M., 1983. Courtship, mating systems and sexual se-
their chelipeds which is held flexed in front of the lection in decapods. In: S. Rebach & D. W. Dunham (eds),
buccal region. Stridulation has been implied in Studies in Adaptation, the Behaviour of Higher Crustacea,
the courtship of /. delsmani (Tweedie, 1954) but J. Wiley and Sons, New York: 143-169.
Salmon, M. & G. W. Hyatt, 1983. Spatial and temporal as-
the acoustic or vibrational signals were not ob- pects of reproduction in North Carolina fiddler crabs (Uca
served in the present study. Clearly, the courtship pugilator Bosc). J. expo mar. BioI. Ecol. 70: 21-43.
displays and sequences of Ilyoplax vary across Sasekumar, A., 1974. Distribution of macrofauna of a Ma-
layan mangrove shore. J. animo Ecol. 43: 51-69.
species. Additional comparative studies are Serene, L. & S. Lundoer, 1974. Observations of the male
needed to detect patterns in form, function and pleopod of the species of [lyoplax Stimpson with a key to
the evolution of these signals and female re- the identification of the species. Phuket mar. bioI. Center
Res. Bull. 3: 1-10.
sponses to them. Tweedie, M. W. F., 1935. Notes on the genus I1yoplax Stimp-
son (Brachyura, Ocypodidae). Bull. Raffles Mus. 10: 53-
61.
Acknowledgements Tweedie, M. W. F., 1954. Notes on grapsoid crabs from the
Raffles Museum, Nos. 3, 4 and 5. Bull. Raffles Mus. 25:
118-127.
We thank H. S. Yong, A. Sasekumar (University of Malaya), Wheeler, D. E., 1978. Semilunar hatching periodicity in the
and U. Bhatia (Phuket Marine Biological Center) for their mud fiddler crab Uca pugnax (Smith). Estuaries 1: 268-
hospitality and assistance through the study. We are much 269.
indebted to J. H. Christy for revising an earlier version of the Zucker, N., 1978. Monthly reproductive cycles in three sym-
manuscript. We are also grateful to S. Goshima, K. Kawai, patric hoodbuilding tropical fiddler crabs (genus Uca). BioI.
T. Koga, M. Matsumasa and S. Takeda for their useful sug- bull. 155: 410-424.
gestions for the field work and to T. Komai and K. Wada for Zucker, N., 1983. Courtship variation in the neo-tropical fid-
dler crab Uca deichmenni: another example of female inci-
helping to identify crabs. P. K. L. Ng and P. Naiyanetr helped
tation to male competition? Mar. Behav. Physiol. 10: 57-
compile the pertinent literature We thank T. Kikuchi, M. 79.
Nishihira, M. Tsuchiya and Department of Biology, Univer- Zucker, N., 1984. Delayed courtship in the fiddler crab Uca
sity of the Ryukyus for permitting Kosuge to participate in this terpsichores. Anim. Behav. 32: 735-742.
Hydrobiologia 285: 101-109, 1994.
Distribution and biodiversity of Singapore gorgonians (sub-class

Octocorallia) - a preliminary survey
Nigel K. C. Goh & L. M. Chou

Department of Zoology, National University of Singapore, 10 Kent Ridge Crescent, Singapore 0511,
Key words: octocorallia, distribution, Singapore, gorgonian, diversity
Abstract
Preliminary surveys of the gorgonian fauna of Singapore show the existence of 6 families, comprising
at least 11 genera and 21 species. The distribution of this fauna is closely related to the availability of
a suitable substrata for settlement and growth. In addition, the greatest number of gorgonians, in terms
of abundance and species richness, is found on the lower slopes and bottoms of the reefs in Singapore.
The 'intermediate disturbance hypothesis' is applied to the data obtained and shown to be useful in
making inferences on the state of succession of gorgonian communities.
Introduction inflammatory (Fenical, 1987), antineoplastic

(Faulkner, 1984) and anti-fouling (Standing et al.,
Gorgonians represent an important natural re- 1984; Bandurraga & Fenical, 1985; Rittschof
source. In terms of geology, their calcareous skel- et al., 1985; Faulkner, 1986) agents having been
etons are an important component of coral reef extracted from gorgonians.
formations (Bayer, 1961). Biologically, gorgonian Gorgonians are widespread in their occurrence,
communities host a wide variety of associated being found from the Arctic to the Antarctic, from
organisms, e.g., bivalves, gastropods, polycha- intertidal waters down to the deep sea abysses,
etes, barnacles, shrimps, copepods, crabs, echi- and in temperatures of _10 C to 30 0C (Alder-
noderms, fish, bryozoans, hydroids, anemones slade, 1984). More species occur in tropical and
and many other invertebrates (e.g., Bruce, 1970; sub-tropical waters, but virtually all benthic ma-
Castro, 1971; Emson & Woodley, 1987; Patton, rine habitats are exploited. Gorgonians are gen-
1972; Utinomi, 1959). The economic importance erally sensitive to salinity changes but some have
of gorgonians is seen in the gorgonian Corallium been known to tolerate brackish water of 17.2%0
rubrum (commonly known as red coral) which salinity (Bayer, 1961).
was sold for US$ 900 per kg in 1980 (Bayer, Many factors have been implicated in the dis-
1981). The usefulness of natural products from tribution of gorgonians, e.g., water depth (Bayer,
gorgonians has been known for a long time (Hick- 1961), irradiance levels (Grigg, 1974), bottom
son, 1924; South China Sea Institute of Ocean- sediment transport (Yoshioka & Yoshioka,
ology, 1978). More recently, antimicrobial 1989a), availability of substrata (Bayer, 1961;
(Burkholder & Burkholder, 1958; Der Mardero- Jordan, 1989; Kinzie, 1973), topographic relief
sian, 1969), antiviral (Faulkner, 1984), anti- (Yoshioka & Yoshioka, 1989a, 1989b), and
o
tv
-
,
~:"-:..?=t;'
••':':";"'",p
. ;/
~ i;)'
{.. .. e
SINGAPORE
,t. ." o Skm
l Retan Laut
l{l: · -'';;'"'~'''
\:.:~.!'~;!=~~. .....
....:-T).. Terumbu Pandan
~ ~~: :;,. (Cyrene)
~~ '~~i&:
-=:
';~ LzRft
ig;
~':~~:!l"'T.' t~ SE Buran
Terumbu Pempang Laut
~
Sultan Shoal ~IT·!. ~. ::": ~
_.. ~
Lighthoun8 "';."-:?,:-. ~~ 'W', ...;~Kunu
T. P. Tengah (TJ{~! ,..~... .. ~
~
~ ~~; "'~an
P. lIantu
a. P • .1ong ~L~~ttl:un
~Q~~" ~ Inland
.;:.
." P. subar
. ;:," ~~~;f~l~~
.~.:;.,:~~ ~ ;';"':: Laut
;~~lf) ~
'............ ...........~
Be~(ii~.l:g ~i.:"::/r}; ~~1' ~
....)-~~ Bemban ····P. Semakau
Benar
~~i~i;';';:!:;\0) #.t~?
,.,. ,~,.r
-:;; ...::,I~ ~ Coral reefs

N ~
r .:-=~
\i.:.. "P
"
1 0-·', O;',.~.:,.., ~ • Survey sites
e.
Raffles Lighthouse
Fig. 1. See p. 103 for legend.

103
strength of surge action (Birkeland, 1974; Jordan, Crest: 1.5-4.2 m.

1989; Kinzie, 1973). Kinzie (1973) described 10 Slope: 4.5-10.9 m.
main zones of gorgonian growth in West Indian Bottom: 6.3-22.2 m.
reefs. Three zones are recognised in this study:
crest, slope, and bottom. The gorgonian fauna of Along each transect, six 1 m 2 quadrats were
these zones can be distinguished in terms of sampled, with a one metre gap between adjacent
species composition and relative abundance. quadrats. This sample size was found to be the
largest practicable. The gaps between quadrats
assured independence of samples. All gorgonians
Materials and methods within the sample quadrats were counted. Those
specimens that could not be identified in situ were
Fifteen sites were chosen for the distribution study collected and dried for preservation and subse-
of the gorgonians (Fig. 1). These sites were quent extraction of sclerites. At all sites, the na-
chosen such that samples were collected from the ture of the substratum was noted.
northern, southern, eastern and western reefs of Identification of gorgonian specimens is mainly
Singapore's southern islands. All specimens were performed using colonial morphology and spicu-
collected using SCUBA. lar structure. The spicules or sclerites were ex-
At each study site, three 11 m transects were tracted according to the method described by
laid on the reef; at the reef crest, reef slope and Bayer (1961), with minor modifications. After
sea bed. All transects were placed parallel to the dissolution of the non-calcareous elements of the
reef crest. The reef crest is easily located not only coenenchyme by 5 % sodium hypochlorite solu-
by the relief of the reef, but also by the presence tion (clorox), the sclerites were washed in distilled
of massive colonies of scleractinian corals. The water to remove the clorox. The sclerites were
sea bed of most study sites is less than 25 m deep, next immersed in 96 % alcohol and then dried and
and can therefore be sampled effectively and mounted on glass slides using D.P.x. mountant.
safely using SCUBA. On two of the sites sampled,
the sea bed was deeper, and transects could not
be completed without exceeding decompression Results
limits. At these sites, a ledge on the slope at about
22 m was taken to be the sea bed. The depth at The transect counts of the 45 transects are shown
which the reef slope transect was located was in Table 1. A total of 715 specimens were re-
determined from the following formula: corded from all the sites. Out of these, 616 col-
onies (86 %) were found at the bottom, 80 colo-
Bottom depth - Crest dePth] nies (11 %) on the slope and 19 colonies (3 %) on
Crest depth + [
2 the crest. Raffles Lighthouse and SE Buran had
the largest total counts of all the sites, with 123
This ensured that the transect at the slope would specimens recorded at each, giving an average
always be midway between the crest and the density of6.8 colonies/m2. When only the bottom
bottom transects. The depth ranges for the vari- transect is considered, SE Buran has a density of
ous zones are as follows: 19.0 colonies/m2 compared with 17.7 colonies in
Fig. 1. Map of Singapore and the southern islands showing survey sites.
Note: The sites corresponding to the site numbers used in the tables are given below:
1: Sultan Shoal Lighthouse 5: Terumbu Pempang Tengah 9: P. Semakau 13: Lazarus Island
2: Retan Laut 6: P. Hantu (West) 10: Raffles Lighthouse 14: Kusu Island
3: Terumbu Pandan (Cyrene) 7: P. Hantu (East) 11: P. Jong 15: SE Buran
4: Terumbu Pempang Laut 8: Beting Bemban Besar 12: P. Subar Laut
104
Table 1. Transect counts of sites and zones. the highest species diversity with 14 and 13
species recorded respectively. Intermediate num-
Site Transect counts
bers of species are found at Terumbu Pandan,
Crest Slope Bottom Total Terumbu Pempang Tengah and SE Buran. All
the other sites had species counts ofless than nine
0 1 31 32 (i.e. less than 50%).
2 0 0 5 5
This preliminary survey resulted in 6 families,
3 1 2 27 30
4 0 1 80 81 comprising 11 genera and at least 21 species being
5 3 0 37 40 collected. It should be noted that the species rich-
6 0 0 0 0 ness reported here is greater than that in Tables 2
7 0 0 10 10 and 3 because taxonomic collections were made
8 0 0 0 0
in addition to the surveys studying distribution
9 1 0 1 2
10 0 17 106 123
patterns.
11 0 7 33 40 Seventeen of the species have not been
12 9 21 81 111 recorded previously from Singapore. A complete
13 0 2 32 34 taxonomic account of the gorgonians of Sin-
14 0 25 59 84
gapore is in preparation and will not be discussed
15 5 4 114 123
in detail in this paper. A summary list of the
Total 19 (3%) 80 (11%) 616 (86%) 715 species found in this study is given below:
Sub-Order SCLERAXONIA
each square metre on the bottom transect at Family ANTHOTHELIDAE

Raffles Lighthouse. No specimens were recorded Solenocaulon sp. A
for P. Hantu (West) and Beting Bemban Besar Solenocaulon sp. B
and only two were found on the P. Semakau
Family SUBERGORGIIDAE
transects.
Subergorgia suberosa (Esper)
Table 2 gives the locations and counts of the
various species of gorgonians included in this Family MELITHAEIDAE
study. Almost all the species sampled are found Melithaea sp. A
in the bottom zone of the reef. Exceptions are ?Acabaria sp. A
found in the family Melithaeidae where more
specimens were recorded from the crest and the Sub-Order HOLAXONIA
slope than from the bottom.
Family ACANTHOGORGIIDAE
Most of the species are cosmopolitan within
Acanthogorgia sp. A
the 15 sites except for Acanthogorgia species A
Acanthogorgia sp. B
and Acanthorgorgia species B. The former is re-
Family ELLISELLIDAE
corded at Raffles Lighthouse and P. Subar Laut
Ellisella laevis (Verrill)
while Acanthogorgia species B is recorded only
lunceella sp. A
from Raffles Lighthouse.
lunceella cf. gemmacea
The species richness of the three zones for each
lunceella gemmacea (Valenciennes)
site is summarized in Table 3. The bottom zone
Ctenocella pectinata (Pallas)
contains most of the species found (95 %), while
the crest supports only two (11 %) species. About Family PLEXAURIDAE
half of the total number of species are found Echinomuricea pulcra Nutting
growing on the slope. Echinomuricea indo-malaccensis Ridley
Raffles Lighthouse and P. Subar Laut showed Echinogorgia sp. A
105
Table 2. Distribution and transect counts of gorgonians on Singapore reefs.
Species Sites Total
2 3 4 5 6 789 10 11 12 13 14 15
Subergorgia suberosa Crest o 0 o 0 o 0 000 o o o o 0 o o

Slope o 0 o 0 o 0 000 o o 1 o 4 o 5
Bottom 2 0 6 4 9 0 100 3 o 11 16 14 47 113
Melithaea sp. A Crest o 0 o 0 2 0 000 o o o o 0 o 2
Slope o 0 o 0 o 0 000 1 o o o 0 o
Bottom o 0 o 0 o 0 000 o o o o 0 o o
?Acabaria sp. A Crest o 0 o 1 0 o 0 o o 9 o 0 5 17
Slope o 0 2 0 o 0 000 2 o 18 o 0 o 22
Bottom o 1 o 0 1 0 000 o o 4 o 0 o 6
Verucella sp. Crest o 0 o 0 o 0 000 o o o o 0 o o
Slope o 0 o 0 o 0 000 3 o 1 6 2 13
Bottom 18 0 7 3 1 0 100 56 17 18 2 28 11 162
Ellisella laevis Crest o 0 o 0 o 0 000 o o o o 0 o o
Slope o 0 o 0 o 0 000 1 1 o o 0 o 2
Bottom 1 0 1 0 o 0 200 o 3 6 o 0 o 13
Junceella sp. A Crest o 0 o 0 o 0 000 o o o o 0 o o
Slope 1 0 o 0 o 0 000 4 5 o o 9 o 19
Bottom 6 0 2 7 7 0 501 11 3 15 3 10 10 80
Junceella cf. gemmacea Crest o 0 o 0 o 0 000 o o o o 0 o o
Slope o 0 o 0 o 0 000 2 1 1 o 4 2 10
Bottom 1 0 2 3 4 0 000 4 2 4 2 3 o 25
Ctenocella pectinata Crest o 0 o 0 o 0 000 o o o o 0 o o
Slope o 0 o 0 o 0 000 o o o o 2 o 2
Bottom o 0 1 1 o 0 000 o o 1 o 2 1 6
Acanthogorgia sp. A Crest o 0 o 0 o 0 000 o o o o 0 o o
Slope o 0 o 0 o 0 000 o o o o 0 o o
Bottom o 0 o 0 o 0 000 2 o 3 o 0 o 5
Acanthogorgia sp. B Crest o 0 o 0 o 0 000 o o o o 0 o o
Bottom o 0 o 0 o 0 000 2 o o o 0 o 2
Echinomuricea pulcra Crest o 0 o 0 o 0 000 o o o o 0 o o
Bottom o 3 4 0 1 0 000 o o o 1 0 5 14
Echinogorgia spp. Crest o 0 o 0 o 0 000 o o o o 0 o o
Slope o 0 o 1 o 0 000 4 o o 1 0 o 6
Bottom 1 0 1 19 4 0 100 15 7 1 7 0 17 73
Unidentified species Crest o 0 o 0 o 0 000 o o o o 0 o o
Bottom 2 1 3 43 10 0 000 13 18 2 23 117
Total 32 5 30 81 40 0 10 0 2 123 40 III 34 84 123 715
Note: Count = total number of colonies recorded in six 1 x 1 m quadrats spaced equally on a 11 m transect.
Echinogorgia Sp. B Discussion

?Heterogorgia sp. A
?Heterogorgia sp. B Gorgonians require a firm substratum for attach-
?Heterogorgia sp. C ment and continued growth (Bayer, 1961; Kinzie,
Plexaurid genus A 1973). The sites at Beting Bemban Besar (Site 8)
106
Table 3. Species richness of sites and zones. were recorded at the reef crest zone. This num-
ber increases with depth such that 80 specimens
Site Number of species
(11 %) were found on the reef slope transects and
Crest Slope Bottom Total 616 (86 %) were counted on the bottom transects.
This trend is also evident in terms of species rich-
1 0 1 8 8 ness (Table 3). Only 11 % of all the species found
2 0 0 3 3
were on the reef crest, 47% on the slope and 95 %
3 1 9 10
4 0 1 8 8 on the bottom transect. Generally, gorgonians in-
5 2 0 10 11 terpret the environment in the bottom zone as
6 0 0 0 0 more favourable for colonisation.
7 0 0 5 5 The paucity of gorgonians on the slope com-
8 0 0 0 0
pared to the number on the bottom may indicate
9 0 1 2
10 0 7 11 14 periodic destruction of the fauna on the slope by
11 0 3 6 6 storms necessitating recolonisation by the deeper
12 1 4 13 13 water fauna. Kinzie (1973) proposed this as a
13 0 2 6 6 reason to explain similar observations at Discov-
14 0 5 6 6
ery Bay, Jamaica. Although Singapore is rela-
15 2 10 12
tively sheltered and does not experience storms of
Total 2 (11 %) 9 (47%) 18 (95%) 19 the intensity of hurricanes or typhoons, the reef
Note: Total species richness for all the sites (19) refers only to slope consists mainly of dead coral. As such,
the species encountered within in transect. Three other spe- the substratum is 'loose' and relatively weak
cies were collected apart from those in the transects and are water turbulence can cause dislodgement of rocks
thus not reflected in this table.
or dead corals on which gorgonians may be
attached.
and P. Semakau (Site 9) clearly demonstrate this The fauna of the crest is largely discontinuous
fact (see Table 1). At the former site, the bottom from that of the slope and bottom (Table 2). None
substratum consists exclusively of fine silt up to of the species occurring in the bottom zone, ex-
30 cm in depth. No gorgonians were recorded or cept ?Acabaria sp. A are found on the reef crest.
even observed. However, at the P. Semakau site, Only two species, Melithaea sp. A and?Acabaria
it was noted that submerged concrete pipes less sp. A are found on the slope as well as at the
than 50 metres from the survey site supported an crest. The fauna of the crest may thus be regarded
abundant gorgonian fauna whereas the three as representing species which are able to tolerate
transects only produced two specimens. Thus, the physical and biological environment at the
the limiting factor could only have been the sub- crest.
stratum. Other factors such as temperature, sa- At least two explanations can be put forward
linity, water quality or irradiance (Grigg, 1974; to interpret these observations: competition and
Weinberg, 1978) and sediment loads (Weinberg, ultraviolet tolerance.
1978), parameters which are known to affect gor- The inability of gorgonians to exploit the envi-
gonian distribution, would have been the same for ronment at the crest may be the result of their
the two populations of gorgonians at P. Semakau. inability to compete with the hard corals, sponges
Generally, it was found that sites with suitable and algae for substrata. It is an established fact
substrata for gorgonian growth and settlement that hard corals in Singapore waters have a de-
(Terumbu Pandan, Terumbu Pempang Tengah, creasing abundance with depth and are generally
Raffles Lighthouse, P. Subar Laut and SE Buran) absent below 10 m (see discussion below). On
showed relatively higher species richness (> 10). Singapore reefs, gorgonians colonise the unex-
With reference to Table 1, out of 715 speci- ploited lower slope and bottom where their main
mens collected, only 19 specimens (3 % of total) competitors cannot survive. Interestingly, this is
107
not the case in the Caribbean where prolific cover for Singapore reefs was 41.9% at the 3 m
growth of gorgonians occur in the reefs and depth dropping to 3.09% at 10 m. In addition, at
lagoons adjacent to scleractinian colonies (Bayer, the 10 m depth, the hard coral colonies are small
1961). Short wavelength ultraviolet radiation can and occur infrequently on the transect.
be physiologically and genetically detrimental to Comparing the transect counts (Table 1) and
coral reef epifauna (Jokie1, 1980). The success of species richness (Table 3) of the 15 sites, several
the shallow water reef organisms suggest that they interesting observations arise in relation to the
must have developed some mechanisms of pro- 'intermediate disturbance hypothesis' (Connell,
tection from ultraviolet radiation. Thus, it may be 1978). This hypothesis suggests that diversity is
conceivable that the absence of most of the spe- highest at intermediate levels of disturbance.
cies of gorgonians from the upper parts of the Immediately after a disturbance, only those spe-
reef may indicate a deficiency in this ultraviolet cies which are efficient colonisers would be re-
protection mechanism. For?Acabaria sp. A and cruited. Diversity is postulated to increase with
Melithaea sp. A, their establishment on the crest time after a disturbance, since more time is avail-
could be related to their ultraviolet tolerance or to able for invasion of species with lower powers of
their ability to colonise vertical and even inverted dispersal and growth. Without any subsequent
substrata. It is quite common to observe colonies disturbance, diversity would decrease. This could
of these species growing underneath rock over- be caused by the competitive dominance of some
hangs on the reef crest where there are no hard species to the exclusion of others or by the in-
corals competing except for the ahermatypic cave ability of some species to withstand damage
corals (Tubastraea spp). At the present time, the caused by the physical environment or by natu-
question of what causes the observed restriction ral enemies.
of gorgonians to the lower depths on Singapore At Terumbu Pempang Laut (Site 4) and Kusu
reefs remains unanswered. Island (Site 14), large numbers of colonies are
Gorgonians in Singapore are found in abun- found within the transects, but with a relatively
dance and relatively high species richness at much low species diversity. According to the 'interme-
shallower depths (as shallow as 8 m) than in other diate disturbance hypothesis', the gorgonian com-
reefs in the region (not less than 25 m), e.g. in the munity at these sites could either be at the first
South China Sea (pers. obs.). Grigg (1974) noted stage after a disturbance or at an advanced stage,
the difference in depth distribution of the gorgo- with a prolonged period without any major dis-
nian Corallium at two sites in Hawaii and ex- turbances. Field observations reveal that the age
plained this observation in terms of its tolerance structure of the two sites is different. At Terumbu
to irradiance levels as determined by water clar- Pempang Laut, the colonies are large (hence ma-
ity. It is possible that gorgonians in Singapore are ture) whereas at Kusu Island, juveniles dominate.
able to colonise the shallower depths because Thus, we can infer that the former site has not
of the relatively low light transmission levels in been disturbed for a long period of time while the
Singapore (Chou & Chia, 1986). On the other latter has been subjected to recent disturbances.
hand, an indirect effect of the decreased irradi- This inference agrees well with the development
ance levels could be the restriction of hard corals and reclamation of Kusu Island in 1975 (Chia
to the top 10 m of the reef, resulting in the re- et al., 1988). Terumbu Pempang Laut, on the
moval of competition for substrata at the inter- other hand is a patch reef that has not been
mediate depths. This is supported by work done disturbed to such an extent in recent years.
on the fringing reef at Pulau Hantu, Singapore At Terumbu Pandan (Site 3) and Terumbu
where it was found that the main 'band' of scler- Pempang Tengah (Site 5), relatively smaller num-
actinian growth is within the upper three metres bers of gorgonians were recorded from the
of the reef slope (Chou, 1988). Chua & Chou transects, but the average species count is higher
(1991) found that the average scleractinian coral compared to Terumbu Pempang Laut and Kusu
108
Island. Raffles Lighthouse (Site 10), P. Subar exists on Singapore reefs in two colour variations,
Laut (Site 12) and SE Buran (Site 15) all had white and reddishbrown. The identity of the genus
relatively high counts as well as species richness ?Heterogorgia is uncertain because of the confu-
for the transects. Since both groups of sites sion in the literature, with different species being
(Terumbu Pandan, Terumbu Pempang Tengah identified under the same name. The species of
and Raffles Lighthouse, P. Subar Laut, SE Buran, ?Heterogorgia are similar in that all have thorn-
respectively) show a relatively high species diver- scales as the predominant sclerite form. How-
sity, both groups would be at an intermediate ever, the three species can be distinguished in the
stage between disturbances. The differences in field by their distinct colours. Species A is purple
abundances could be attributed to the availabil- with purple polyps, species B is yellow with yel-
ity of substrata. The importance of this factor for low polyps and species C is yellow with purple
settlement and growth has already been dis- polyps.
cussed. At Terumbu Pandan and Terumbu Pem-
pang Tengah, the bottom consists mainly of sand
with scattered boulders and rocks. The bottom of Acknowledgements
SE Buran also consists of sand and rocks, but
with a higher proportion of rocks. At Raffles The authors are grateful to Dr Philip Alderslade
Lighhouse and P. Subar Laut, the bottom is and Dr Perry Alino for their constructive critique
mainly rocky. Thus, in the former group of sites, of this manuscript. We would like to acknowledge
the lack of suitable substrata would limit the members of the Reef Ecology Study Team, NUS
abundance of gorgonians. This limitation would and friends for help in the field work and Dr Peter
not occur at the latter group of sites. K. L. Ng for advice on taxonomic matters.
Gorgonian taxonomy for this region is known
to be in a- poor state, with major revisions needed
in many families. Until such time when these References
problems are ironed out, species identification re-
mains uncertain. Alderslade, P., 1984. Subclass Alcyonaria. In P. Mather &
In the genus Solenocaulon, two distinct forms I. Bennet (eds), A coral reef handbook, 2nd edn. The
are recognised; one red with white polyps, the Australian coral reef society: 45-48.
Bandurraga, M. M. & W. Fenical, 1985. Isolation of the mu-
other white with white polyps. Subergorgia ap- ricins. Evidence of a chemical adaptation against fouling in
pressa, together with S. Pulcra were synonymised the marine octocoral Muriceafruticosa (Gorgonacea). Tet-
under S. suberosa by Stiasny in 1937. The iden- rahedron 41: 1057-1065.
tification of this specimen was based on S. Bayer, F. M., 1961. The shallow water octocorallia of the
appressa described by Nutting. Species A and B West Indian region. Martinus Nijhoff, The Hague, 373 pp.
Bayer, F. M., 1981. Status of knowledge of octocorals of
of the genus Acanthogorgia resemble A. boninensis world seas. In Seminarios de Biologia Marinha, Academia
Aurivillius and A. dofleini Kukenthal & Gorzaw- Brasileira de Ciencias, Rio de Janiero: 3-11.
sky. However these species have type localities Birkeland, C., 1974. The effect of wave action on the popu-
that are sub-tropical and temperate. On zoogeo- lation dynamics of Gorgonia ventalina Linnaeus. Stud. Trop.
graphical grounds, it is thus unlikely, though Oceanogr. 12: 115-126.
Bruce, A. J., 1970. Report on some commensal pontoniinid
not impossible that their ranges extend to the shrimps (Crustacea:Palaemonidae) associated with an
tropical Singapore waters (Philip Alderslade, Indo-Pacific gorgonian host (Coelenterata:Gorgonacea). J.
pers. comm.). The species of the ellisellid genus Zoo!., Lond. 160: 537-544.
Junceella are differentiated mainly by colonial Burkholder, P. R. & L. M. Burkholder, 1958. Antimicrobial
morphology since their sclerite forms are very activity of horny corals. Science 127: 1174-1175.
Castro, P., 1971. The Natantian shrjmps (Crustacea:
similar. Colour variations occur in Junceella sp. A Decapoda) associated with invertebrates in Hawaii. Pacific
and Junceella cf. gemmacea with colonies taking Science 25: 395-403.
on a red or white colouration. C. pectinata also Chia, L. S., H. Khan & L. M. Chou, 1988. The coastal envi-
109
ronmental profile of Singapore. ASEAN/US Coastal Jokiel, P. L., 1980. Solar ultraviolet radiation and coral reef
resources management project technical publications epifauna. Science 207: 1069-1071.
series 3, ICLARM, Manila, 91 pp. Jordan, E., 1989. Gorgonian community structure and reef
Chou, L. M., 1988. The impact of human influence on the zonation patterns on Yucatan coral reefs. Bull. mar. Sci.
fringing reef of Pulau Hantu, Singapore. Proc. 6th Interna- 45: 673-696.
tional Coral Reef Symposium, Australia 2: 201-205. Kinzie, R. A. III, 1973. The zonation of West Indian gorgo-
Chou, L. M. & L. S. Chia, 1986. The marine environment. In nians. Bull. mar. Sci. 23: 93-155.
L. S. Chia,A. Rahman&D. B. H. Tay{eds), Thebiophysi- Patton, W. K., 1972. Studies on the animal symbionts of the
cal environment of Singapore, Singapore University Press: gorgonian coral Leptogorgia virgulata (Lamarck). Bull. mar.
155-184. Sci. 22: 419-431.
Chua, C. Y. Y. & L. M. Chou, 1991. The scleractinian com- Rittschof, D., I. R. Hooper, E. S. Branscomb & J. D.
munity of southern islands' reefs, Singapore. Proc. Regional Costlow, 1985. Inhibition of barnacle settlement and be-
Symp. on Living Resources in Coastal Areas, Manila, haviour by natural products from whip corals Leptogorgia
Philippines: 41-46. virgulata (Lamarck, 1815). J. chern. Ecol. 11: 551-563.
Connell, J. H., 1978. Diversity in tropical rain forests and South China Sea Institute of Oceanology, Academic Sinica,
coral reefs. Science 199: 1302-1310. 1978. Marine life of medicinal value in the South China
Der Marderosian, A., 1969. Marine pharmaceuticals. J. Sea. Kexue Chubanshe, Guangzhou, 153 pp.
Pharm. Sci. 58: 1-33. Standing, J. D., I. R. Hooper & J. D. Costlow, 1984. Inhibi-
Emson, R. H. & J. D. Woodley, 1987. Submersible and labo- tion and induction of barnacle settlement by natural
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31-45. Utinomi, H., 1959. A new gall-forming barnacle embedded in
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Hydrobiologia 285: 111-121, 1994.
Distribution and abundance of marine wood borers on the west coast of

Peninsular Malaysia
Harinder Rai Singh & A. Sasekumar

Department of Zoology, Faculty of Science, University of Malaya, 59100 Kuala Lumpur, Malaysia
Key words: Subtidal, mid-intertidal (intertidal), teredinids, Martesia striata, Sphaeroma, mangroves
Abstract
Twenty one species of marine wood borers were recorded in this study. There are seventeen species of
bivalves of which sixteen are from the family Teredinidae and one from the family Pholadidae. The
crustaceans comprised the family Sphaeromatidae with three species and the family Limnoriidae with
one species. Seven of the fourteen known genera of the teredinids were recorded. The bivalve wood
borers were widely distributed in the local mangroves. Martesia striata was the most abundant wood borer
at the Degaussing Range jetty, Lumut. Lyrodus pedicellatus was the most abundant shipworm in wooden
panels. Shipworms were abundant on the subtidal panels where the genus Bankia, was most common
while M. striata was abundant on the intertidal panels. M. striata numbers were higher on the top sur-
faces as compared to the under surfaces of wooden panels.
Introduction the Crustacea (fam: Sphaeromatidae, genus:

Sphaeroma and fam: Limnoriidae, genus: Limno-
Marine wood borers or termites of the sea ria); the Amphipoda (fam: Cheluridae, genus
(gribbles, pill bugs, piddocks and shipworms) are Chelura) (Kuhne, 1971); and the Bivalvia (fam:
of ecological interest and economic concern. They Teredinidae and Pholadidae) (Turner, 1971).
are responsible for wood deterioration in man- Fourteen genera of Teredinidae are known
grove, marine and brackish water ecosystems. (Turner, 1966) but only two (Teredo and Bankia)
Their existence was probably first realised by man are common in Malaysian marine and brackish
when he started his maritime activities using waters (Chong, 1979; Tan, 1970). Martesia is the
wooden rafts and boats. only representative of the Pholadidae encountered
Fossil records of the teredinids show that they in this region to date (Chong, 1979).
arose in the Cretaceous about 65 million years Literature on marine wood borers in Malaysian
ago (Turner, 1966). Even as far back as 412 B.C., waters is scarce. The earliest report recorded and
man was already familiar with and was using described Sphaeroma triste (Lancaster, 1902). The
chemicals to fight their ravages (Castagna, 1973). presence of Martesia, Teredo and Bankia was
Pliny, Ovid, Aristophanes and Homer also men- shown in Singapore waters (Walker, 1941;
tioned shipworms in their writings. Columbus on Chong, 1979). Although Limnoria and Sphaeroma
his ocean voyages, lost his ships, the 'Capitan a' were found, it was suggested that they did not
and 'Santiago' to these dreaded 'vermes'. pose a threat (Menon, 1957).
Marine wood borers are grouped into 3 taxa: 'Yood biodeterioration studies in Penang har-
112
bour showed the presence of Lyrodus, Teredo, and mud mixture. Wood litter was abundant on
Bankia and Limnoria (Jones et al., 1972). The sys- the mangrove floor at Sementa, Pulau Klang and
tematics and distribution of teredinids in Sin- Lumut. Destruction of mangroves by human ac-
gapore waters is given by Tan (1970) while Eaton tivity was observed at all the sampling stations.
(1982), showed the presence of one limnoriid, six
teredinids and three pholads in log storage ponds
in Sabah waters. The occurrence of Chelura tere- Degaussing Range jetty, Lumut
brans and Martesia in the local mangroves was
reported by Berry (1972). The record of Limnoria The Degaussing Range jetty is situated in marine
lignorum by Berry (1972) is uncertain since this waters [average salinity, 30 parts per thousand
species is found in cold waters. Six species of (Range: 28 ppt-32 ppt) (Singh, 1991)]. The sea
teredinids, one pholad and one sphaeromatid floor is a mud and sand mixture where the former
were recorded by Jubir (1985) in the mangroves consists mainly of silt that is deposited by flood
at Sementa, Klang. tides from the outflow of Sungai Dinding (Fig. 2).
The west coast of Peninsular Malaysia has an The water becomes turbid due to resuspension of
extensive mangrove coastline (approximately silt from the sea floor during tidal shifts.
98334 hectares, Darus & Haron, 1988) and this
would harbour a large number and species of
wood borers having a wide distribution. This large Materials and methods
store-house of wood acts as a natural reservoir
for larvae, juveniles as well as adult marine wood Mangrove stations
borers.
This study gives an account on the species, Three sampling trips (January, February, March
abundance, composition and distribution of ma- 1988) were made to each mangrove station while
rine wood borers at five mangrove localities and monthly samples for a period of twelve months
also their distribution and abundance at the De- (1988) were collected from the Degaussing Range
gaussing Range jetty, Lumut, Perak as assessed jetty. Wood samples of healthy mangrove trees
in a 12 month study carried out in 1988. and broken branches showing attack were split
open with a knife to ascertain the presence and
intensity of attack by wood borers. Wood show-
Description of study site ing attack were cut into one foot lengths and
brought to laboratory for analysis to determine
Mangrove stations wood borer species and their tunnel lengths
(where possible).
Five mangrove stations - Sementa, Pulau Klang,
Morib, Kuala Selangor and Lumut were sampled Degaussing Range jetty, Lumut
for marine wood borers (Fig. 1). Sampling was
done in the lower intertidal zone closer to the Wooden panels of Shorea leprosula (dimension:
seaward edge of the mangroves. The flora is pre- 26 cm x 11 cm x 2 cm) were suspended by a
dominantly of A vicennia sp. and Sonneratia sp. length of high tension nylon rope from the De-
The tidal range at Sementa, Kuala Selangor and gaussing Range jetty, Lumut. To ensure that the
Pulau Klang is approximately 5 metres while that panels were kept in a horizontal position, a 1.5 in.
at Lumut and Port Dickson is approximately 3 length PVC (polyvinyl-chloride) pipe (1 in. diam-
metres. eter) was placed between the wooden panels.
The mangrove floor at Sementa and Kuala Twenty ropes, each containing 12 panels were
Selangor is soft and muddy whereas at Morib, placed at the mid-intertidal (total = 240 panels)
Lumut and Pulau Klang it is firmer with a sand and subtidal (total = 240 panels) zones. The pan-
113
+
N
...,
~-
,
,'" , " __ • I
.... ~ .. .lflii
II Mangrove Forest
Fig. 1. Map of Peninsular Malaysia showing the distribution of mangrove forest reserves (source: Anon., 1982) and sampling
stations (a - Lumut, b - Kuala Selangor, c - Sementa, d - Morib and e - Pulau Klang).
els at the subtidal zone were placed approximately tracted specimens were kept in an ethanol (70 %)
0.3 metre below the extreme low water spring tide and glycerine mixture (70:30 vjv).
(ELWS) and hence were always submerged. Pan- For mangrove studies, individual species of
els in the intertidal zone were placed in the mid- shipworms numbering 25 and more in wood
intertidal zone (henceforth referred to as the in- samples were classified as abundant (A) while
tertidal zone) (between 1-1.5 metre above low those numbering 5 and below were classified as
water) and hence were subjected to periods of rare (R). For the Pholadidae, 100 and above
'wetting' and 'drying'. Eight samples were col- individuals were classified as abundant while
lected on a monthly basis from each zone. This 25 and below were classified as rare. For the
was done for a twelve month period. All samples Sphaeromatidae 50 and above individuals in
were packed in plastic bags and brought back to wood samples were classified as abundant while
the laboratory for analysis. 10 and below individuals were classified as rare.
In the laboratory the wooden samples Only the abundant and rare infestations were
(branches and panels) were placed in 75% etha- considered so as to determine mangrove areas
nol (commercial). Wood samples were split open that are either heavily or rarely infested by the
very carefully with a hammer and chisel, and ex- different groups of wood borers.
114
Measurement of abundance (1966 & 1971), Tan (1970), Kuhne (1971), Naylor
(1972) and Cragg & Icely (1982).
For the Degaussing Range jetty study, direct
counts of Martesia striata entrance holes were
made from top, under and side surfaces of Results
wooden panels. Shipworm counts were based on
the number of tunnels observed. Twenty one species of marine wood borers were
recorded: seventeen (17) species of bivalves (six-
Identification of marine wood borers teen from the family Teredinidae and one from
Pholadidae); four (4) species of crustaceans (three
Marine Wood Borers were identified with the aid species from the family Sphaeromatidae and one
of keys in Barnard (1936), Pillai (1961), Turner from the family Limnoriidae) (Table 1). Seven of
+
N
"tI
l>
Z
--:.
Q
"
0
:l:I
0
0
CJ)
r-
%
l>
Z ~
c ."
~
"'(I)"
LEGEND:
GBI SA ND Y 6EAC H
~ F LOOD U DE
-. ~ Eee T .OE
~ MM" GROVE
Fig. 2. Map of study site showing Degaussing Range jetty (A) and mangrove station (B).
115
Table 1. List of marine wood borer species recorded in the the three groups of wood borers, the teredinids
present study. are represented by ten (10) species (Lyrodus pedi-
Phylum: Mollusca cellatus, Dicyathifer manni, Bactronophorus thorac-
Class: Pelecypod a ites, Teredo mindanensis, Teredo furcifera, Teredo
Family: Teredinidae bartschi, Teredothyra sp., Bankia rochi, Bankia
Genus: Bactronophorus Tapparone-Canefri gracilis and Bankia campanellata. The sphaero-
Bactronophorus thoracites (Gould) 1856
matids comprises two species Sphaeroma tere-
Genus: Dicyathifer Iredale
Dicyathifer manni (Wright) 1866 brans and Sphaeroma triste while the pholad is
Genus: Teredothyra Bartsch solely represented by Martesia striata.
Teredothyra sp. M. striata was the most widely distributed
Genus: Teredo Linnaeus wood borer and was found at all five stations
Teredo bartschi Clapp 1923
(Table 2). It was abundant at Lumut, Pulau Klang
T. johnsoni Clapp 1924
T. furcifera von Martens 1894 and Kuala Selangor, mainly in dead branches
T. mindanensis Bartsch 1923 and drift wood lying on the mangrove floor. In the
Genus: Lyrodus Gould mangroves at Kuala Selangor, M. striata was ob-
Lyrodus pedicelatus (Quatrefages) 1849 served in living A vicennia sp. trunks. M. striata
Genus: Psiloteredo Bartsch
was either co-existing with the shipworms or was
Psiloteredo sp.
Genus: Bankia Gray found solitarily in wood.
Bankia gouldi (B artsch) 1908 Among the Teredinidae, L. pedicellatus,
B. orcutti Bartsch 1923 D. manni and B. rochi were the most widely dis-
B. carinata (Gray) 1827 tributed and were found at three stations
B. gracilis Moll 1935
(Table 2). The former two species were abundant
B. campanellata Moll & Roch 1931
B. philippinensis Bartsch 1927 at three stations, while B. rochi was only abun-
B. rochi Moll 1931 dant at two stations. T. mindanensis, T. bartschi,
Family: Pholadidae
T.furcifera, Teredothyra sp. and B. campanellata
Genus: Martesia Sowerby 1824 were collected at only one station and their oc-
Martesia striata (Linnaeus) 1758 currence was rare. This suggests they are uncom-
Phylum: Arthropoda mon species in Malaysian waters. B. gracilis, al-
Class: Crustacea though not widely distributed, but when observed,
Order: Isopoda was abundant in infested wood. B. thoracites was
Family: Sphaeromatidae observed only at Kuala Selangor and together
Genus: Sphaeroma Latreille 1802
with D. manni (at Morib) infested live, as well as
Sphaeroma terebrans Bate 1866
S. triste Heller 1868 dead, mangrove trees. The living mangrove trees
S. walkeri Stebbing 1905 that were attacked were mainly A vicennia sp.
Family: Limnoriidae
B. thoracites and D. manni attacked both healthy
Genus: Limnoria Leach and diseased trees while M. striata attacked
Limnoria sp. mainly diseased trees.
B. thoracites and D. manni were the largest spe-
cies, with tunnel lengths ranging from 60 to 90 cm,
the fourteen known genera of teredinids are with a maximum burrow diameter of approxi-
present (Turner, 1966 has listed 14 genera). mately 2 cm (3/4 in.). Extraction of large wood
borer specimens from living trees was difficult for
Distribution and abundance of wood borers at five three reasons. Firstly, their tunnels run quite deep
mangrove stations below the mudline. Secondly, large specimens
bore deeply into the trunks making it difficult for
Table 2 gives the distribution and abundance of their extraction. Also, mangrove wood is dense
marine wood borers at 5 mangrove stations. Of and very hard.
116
Tab/e 2. Distribution and abundance of marine wood borers at five mangrove stations on the west coast of Peninsular Malaysia.
Species Mangrove stations
Lumut Pulau Klang Sementa Morib Kuala Selangor
Fam: Sphaeromatidae
Sphaeroma triste +(A) + (A)
Sphaeroma terebrans + (A)
Fam: Teredinidae
Lyrodus pedice//atus +(A) + (A) + (A)
Dicyathi fer manni + (A) + (A) + (A)
Bactronophorus thoracites + (A)
Teredothyra sp. +(R) +(R)
Teredo mindanensis +(R)
Teredo bartschi +(R)
Teredo furcifera + (A)
Bankia rochi + (A) + (A) +(R)
Bankia graci/is + (A) + (A)
Bankia campane//ata + (A)
Fam: Pholadidae
M artesia striata + (A) + (A) + (R) +(R) + (A)
+ = Presence of species; - = absence of species.
S. terebrans and S. triste were not common but The results indicate that M. striata prefers the in-
when observed were abundant only in dead wood. tertidal zone while the shipworms prefer the sub-
The crustaceans infested mainly the 'knee' roots tidal zone.
of dead Bruguiera sp. as well as dead branches of The number of species of shipworms however
A vicennia sp. and Sonneratia sp. Attack by did not differ markedly between the subtidal and
Sphaeroma and Martesia was by single species
while the shipworm attack was by multi-species.
The mangroves at Lumut had the highest spe-
cies count of wood borers (9 species) followed by
"'....
Pulau Klang (6 species), Sementa and Morib J:
(4 species) and Kuala Selangor (3 species).
e
=
=
....
~
o
...
~
Distribution and abundance of wood borers at ; 1

Degaussing Range jetty, Lumut .=
~
M. striata was the most abundant wood borer. Its
mean numbers were higher at the intertidal zone Mid-intertidal Subtidal
as compared to the subtidal zone (Fig. 3). Zones
M. striata numbers (mean) were higher on the top
surfaces of wooden panels as compared to the 1m Shipworms _ ~
under surfaces of wooden panels (Fig. 4). This
Fig. 3. Mean Martesia striata and shipworm numbers from
was observed for both the intertidal and subtidal wooden panels for a twelve month period in the mid-intertidal
zones. Shipworm numbers were higher at the and subtidal zones. [N(Mid-Intertidal) = 88 panels; N(Sub-
subtidal zone as compared to the intertidal zone. tidal) = 86 panels].
117
T. bar
.
~
~ 1 Cn JV1- - -
T. joh
~0:
OJ
'0
·~ I "~
II..
§
0
~
c..
'.E
V)
0:
~ L. ped
PSi. sp.
Mid-intertidal Subtidal 0 400 2000
Zones Shipworm numbers
1_ Top surtace _ Under surtace

ID sublidal _ Mid-intertidal
Fig. 4. Mean Martesia striata numbers from top and under
surfaces of woden panels for a twelve month period at Fig. 5. Total shipworm numbers and species abundance from
th e mid-intertidal and subtidal zones. [N(Mid-Intertidal) = 156 wooden panels for a twelve month period at the mid-
88 panels; N(Subtidal} = 86 panels]. intertidal and subtidal zones. [N(Mid-Intertidal) = 72 panels;
N(Subtidal} = 84 panels]
(Number of empty tunnels measured = 2441).
the intertidal zones (12 and 13 species respec- T. bar - Teredo bartschi B. gra - B. gracilis
tively) (Table 3). Only B. rochi was not recorded T. joh - T. johnsoni B. cam - B. campanellata
at the subtidal zone. The most important ship- T. fur - T. furcifera B. phi - B. philippinensis
B. gou - Bankia gouldi B. roc - B. rochi
worm species were indicated to be L. pedicellatus, B. orc - B. orcutti L. ped - Lyrodus pedicellatus
B. campanellata, T.furcifera and T. bartschi in a B. car - B. carinata Psi.sp. - Psiloteredo sp.
decreasing order of abundance (Fig. 5). L. pedi-
Table 3. Shipworm species distribution at the mid-intertidal cellatus was the most abundant shipworm at the
and subtidal zones, Degaussing Range jetty, Lumut, Perak.
jetty site. The genus Bankia was the most com-
Shipworm species Experimental zones mon with seven species collected, followed by
Teredo, with three species. Teredothyra sp., Psilo-
Mid-intertidal Subtidal teredo sp., T. johnsoni, B. gouldi, B. rochi, B. phil-
ippinensis and B. orcutti were uncommon species.
Psiloteredo sp. + +
Lyrodus pedicellatus
Infestation by the shipworms on wood panels
+ +
Teredo furcifera + + was of multi-species type.
Teredo johnsoni + + Only two specimens of S. walkeri were re-
Teredo bartschi + + corded from wooden panels. It was observed co-
Teredothryra sp. + + habiting among fouling organisms and was not
Bankia rochi +
Bankia campanellata
found boring in wooden panels. This was not
+ +
Bankia philippinensis + + surprising as this sphaeromatid is generally con-
Bankia gracilis + + sidered to be a non-woodboring species. Six
Bankia carinata + + specimens of Limnoria were recorded from pan-
Bankia orcutti + + els even though the panels were heavily attacked
Bankia gouldi + + by the bivalve borers (M. striata and shipworms).
+ = present; - = absent. The sphaeromatids and the limnoriids were not
118
important wood borers at the Degaussing Range no surprise. M. striata prefers marine and slightly
jetty, Lumut. brackish waters (Castagna, 1973). Turner &
Johnson (1971) showed that adult M. striata can
survive in salinities as low as 6 ppt. The larvae
Discussion however can only tolerate salinity as low as
10 ppt. This range of salinity tolerance of both the
Of the four families of marine wood borers, the adult and larvae allows it to survive in areas that
Teredinidae and the Pholadidae (M. striata) are are subjected to heavy rains and flooding (Boyle
widely distributed in local marine waters and & Turner, 1976).
mangroves, as compared to the Spaeromatidae The low abundance of M. striata in Sementa
and the Limnoriidae. and Morib was probably a result of competition
for wood with the shipworms. At Morib, most of
the mangrove forests have been reclaimed (reduc-
Mangrove stations ing the amount of wood), and would therefore
intensify competition for wood between M. striata
The absence of Limnoria in the local mangroves and the shipworms. Low abundances of
is not unusual. Limnoria is a true marine animal Sphaeroma, in conjunction with higher Limnoria
(Pillai, 1965), with low tolerance to reduced numbers was observed by Miller (1926) in bay
salinity (Becker, 1971). During monsoons, the pilings. He suggested that Sphaeroma is unable to
salinity in the mangroves can be low, a condition compete with Limnoria.
which does not favor Limnoria. Eltringham (1961) There are 33 species of marine wood borers
showed that Limnoria cannot survive in waters of (Tan, 1970; Eaton, 1982 and present study) in
low salinity, while Miller (1926) stated that the Malaysian and Singapore marine and brackish
distribution of L. lignorum in the San Francisco waters, of which 25 are teredinids. Tan (1970)
Bay was limited by its inability to adapt to lower listed 12 species of shipworms in Singapore
salinity. waters, but stated that some species may be
In the Malaysian waters, Limnoria sp. is prob- synonyms. Forty-two species of shipworms are
ably restricted to marine and coastal waters present in the Australian-Papua New Guinea re-
(Jones el al., 1972). Although Limnoria is found gion (Ibrahim, 1984). Thirty four species occur in
in the local mangroves it is neither abundant nor Papua New Guinea and the seas around the
destructive (Berry, 1972) and is generally not a Solomon Islands and Coral Sea and Bismark
pest (Menon, 1957). In the mangroves of India Archipelago (Rayner, 1983).
nine species of Limnoria are known but are not In India, there are 32 shipworm species, twenty
important wood destroying agents (Nair, 1965). six of which are found in the mangroves (San-
Sphaeroma is euryhaline and can tolerate wide thakumaran, 1983). The total wood borer species
fluctuations in salinity. Thus the presence of recorded in India is fifty five (Santhakumaran
S. terebrans and S. triste in the local mangroves et al., 1985). Wood borers recorded from this
and also along the coastal waters is not surpris- study are also found in mangroves and coastal
ing. Both S. triste and S. terebrans are active wood waters of India, Australia, Indonesia, Singapore,
borers. S. walkeri is not a wood borer (as ob- Philippines and Papua New Guinea (Table 4).
served in this study) and prefers marine condi- The widely distributed wood borers in this study,
tions. Its mandibles and maxilla are not fully M. striata and L. pedicellatus also has wide dis-
chitinised and suggests a weak link to the rasp- tribution in India (Kalyanasundaram & Ganti,
ing of wood (PiIlai, 1965). 1975).
Since the mangrove sampling stations in this Species diversity of wood borers in mangroves
study were all near the seaward edge, having and coastal waters is high in the tropics. There
higher salinity, the abundance of M. striata was of are three possible reasons for this:
119
Table 4. Marine wood borers found in the current study and tion of teredinids is in the marine environ-
their distribution in the tropical Indo-Pacific region.
ment and is correlated with the evolution of
Species Distribution woody plants and as such, involves efficient
use of wood for food and as a protective
Pam: Limnoriidae substrate.
Limnoria sp. India, Australia, Malaysia (iii) Niches of different groups or species of wood
Pam: Sphaeromatidae borers in areas of extensive wood resource
Sphaeroma triste India, Malaysia, Australia do not overlap greatly and this reduces intra
& Papua New Guinea (PNG) and interspecific competition.
S. terebrans India, Malaysia & Australia
S. walkeri India, Malaysia & Australia Species numbers recorded in the Malaysian and
Fam: Teredinidae Singapore waters were lower than in other tropi-
Dicyathifer manni India, Singapore, Australia, cal areas (such as India, Australia and Papua
PNG, Philippines & Malaysia
New Guinea). This is probably due to the lack of
Bactronophorus thoracites India, Singapore, Australia,
Indonesia, Philippines, PNG sampling intensity especially in Malaysian marine
& Malaysia and brackish waters rather than their actual pau-
Lyrodus pedicellatus India, Malaysia, Singapore, city. Occurrence of wood borers in local riverine
PNG, Australia & Philippines conditions with lower salinities has yet to be ex-
Teredo bartschi Australia, India, Indonesia
plored.
& Malaysia
T. mindanensis PNG & Malaysia Nausitora hedleyi, D. manni, L. pedicellatus,
T. furcifera India, Indonesia, PNG M. striata, B. rochi, B. campanellata, B. thoracites,
& Malaysia T.furcifera, S. terebrans and S. annandalei attack
T. matocotana Malaysia & India living mangroves in India (Srinivasan & Mohan,
Psiloteredo sp. Malaysia & India
1973; Santhakumaran & Pillai, 1974; Santhaku-
Bankia campanellata India, PNG & Malaysia
B. carinata India, Indonesia, Singapore, maran, 1983). Only D. manni and B. thoracites
Philippines, PNG & Malaysia were observed attacking living A vicennia sp. in
B. rochi India, Singapore, PNG this study.
& Malaysia
B. gracilis PNG, Singapore & Malaysia
B. gouldi Malaysia
Degaussing Range jetty
Fam: Pholadidae
Martesia striata India, Australia, Malaysia
Singapore, Philippines, PNG
The heavier infestation of M. striata at the inter-
& Indonesia tidal zone implied that either,
Source: Berry (1972); Chong (1979); Eaton (1982); Ibrahim (i) it preferred an intertidal niche or that
(1984). Jubir (1985); Kalyanasundaram & Ganti (1975); (ii) competition for wood space with shipworms
Lancaster (1902); Menon (1957); Nair (1965); Pillai (1961);
in wooden panels at the subtidal zone
Rayner (1983); Santhakumaran et al. (1985); Tan (1970).
restricted M. striata to the intertidal zone.
(i) Uniformly high water temperature through- The maximum intensity of attack by M. striata is
out the year allows for continuous breeding. in the one metre zone above and below the low
Fairly constant water temperature is an im- water mark, below which there is a sharp de-
portant factor that influences the distribution crease (Ganapati & Nagabushanam, 1955). This
of both adults in driftwood and of the free was also observed by Cheriyan (1964), where
swimming larvae (Rayner, 1983). M. striata numbers were observed above the low
(ii) There are extensive areas of mangroves for tide level but in competition with Sphaeroma
propagation and speciation. Hoagland & where the latter is abundant. He stated however,
Turner (1980) stated that the adaptive radia- that numbers were also high below the low water
120
mark but decreased towards the mudline, where and shipworms. Supporting this was the fact that
more teredinids are found. Limnoria sp. were recorded in panels that were
The absence of S. terebrans and S. triste could already heavily attacked by shipworms and
be related to their lower tolerance to higher sa- M. striata.
linities. Pillai (1961) in his monograph of the wood
boring crustaceans ofIndia writes that Sphaeroma
Acknowledgements
prefers estuarine conditions. Singh (1991) showed
that the salinity at the Degaussing Range was
between 28 parts per thousand (ppt) to 33 ppt. This study was undertaken with financial assis-
The high salinity would tend to favour Martesia tance provided by University of Malaya (Vote F).
and hence its higher abundance. With the ab- We thank Mr Jamri Tohid, Mr Mohammad Zuki
& Mr Jalal for field assistance rendered.
sence of competition from Sphaeroma at the in-
tertidal zone, Martesia would be the dominant
wood borer. References
The attack by M. striata at both the intertidal
and subtidal zones observed in this study is in Anonymous, 1982. Annual Report. Forestry Department,
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zone competition would increase and this would
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In this study the cause-effect relationship (com- the wood boring piddock Martesia striata (Linn.) (Mol-
petition) in the settlement pattern of the wood lusca: Bivalvia: Pholadidae). J. expo mar. BioI. Ecol. 22:
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The larger settlement of M. striata on top as Cochin harbour. J. Timb. Dry. Preserv. Assoc. India. 10:
compared to under surfaces of panels suggests 26-33.
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Turner (1976) reported that during settlement, of the tolerance by Sphaeroma (Crustacea: Isopoda) of
M. striata larvae display a bottom seeking behav- CCA-treated timber. Internat. Res. Grp. Wood Preserv.
iour. This then could probably explain the larger Doc. No. IRGjWPj491.
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of Matang mangrove forest reserves in Peninsular Malay-
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A higher settlement of shipworm larvae on upper Area Management in Malaysia. October 25-27, 1988,
surfaces of horizontal panels was also observed Johor Baru.
by Tsunoda (1979) and Walden et al. (1967). Eaton, R. A., 1982. Development of sawmilling, kiln drying
Jones et al. (1972) found moderate numbers of and preservation research in Sabah: Log deterioration
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L. indica in Penang. The low abundance of Lim- Eltringham, S. K., 1961. The effects of salinity upon the boring
noria sp. at the degaussing range, Lumut, could be activity and survival of Limnoria (Isopoda). J. mar. bioI.
due to competition with early settling M. striata Ass. U.K. 41: 755-797.
121
Ganapati, P. N. & R. Nagabushanam, 1955. Notes on Rayner, S. M., 1983. Distribution of Teredinids (Mollusca:
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Ibrahim, J. V., 1984. Wood boring molluscs. Workshop on Santhakumaran, L. N., 1985. Destruction of mangrove veg-
Marine Timber Piled Structures. Monash University, etation by marine wood borers along the Goa coast, with
Australia: 1-13. notes on their distribution in mangroves along the Indian
Jones, E. B. G., H. Kuhne, P. C. Trussel & R. D. Turner, coasts. The Mangroves: Proc. Nat. Sym. BioI. Util. Cons.
1972. Results of an international cooperative programme Mangroves: 492-498.
on the biodeterioration of timber submerged in the sea. Santhakumaran, L. N. & S. R. M. Pillai, 1974. Incidence of
Mat. u. Organismen. 7: 93-118. marine wood borers in mangroves in the vicinity of Bombay
Jubir, H. B., 1985. The role of marine fauna in in situ decom- harbour. J. Timb. Dev. Assoc. India. 20: 18-21.
position of mangrove forest. B. Sc (Hons.) thesis. Dept. of Santhakumaran, L. N., S. U. Bhaskar & V. V. Srinivasan,
Zoology. University of Malaya, Kuala Lumpur, Malaysia, 1985. Occurrence and distribution of marine wood borers
67 pp. of India. J. Indian Acad. Wood Sci. 16: 40-60.
Kalyanasundaram, N. & S. S. Ganti, 1975. The intensity and Singh, H. R., 1991. Distribution and abundance of marine
distribution of marine wood borers at various ports in India. wood borers and fouling associates in the Degaussing
Bull. Dept. Mar. Sci. Univ. Cochin. 7: 637-644. Range, Lumut, Perak. M. Sc. thesis. Department of Zool-
Kuhne, H., 1971. The identification of wood boring crusta- ogy. University of Malaya, Kuala Lumpur, Malaysia,
ceans. In E. B. G. Jones & S. K. Eltringham (eds), Marine 280 pp.
borers, fungi and fouling organisms of wood. Proceedings Srinivasan, V. V. & K. C. Mohan, 1973. Notes on a collection
of the OECD Workshop, Paris: 5-88. of Teredinids infesting wooden structures from south east
Lancaster, W. F., 1902. On the crustacea collected during the coast of India. J. Timb. Dev. Assoc. India. 19: 17-20.
'Skeat Expedition' to the Malay Peninsular. Proc. zool. Tan, W. H., 1970. Some Singapore shipworms (Teredinidae).
Soc. Lond. 27: 363-389. J. Singapore nat. Acad. Sci. 2: 1-13.
Menon, K. D., 1957. A note on marine borers in Malayan Tsunoda, K., 1979. Ecological studies of shipworm attack on
waters. Malay. For. 20: 32-37. wood in the sea water log storage site. Wood. Res. Inst.
Miller, R. C., 1926. Ecological relations of marine wood bor- Kyoto 65: 11-53.
ing organisms in San Francisco. Ecology. 7: 247-254. Turner, R. D., 1966. A survey and illustrated catalogue of the
Nair, N. B., 1965. Marine timber boring organisms of the teredinidae. The Mus. Compo Zool., Harvard University,
Indian coast. J. Bombay nat. Hist. Soc. 62: 120-131. Cambridge, Mass, 265 pp.
Nair, N. B., 1966. Vertical distribution of marine wood boring Turner, R. D., 1971. Identification of marine wood boring
animals in Cochin harbour, South West Coast of India. molluscs. In E. B. G. Jones & S. K. Eltringham (eds),
Hydrobiologia. 27: 248-259. Marine borers, fungi and fouling organisms of wood. Pro-
Naylor, E., 1972. British marine isopods. Synopsis of the ceedings of the OECD Workshop, Paris: 18-64.
British Fauna No.3. Linnean Soc. Lond., 86 pp. Turner, R. D. &A. C. Johnson, 1971. Biology of marine wood
Owen, G., 1953. Vertical distribution of Teredo norvegica. boring molluscs. In E. B. G. Jones & S. K. Eltringham
Nature 171: 484-485. (eds), Marine borers, fungi and fouling organisms of wood.
Pillai, N. K., 1961. Monograph. Wood boring crustacea of Proceedings of the OECD Workshop, Paris: 259-301.
India. Manager of Publications, Govt. of India Press, New Walden, C. C., I. V. F. Allen & P. C. Trussel, 1967. Estima-
Delhi, 61 pp. tion of marine borer attack on wood surfaces. J. Fish. Res.
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Hydrobiologia 285: 123-129, 1994.
New records of Malaysian marine algae
Siew-Moi Phang
Institute of Advanced Studies, University of Malaya, 59100 Kuala Lumpur, Malaysia
Key words: marine algae, Malaysia, new records, taxonomy
Abstract
Checklists of the marine algae of Malaysia have been published. The last checklist included the marine
algae of Singapore. The checklist recorded 212 taxa for the region. This paper lists an additional 7 taxa
of Chlorophyta, 18 taxa of Rhodophyta and 5 taxa of Phaeophyta, which had been left out of the last
checklist. These include marine algae from Pulau Pinang, Pulau Redang and Pulau Sibu. In addition,
new records of Cyanophyta (2 species), Chlorophyta (4 species), Rhodophyta (10 species) and Phaeo-
phyta (3 species) are reported.
Introduction 12 species of Cyanophyta, 13 families, 24 genera

and 74 species of Chlorophyta, 19 families, 39
Collections made during the Preussische Expedi- genera and 59 species of Rhodophyta and 8 fami-
tion nach Ost-Asien from 1860 to 1862 contributed lies, 13 genera and 46 species of Phaeophyta.
to the early records of marine algae in the South-
east Asian region. The algae were enumerated by
Georg von Martens in 1866. During 1899 and Materials and methods
1900, another collection was made in the Indo-
nesian region during the Siboga Expedition. The After the checklist of Phang and Wee (1991) was
records were published as monographs including published, a few other published papers reporting
those on Halimeda (Barton, 1901), the Codiaceae species collected from Pulau Redang (Green,
(Gepp & Gepp, 1911), the Corallinaceae (Weber 1978), Sabah (Doty, 1988), Pulau Pinang (Abbott
van Bosse & Foslie, 1904) and in the 'Liste des et al., 1991; Xia & Abbott, 1987) and Pulau Sibu
Algues du Siboga Part 1 and 2 (Weber van Bosse, (Ahmad & Suzana, 1989) were obtained. This
1913; 1921). paper thus lists an additional 7 taxa of Chloro-
Checklists of the marine algae of Malaysia and phyta, 18 taxa of Rhodophyta and 5 taxa of
Singapore have been published (Phang, 1984, Phaeophyta from these papers. These additions
1986; Phang & Wee, 1991; Teo & Wee, 1983). to the checklist are given in Table 1.
The checklists have been based on all available In addition, from the collection of marine algae
published reports and on collections made by the at the Institute of Advanced Studies, University
authors. Many of the taxa reported, however, of Malaya, 19 new records are now reported.
could not be verified because of absence of de- The specimens have been collected from Port
posited specimens for examination. Dickson, Pulau Tioman, Pulau Tulai, Pulau
The last checklist (Phang & Wee, 1991) of the Langkawi, Pulau Besar, Pulau Perhentian, Kuala
marine benthic algae of Malaysia and Singapore Selangor and Pulau Tenggol in Peninsular Ma-
recorded 212 taxa with 5 families, 13 genera and laysia, and Pulau Labrador and Pulau Sentosa in
124
Table 1 Additions to the checkhst Table 1 (Contmued)
Taxa Collection site Taxa Collection site
CHLOROPHYTA a Ceramium cormeulatum Montagne Pulau Redang

Order CLADOPHORALES 2 Family RHODOMELACEAE
1 Family BOODLEACEAE U Laureneza IInpflcata J Agardh Pulau Redang
• Boodlea eoaeta Murray et de Tom Pulau Redang • Laureneza papillosa (C Agardh) Grevllle, Pulau Redang
2 Family CLADOPHORACEAE Setchell et Gardner
a Chaetomorpha aerea (Dillwyn) KOtzmg Pulau Redang a Herposlphoma secunda (C Agardh) Ambronn Pulau Redang
b Chaetomorpha mlmma Collins et Hervey Pulau Sibu
a Cladophora repens (J Agardh) Harvey Pulau Redang PHAEOPHYTA
d Rhlzoclomum hooken KOtzmg Pulau RedangA Order ECTOCAPALES
(Syn Rhlzoclomum ajneanum Klltzmg) 1 Family ECTOCARPACEAE
d Feldmanma Simplex (Crouan & Crouan) Pulau Redang
Order CAULERPALES
Hamel
1 Family UDOTEACEAE
(Syn Eetoearpus eylmdneus Saunders)
d Udotea cyathyormls Decalsne Pulau Redang
(Syn Udotea subilttorafls Taylor) Order DICTYOTALES
I Family DICTYOTACEAE
Order DASYCLADALES
U Dletyota dlvancata Lamouroux Pulau Redang
1 Family ACETABULARIACEAE
• Lobophora varlegata (Lamouroux) Womersley Pulau Redang
a Acetabulana pusilla Howe Pulau Redang
(Syn Pococklella vanegata (Lamouroux)
Papenfuss)
RHODOPHYTA
d Padma caulescens ThlVY Pulau Redang
Order NEMALIALES
I Family LIAGORACEAE Order FUCALES
U Llagora ceranoldes Lamouroux Pulau Redang I Family SARGASSACEAE
(Syn Llagora leprosa J Agardh) a Turbmarla meostata Barton Pulau Redang
Order CRYPTONEMIALES (Note References a Green, 1978, b Ahmad & Suzana, 1989;
1 Family CORALLINACEAE
C Xla & Abbott, 1987, d Abbott et ai, 1991, e Doty,1988)
a Amphiroa mbulus (Ellis et Solander) Pulau Redang
Lamouroux
• Jama adherens Lamouroux Pulau Redang
a Jama caplilaeea Harvey Pulau Redang
Singapore. The specimens were washed and pre-
• Fosflella dispar Foslie Pulau Redang served in 4 % formalin in seawater and brought
Order GIGARTINALES back to the laboratory for examination. The larger
1 Family GRACILARIACEAE speCImens were made into herb anum specimens.
C Gracllana subl/ils (Xla & Abbott) Pulau Pmang Free-hand sections were prepared and mounted
Xla& Abbott
(Syn Polycavernosa subtlfls Xla & Abbott) in glycerine jelly and stained with 1% aniline blue
C Gracliana eduils (Gmehn) Silva Pulau Pmang for microscopic examination. All specimens are
(Syn Polycavernosa jastlglala Chang & Xla) deposited in the Seaweed Herbarium, Institute of
d Gracliana changll (Xia & Abbott) Pulau Pmang
Abbott, Zhang & Xia Advanced Studies, University of Malaya.
(Syn Polyeavernosa changll Xla & Abbott)
, Gracliana urvlilel (Montagne) Xla & Abbott Sabah, Melaka,
Smgapore Description of new records
(Syn Polycavernosa urVillel (Montagne)
Xla & Abbott) CYANOPHYTA
2 Family SOLIERIACEAE
e Kappaphycus alvarezll (Doty) Doty Sabah
Order HORMOGONALES
Order RHODYMENIALES
I Family CHAMPIACEAE
1. Family NOSTOCACEAE
• Gastroclomum eompressum (Hollenberg) Pulau Redang Calothrix crustacea Schousboe & Thuret
Chang & Xla Pulau Tioman (PSM 645)
(Syn Coeloselra compressa Hollenberg)
Order CERAMIALES Attached to eroded corals; generally epiphytic
1 Family CERAMIACEAE on a variety of substrates; occur at intertidal or
• Call1thammon jelflpponel Howe Pulau Redang splash zones.
• Ceramium fiaeeldum (KOtz ) Ardlssone Pulau Redang
Thallus consists of an expanded blackish-green
125
turf becoming pulvinate and gelatinous; trichomes Attached to corals, rocks, on sand between
terminate in long hairs; sheaths thick and colour- corals; occur at intertidal zone.
less; false branching present; heterocysts more Thallus tufted, 0.5 to 2.5 cm tall; blades crisped,
commonly at base than intercalary. clustered at base; much lobed in older plants;
Plants function as important nitrogen-fixers. blades characterised by dichotomously arranged
C. crustacea epiphytic on Sargassum plants en- 'ribs' as compared to the palmate 'ribs' of A. stel-
able them to grow abundantly. C. crustacea has lata (Wulfen) C. Agardh, the only other species
been known to penetrate skeletal limestone. In found here; thallus attached by rhizoids.
China it is commonly used for food. Distribution: Indonesia, Philippines, Northern
Distribution: Cosmopolitan Australia
Chapman, 1961, p.43-44, Fig. 45; Humm & Cribb, 1985, p.26; Lewis, 1987, p.6; Weber
Wicks, 1980, p. 84, Fig. 29; Islam, 1976, p.74, van Bosse, 1913, p. 26, Fig. 16, 17.
Fig. 433; Tseng, 1984, p.32, pI. 20, Fig. 5;
Umezaki, 1961, p. 95, pI. 17. 2. Family VALONIACEAE
Valonia aegagropila C. Agardh
2. Family STIGONEMATACEAE Pulau Tenggol (PSM 575); Cape Rachado,
Brachytrichia quoyi (C. Agardh) Bornet & Port Dickson (PSM 592, 712, 729, 853, 886);
Flahault Pantai Dickson, Port Dickson (PSM 1052).
Cape Rachado, Port Dickson (PSM 1076)
Attached to corals, rocks, wood; occur at in-
Attached to corals and rocks; generally epi- tertidal to sub-tidal zones.
phytic on a variety of substrates; occur at inter- Thallus consists of dense clusters of dark green
tidal zone. vesicles 5 to 10 mm long, 1 to 3 mm broad;
Thallus is a firm gelatinous mass, ranging from vesicles almost cylindrical branching irregularly
flat to spherical to irregular, and of a few mm to from sides of vesicles but more commonly nearer
several cm in diameter; thallus becomes folded the ends.
and hollow with age; trichomes 4 to 5 /lm in di- Our studies show that V. aegagropila is an
ameter at base becoming broader (6 to 9 microns) effective bioaccumulator of heavy metals. This
before tapering to 1 /lm at tip; heterocysts inter- fact may be due to its coenocytic nature.
calary; false branches at tips form reverse Distribution: Cosmopolitan in warm seas
V-shape. Other species found; V. jastigiata Harvey ex J.
U sed as food in China Agardh Chapman, 1961, p. 98, Fig. 111; Tseng,
Distribution: Cosmopolitan in warm temper- 1984, p. 270, Fig. 3
ate-tropical areas
Humm & Wicks, 1980, p.92, Fig. 34; Tseng, 3. Family CLADOPHORACEAE
1984, p.40, pI. 24 Fig. 5a,b; Umezaki, 1961, Cladophora prolifera (Roth) Kiltzing
p. 82, pI. 13. Pantai Dickson, Port Dickson (PSM 166,
839, 911)
CHLOROPHYTA Submerged and attached to sandy substrate in
tidal pools and in subtidal zone.
Order CLADOPHORALES Thallus tufted, 1 to 4 cm tall; profusely
1. Family ANADYOMENACEAE branched, branching irregularly dichotomous;
Anadyomene plicata C. Agardh dark green in colour; cell wall thick.
Teluk Salang, Pulau Tioman (PSM 642); Distribution: Mauritius, Philippines, Ceylon,
Kampung Mukut, Pulau Tioman (PSM Taiwan, Northern Australia.
1024); Cape Rachado, Port Dickson (PSM Other species found: C. vagabunda (L.) van der
1070) Hoek (Syn.: C.jascicularis (Mert.) Kiltzing,
126
C. inserta Dickie and C. mauritiana Kiitzing), 2. Family GELIDIACEAE

C. nitida Kutzing, C. patentiramea (Mont.) Kutz- Pterocladia caerulescens (Kutzing) Santelices
ing Pasir Hitam, Pulau Langkawi (PSM 404);
Womersley, 1984, p. 193, Fig. 62a Cape Rachado, Port Dickson (PSM 711,
737, 862, 898); Pulau Besar, Melaka (PSM
4. Family UDOTEACEAE 817)
Tydemannia expeditionis Weber-van Bosse
Attached on sand, corals, shells at intertidal
Teluk Salang, Pulau Tioman (PSM 651);
zone.
Pulau Tulai (PSM 1010)
Thallus with stolon and subcylindrical horizon-
On sand between corals at intertidal zone. tal axis; erect blade 0.2 to 0.5 cm broad, 0.5 to
Thallus greyish-green, lightly calcified, erect; 2.0 cm long; branching almost bipinnate; all axes
axis simple, sparingly branched above; 5 to 20 cm flat and submembranous; tetrasporangia in
tall; axis produce contiguous tufts by repeated simple blades on ultimate unmodified branchlets;
dichotomous division of lateral branchlets which branches slightly swollen where cystocarps are
are arranged in whorls on the axis; the axils be- located. Colour usually greenish-brown.
tween branches are wide. Distribution: Pacific Islands (New Caledonia,
Distribution: Indo-west Pacific region, Indo- type locality), Caribbean, east coast of Australia
nesia, Philippines, Taiwan, Northern Australia Cribb, 1983, p. 34, pI. 6 Fig. 3.
Trono & Ganzon-Fortes, 1988, p. 63, Fig. 41;
Tseng, 1984, p. 292, pI. 145, Fig. 2 Order CR YPTONEMIALES
1. Family CORALLINACEAE
Amphiroa foliacea Lamouroux
RHODOPHYTA Cape Rachado, Port Dickson (PSM 77, 858)
Attached on corals at intertidal zone; abun-
Order NEMALIALES
dant at reef edge where it is SUbjected to strong
1. Family BONNEMAISONIACEAE
wave action.
Asparagopsis taxi/ormis (Delile) Trevisan
Thallus white to pink-purple, 3 to 5 cm tall;
Teluk Salang, Pulau Tioman (PSM 1098)
abundantly di- to trichotomously branched; di-
Attached to sand, corals and rocks at intertidal morphic; horizontally creeping branches have
zone. conspicuously-winged segments; segments with
Thallus in feathery tufts; erect shoots arising conspicuous 'midrib'; vertical segments com-
from irregularly ramified creeping stolon, the pressed; sporangial conceptacles scattered over
shoots sparingly branched forming pyramid up to surface of intergenicula.
20 cm tall; branches absent at base but abundant Distribution: Common in tropical waters,
pinnate branches above; branches soft, tapered northern Australia
and long; spermatangia and procarps produced Other species: A.fragilissima (Linnaeus) Lam-
near tips of clavate branches. ouroux, A. rigida Lamouroux
U sed as food in Guangdong, China as 'Haicai'. Cribb, 1983, p. 45-46, pI. 11 Fig. 2,3; Dawson,
Distribution: Widely distributed in the tropics, 1954, p. 430, Fig. 40c.
Mediterranean, Bermuda, West Indies, Pacific
region 2. Family HALYMENIACEAE
Abbott & Hollenberg, 1976, p. 340; Chapman, Halymeniaformosa Harvey ex Kutzing
1961, p. 56-57, Fig. 53; Cribb, 1983, p.28-29, Labrador, Singapore (WYC 5075), Pulau
pI. 4 Fig. 1,2; Islam, 1976, p.57-58, Fig. 370; Sentosa, Singapore (WYC 5636)
Tseng, 1984, p. 64, pI. 35 Fig. 3.
Attached to corals at intertidal zone.
127
Bushy bright red thallus; main axis broadly Order RHODYMENIALES

flattened with almost pinnately branched second- 1. Family RHODYMENIACEAE
ary branches arising from the sides; secondary Botryocladia leptopoda (J. Agardh) Kylin
branches are further twice divided. Pantai Dickson, Port Dickson (PSM 843,
Distribution: Philippines, Taiwan 877, 913, 943, 1045)
Other species; H. durvillaei Bory de Saint-
Attached to mangrove roots, corals, sand and
Vincent, H. microcarpa (Mont.) Silva (Syn.: H.
stones at intertidal zone.
durvillaei var. ceylanica Kiitzing), H. floresii
Thallus up to 20 cm long; axis frequently
(Clemente) C. Agardh
branched bearing hollow vesicles on main thallus
Velasquez et al., 1975, p. 149.
or branches; vesicles broadly-oval, 2 to 10 mm
long, 1 to 5 mm broad; vesicles bear gland cells.
Order GIGARTINALES
Distribution: Indo-west Pacific region, Central
1. Family HYPNEACEAE
America, Northern Australia
Hypnea cornuta (Kiitzing) J. Agardh.
Abbott & Hollenberg, 1976, p. 550; Chapman,
Tanjung Rhu, Pulau Langkawi (PSM 374,
1961, p. 117; Tseng, 1984, p. 118
378)
On sandy-muddy substrate at intertidal zone.
Order CERAMIALES
Thallus caespitose up to 20 cm long; axis with
1. Family CERAMIACEAE
alternate secondary to tertiary branches; all
Spyridiajilamentosa (Wulfen) Harvey
branches bear short spinelike branchlets many
Pulau Perhentian (PSM 525, 541)
with reflexed barbs.
Source of carrageenan. Attached to sand, corals at intertidal zone.
Distribution: Central America, Japan, Mauri- Thallus dull pink to brownish, tufted, 5 to 20
tius, Jamaica, Philippines cm long; rhizoidal disc holdfast; main axis corti-
Other species: H. coenomyce J. Agardh, cated, attenuating upwards; alternately or irregu-
H. musciformis (Wulfen) Lamouroux, H. spinella larly branched; single, spherical, sessile, tetrahe-
(C. Agardh) Kiitzing dral tetrasporangium at node of spine-like
Chapman, 1961, p. 115, Fig. 119; Dawson, ramulus; spermatangia in colourless patches
1954, p. 435, Fig.46c at node; cystocarp surrounded by branched in-
volucre.
2. Family RHIZOPHYLLIDACEAE This species has very variable morphology
Portieria hornemannii (Lyngbye) Silva influenced by the environment.
(Syn.: Chondrococcus hornemannii (Lyngbye) Distribution: Widely distributed in tropical and
Schmitz) subtropical waters, West Indies, Central America,
Kampung Mukut, Pulau Tioman (PSM 669, Bermuda, warmer parts of Atlantic, Mediterra-
1027) nean, Red Sea, Indian Ocean, China
Chapman, 1961, p. 172-173, Fig. 178; Cribb,
Attached to corals and sand at intertidal zone.
1983, p. 94, pI. 26 Fig. 2-4; Tseng, 1984, p. 132,
Thallus deep red to orange; rigid, caespitose;
pI. 69 Fig. 3
attached at base by small disc-shaped holdfast;
branching 4 or more times-pinnate in a disticho-
Wrangelia bicuspidata B0rgesen
alternate manner; subflabellate; branches com-
Pantai Dickson (PSM 190, 957); Tanjung
pressed and irregularly alternate.
Rhu, Pulau Langkawi (PSM 377); Pulau
Distribution: Indo-west Pacific region
Tioman (PSM 992)
Cribb, 1983, p. 35-36, pI. 8 Fig. 2; Tseng, 1984,
p. 70, pI. 38 Fig. 2 Attached to mangrove roots, corals and sandy-
muddy substrate at intertidal zone.
128
Thallus sparsely, irregularly branched; axis up Distribution: Vietnam, Philippines, India,

to 10 cm long, corticated at nodes by entangled Bangladesh, China, Taiwan
filaments; cells of main branch sub-cylindrical, Dawson, 1954, p. 404, Fig. 19a; Misra, 1966,
and bear whorls of determinate branchlets at p. 126; Islam, 1976, p.45-46; Tseng, 1984,
upper ends; determinate branchlets repeatedly p. 188, Fig. 3
subdichotomously branched, ending in double-
spined tips, which may be deciduous; tetrahedral Order DICTYOTALES
sporangia covered by few involucral filaments. 1. Family DICTYOTACEAE
Distribution: Florida, Bahamas, Mauritius, Dictyota friabilis Setchell
Jamaica Sementa, Kuala Selangor (PSM 773, 774,
Chapman, 1961, p. 164, Fig. 171 775, 776)
Attached to mangrove roots, mud.
3. Family RHODOMELACEAE
Thallus forms entangled mass, creeping, over-
Bostrychia tene/la (Lamouroux) J. Agardh
lapping one another; attached by numerous rho-
Pantai Dickson (PSM 845,261); Sementa,
zoids; thallus thin membranous, decumbant,
Kuala Selangor (PSM 770); Kampung
friable, 3 cm or more long, 3 to 6 mm broad;
Kisap, Pulau Langkawi (PSM 786, 787)
branching almost dichotomous; branches with
Attached on mangrove roots and trunks, rocks obtuse apices and undulate margins.
and mud. This species is noted to occupy low-light inten-
Thallus forming mats 1 to 2 cm thick, attached sity habitats.
by rhizoids; intricately branched pinnate or se- Distribution: Cosmopolitan in tropical seas,
cund branchlets bear long, incurved monosipho- Bangladesh, Tahiti, Southeast Asia, Australia,
nous hyaline hairs. Vietnam, Guam, China
This species has very variable morphology and Other species: D. apiculata J. Agardh,
is usually associated with mangroves. D. atomaria Hauck, D. bartayresiana Lamouroux,
Distribution: Widely distributed in warm seas, D. beccariana Zanardini, D. cervicornis Kiitzing,
China, Australia D. ciliolata Ki1tzing, D. mertensii (Martius) Kiitz-
Other species; B. moritziana (Sonder ex Ki1tz- ing (Syn.: D. dentata Lamouroux), D. dichotoma
ing) J. Agardh (Hudson) Lamouroux, D. indica Sonder ex Kiitz-
Chapman, 1961, p. 130, Fig. 135; Cribb, 1983, ing, D. linearis (c. Agardh) Greville, D. pardalis
p. 106-107, pI. 66 Fig. 3-4; Islam, 1976, p.66; Kiitzing.
Tseng, 1984, p. 144, pI. 75 Fig. 3 Dawson, 1954, p. 401, Fig. 16a,b; Islam, 1976,
p.37, pI. 21, Fig. L; Tseng, 1984, p. 194, pI. 98
PHAEOPHYTA Fig. 1
OmerSCYTOSIPHONALES Dictyopteris acrostichoides (J. Agardh) B0rgesen

1. Family SCYTOSIPHONACEAE Cape Rachado, Port Dickson (PSM 213,
Rosenvingea orientalis (J. Agardh) B0rgesen 545); Pantai Dickson (PSM 1048)
Tanjung Rhu, Langkawi (PSM 355)
Attached to corals, sand at intertidal zone.
On sand and attached to corals at intertidal Thallus light brown, thin with irregularly di-
zone. chotomously branched blades with distinct
Thallus yellow-brown, erect, tubular when ma- 'midribs'; sporangia in sori scattered all over blade
ture with abundant irregular, alternate or subdi- surface.
chotomous branches; thallus a twisted entangled Distribution: Philippines, India, Australia
mass; branches cylindrical, 2 to 3 rom diameter, Misra, 1966, p. 147; Womersley, 1987, p. 226,
tapering to apex. Fig.78A
129
(Note: Expedition including a monograph of Flabellariaceae and

Udoteae. Siboga-Expeditie Monographie 62.
Collector's reference: - PSM - Phang Siew-Moi
Green J. P., 1978. A Survey and Proposal for the Establish-
WYC- WeeYeowChin) ment of the Pulau Redang Archipelago National Park.
Report submitted to the World Wildlife Fund, Malaysia.
Humm, H. J. & S. R. Wicks, 1980. Introduction and Guide
Acknowledgements to the Marine Blue-green Algae. John Wiley & Sons, 193 pp.
Islam, A. K. M. N., 1976. Contribution to the Study of the
Marine Algae of Bangladesh. Bibliotheca Phycologia Band
Acknowledgements are due to the International 19. J. Cramer, 253 pp.
Foundation for Science, Sweden and IRPA Pro- Lewis, J., 1987. Checklist and Bibliography of Benthic Ma-
gramme (4/180) of the Ministry of Science, Tech- rine Macroalgae recorded from Northern Australia. III
nology and the Environment, Malaysia, for fund- Chlorophyta. Department of Defence, Materials Research
ing research contributing to this paper. Thanks Laboratory Report MRL-R-1063, 55 pp.
Martens, G. von., 1866. In Die Preussische Expedition nach
are also due to Professor Isabella A. Abbott, Uni- OstAsien; nach amlitchen quellen: Botanischer Theil,
versity of Hawaii and Dr. Gerald T. Kraft, Mel- Berlin.
bourne University for help in confirming the iden- Misra, J. N., 1966. Phaeophyceae in India. Indian Council of
tification of the specimens. The comments and Agricultural Research, New Delhi, 203 pp.
suggestions for improvement of the manuscript in Phang, S. M., 1984. Seaweed Resources of Malaysia. Wal-
laceana 33: 3-8.
Professor Michael J. Wynne's referee report are Phang, S. M., 1986. Malaysia's Seaweed Flora. Proc. Ninth
appreciated. Ann. Seminar Malaysian Society of Marine Sciences: 17-
45.
Phang, S. M. & Y. C. Wee, 1991. Benthic Marine Algae. In
Kiew, R. (ed.), The State of Nature Conservation in
References Malaysia, Malayan Nature Society, Kuala Lumpur: 51-61.
Teo, L. W. & Y. C. Wee, 1983. Seaweeds of Singapore.
Abbott, I. A. & G. J. Hollenberg, 1976. Marine Algae of Singapore University Press, Singapore.
California. Stanford University Press, Stanford, 827 pp. Trono Jr., G. C. & E. T. Ganzon-Fortes, 1988. Philippine
Abbott, I. A., J. Zhang & B. Xia, 1991. Graci/aria mixta, sp. Seaweeds. National Book Store, Manila, 330 pp.
nov. and Other Western Pacific Species of the Genus Tseng, C. K., 1984. Common Seaweeds of China. Science
(Rhodophyta: Gracilariaceae). Pacific Science 45: 12-27. Press, Kugler Publications, Amsterdam/Berkeley, 316 pp.
Ahmad Ismail & Suzana Samsuri, 1989. Alga Marin Pulau Umezaki, I., 1961. The Marine Blue-green Algae of Japan.
Sibu, Pantai Timur, Semenanjung Malaysia: Chlorophyta. Memoirs of the College of Agriculture, Kyoto University
Sains Malaysia 18: 139-153. No. 83, Fisheries Series No.8, 149 pp.
Barton, E. S., 1901. The genus Halimeda. Siboga Expeditie Velasquez, G. T., G. C. Trono Jr. & M. S. Doty, 1975. Algal
Monographie 60. species reported from the Philippines. Philipp. J. Sci. 101:
Chapman, V. J., 1961. The Marine Algae of Jamaica, Pt.! & 115-169.
2. Bulletin of the Institute of Jamaica Science Series No. 12 Weber van Bosse, A., 1913. Liste des algues du Siboga I.
Pt.!, 159 pp., Pt.2, 201 pp. Myxophyceae, Chlorophyceae, Phaeophyceae. Siboga
Cribb, A. B., 1983. Marine Algae of the Southern Great Expedition 59: 1-187.
Barrier Reef - Rhodophyta. Australian Coral Reef Society Weber van Bosse, A., 1921. Liste des Algues du Siboga. II.
Handbook No.2, 173 pp. Rhodophyceae. Siboga Expedition 59: 1-124.
Cribb, A. B., 1985. Marine Algae of the Cape Tribulation Weber van Bosse, A. & M. Foslie, 1904. The Corallinaceae
Area. Qd. Nat. 26: 26-29. of the Siboga-Expedition. Siboga Expeditie Monograph 61:
Dawson, E. Y., 1954. Marine Plants in the vicinity of the 78-110.
Institut Oceanographique de Nha Trang, Viet Nam. Pacific Womersley, H. B. S., 1984. The Marine Benthic Flora of
Science 8: 373-471. Southern Australia, Pt. 1. Government Press, Adelaide,
Doty, M. S., 1988. Prodromus ad Systematica Eucheuma- 329pp.
toideorum: A Tribe of Commercial Seaweeds related to Womersley, H. B. S., 1987. The Marine Benthic Flora of
Eucheuma (Solieriaceae, Gigartinales). In I. A. Abbott (ed.), Southern Australia PUI. Government Printers, Adelaide,
Taxonomy of Economic Seaweeds Vol. II California Sea 484pp.
Grant College Program Report No. T-CSGCP-018: 159- Xia, B. & I. A. Abbott, 1987. New species of Polycavernosa
208. Chang & Xia (Gracilariaceae, Rhodophyta) from the
Gepp, A. & E. S. Gepp, 1911. The Codiaceae of the Siboga western Pacific. Phycologia 26: 405-418.
Hydrobiologia 285: 131-137, 1994.
The community structure of macroalgae in a low shore mangrove forest

in Selangor, Malaysia
Sarala Aikanathan * & A. Sasekumar

Department of Zoology, University Malaya, 59100 Kuala Lumpur, Malaysia; *Present address: nrorld
Wide/or Nature Malaysia, Locked Bag No. 911, lalan Sultan P.O., 46990 Petalinglaya, Malaysia
Abstract
The macro algal communities associated with pneumatophores, basal area of tree trunks and sediment
surface in the mangrove forest at Sementa, Selangor consisted of nine main species. Biomass, frequency
of occurrence and relative cover of the species along a belt transect, showed two major trends, a de-
crease in these parameters in the landward direction for Colpomenia sp. Gracilaria blodgettii and Gracilaria
crassa and an increase in the landward direction for Dictyota dichotoma, Catenella nipae, Rhizoclonium
sp. and Bostrichia radicans. Algal dominance varied with substratum. Pneumatophores were dominated
by Caloglossa lepreurii and sediment surface by D. dichotoma. The 40 cm zone at the base of tree trunks
was dominated by two algal species. The 0-20 cm region above the sediment surface was colonized by
C. nipae, while the 20-40 cm region was dominated by Rhizoclonium sp. The study identified the
importance of substrate in macro algal colonization.
Introduction the stable carbon isotope ratios of consumers.

Deposit feeders assimilated significant quantities
Numerous accounts of mangrove fauna and flora of carbon with the range of ratios of the macroal-
have appeared since 1950 (Lugo & Snedaker, gae and diatoms examined (Rodelli et al., 1984).
1974), but mangrove algal communities have re- Beanland & Woelkering (1983) suggested fre-
ceived little attention. Post (1963, 1964a & b, quency distribution ofmangrove-associated algae
1966; Lambert et al., 1987) gave detailed descrip- in South Australia may be influenced by canopy
tions of various macro algal species and identified cover. Thirty two macro algae taxa were associ-
a bostrychietum as an association of algae com- ated with pneumatophores of mangroves at the
prising several macroalgae such as Bostrychia, New South Wales coast (King & Wheeler, 1985).
Caloglossa, Catenella and Murrayella. The species The mangroves of Spencer Gulf, South Australia,
composition and ecology of macro algae in the are the habitat for 49 macroalgal species includ-
mangroves of east Indonesia have been studied ing 10 Chlorophyta, 2 Cyanophyta, 9 Phaeophyta
by Chihara & Tanaka (1988) and Tanaka & and 28 Rhodophyta (Beanland & W oelkering,
Chihara (1988). Benthic macro algae were found 1982). Davey & Woelkering (1985) showed a de-
to have a total gross primary productivity of crease in occurrence, relative cover and biomass
240 kg O 2 d - 1, which accounted for 12 % of the of algae towards the landward side in Western
total gros s primary productivity of the mangroves Port Bay, Victoria.
in Shurat Arwashie, Israel (Dor & Levy, 1984). Few of the studies mentioned provide quanti-
The trophic importance of algae in the food web tative data on algal community structure, and
of the mangrove ecosystem was demonstrated in have been concerned only with macroalgae
132
attached to pneumatophores. The aim of this Selangor (Fig. 1). The forest was bordered by the
study is to determine the frequency of occurrence, Straits of Malacca on the western side and on the
relative cover and biomass of macro algae on eastern side by a bund which prevents sea water
pneumatophores, tree trunks and sediment sur- from intruding into reclaimed land. Starting at
face along a belt transect from the A vicennia forest the seaward edge and for a distance of about
near the mudflat to the seaward edge of the 300 metres to the bund the forest can be sub-
Rhizophora-Bruguiera mixed forest. divided into three distinct zones:
(i) an A vicennia zone, comprising exclusively of

Methods A vicennia alba.
(ii) a Sonneratia zone, with Sonneratia alba in-
The study site was situated at Kapar Mangrove termingled with A vicennia alba; and
Forest Reserve, 6 km north of Port Klang, (iii) a mixed forest zone on the high shore near
Table 1. The percentage cover, frequency of occurrence and biomass of macroalgae on pneumatophores along the belt transect
in the mangrove forest in Sementa, Selangor. Quadrat A is near the high shore.
Quadrat algae A B c D E F G H J Mean
Percentage cover
Catenella nipae 5.2 4.7 4.3 7.1 1.0 2.23
Caloglossa lepreurii 4.7 4.0 5.65 2.4 2.05 0.2 5.0 2.2 9.6 2.1 3.79
Bostrichia radicans 8.3 1.8 3.5 1.5 0.9 0.2 0.2 1.64
Enteromorpha sp. 0.2 0.02
Cladophora 4.0 0.40
Graci/aria blodgettii 0.2 3.0 0.1 0.33
Caloglossa adnata 0.05 0.7 0.1 0.2 0.6 3.0 0.1 0.47
Colpomenia sp. 0.2 0.1 0.03
Rhizoclonium sp. 0.1 8.2 0.83
Dictyota dichotoma 0.2 0.02
Frequency of occurrence
Catenella nipae 0.4 0.4 0.3 0.3 0.1 0.15
Cladophora 0.1 0.01
Gracilaria blodgettii 0.1 0.1 0.1 0.03
Caloglossa adnata 0.1 0.2 0.1 0.1 0.2 0.1 0.1 0.10
Compomenia sp. 0.1 0.1 0.02
Biomass (mg/cm2)
Catenella nipae 2.332 3.961 2.249 10.55 1.626 2.0718
Cladophora
Gracilaria blodgettii 0.608 10.883 0.233 1.1724
Caloglossa adnata 0.037 0.423 0.100 6.343 0.6903
Colpomenia sp. 0.707 0.032 0.0739
133
mudflat in the A vicennia forest and ended at the

landward edge of the mIxed forest of Sonneratia
and A vicennia. This did not include the landward
PALM
mixed forest of Rhizophora and Bruguiera.
OIL Three parameters were considered: biomass,
ESTATE
cover and frequency of occurrence. All these
measurements were obtained for macroalgae on
pneumatophores, tree trunks (from sediment level
up to a height of 40 cm) and sediment surface.
i
Dry weight biomass was obtained by picking the
algae into aluminium foil 'boats', removing any
-N- flecks of barks, then drying at temperature of
70°C for two days. Biomass has been expressed
as algal weight per unit area of the sample (mg
cm - 2). Frequency of occurrence was obtained by
counting the total number of occurrences for each
species divided by the total number of samples.
Cover data represent the total area occupied by
C.
I;: algae as a percentage of total area sampled in
~ each quadrat. Pneumatophores were considered
I'
\. KAPAR FOREST
as cylinders of 1 cm diameter for calculation of
F. Q. RESERVE their surface area.
I'
I;. Along its length the transect was divided into
\:
f. quadrats of 10 m x 10 m, identified as A through
o 2 km
J from the landward to the seaward end. Ten
pneumatophores were taken from each quadrat
L
' . ._ _ _ _ _ _ _- ' ,
and the algae were collected from 20 random

b'::::-:i MUDFLAT square areas (0.5 m on a side) of the sediment.
I<;> ~ IMANGROVE FOREST
Also in each quadrat samples were taken from
five trees of greater than 5 cm DBH. For these
...... BUND samples a knife was used to peel off 5 cm x 5 cm
=MAIN ROAD squares of the bark at 10 cm heights upward from
the sediment to a maximum of 40 cm. Takmg
FIg 1 Map of the study sIte III the Kapar Forest Reserve, such peelings in equal numbers from the seaward
Selangor. and landward side of the trunks provided eight
samples. All samples from the sections were pre-
served in 4 % formalin for later analyses.
The distances from the sediment surface to the
the bund, predominantly comprised of Bru-
water level as noted on the three trunks was mea-
guiera parviflora and Rhizophora mucronata.
sured at both ends of the transect on a relatively
The tides are semi-diurnal and at extreme low tide high tide. With this information and the Chart
(0 metre on the Chart Datum) exposes a mudflat Datum as a reference, it was determined that the
seaward of the mangrove foreshore for about seaward end of the transect was 3.0 m above (CD)
300 m. The extreme highs of 5.3 m reaches the while the landward end was 4.1 m above. In other
top of the bund. words the sediment surface sloped upward 1.1 m
The study site proper was a transect of 100 m toward the landward end (Fig. 2b).
long and 10 m wide, which began just above the
134
Table 2. The percentage cover, frequency of occurrence and biomass of macro algae on lower tree trunks (up to a height of 40 cm)
along the belt transect in the mangrove forest in Sementa, Selangor. Quadrat A is near the high shore.
Quadrat algae A B C D E F G H J Mean
Percentage cover
Rhizoc/onium sp. 0.1 19.15 25.75 30.55 36.65 20.75 4.95 8.45 21.4 16.76
Caloglossa adnata 3.0 8.85 15.25 7.0 3.95 4.2 3.05 14.3 6.8 1.47 6.78
Caloglossa lepreurii 0.15 2.25 0.55 0.05 0.3 0.33
Bostriehia radieans 6.6 32.30 28.5 39.7 30.2 2.05 3.05 6.75 1.05 14.97
Catenella nipae 44.75 29.7 11.6 15.3 4.25 0.5 10.0 4.35 0.05 12.05
Cladophora 0.3 0.05 4.6 0.39
Colpomenia sp. 0.45 0.4
Dietyota diehotoma 0.16 0.5 0.06
Rhizoelonium sp. 0.05 0.6 1.35 1.35 0.9 1.05 0.85 0.95 1.1 0.90
Caloglossa lepreurii 0.05 0.15 0.15 0.10 0.04
Catenella nipae 0.6 0.85 1.05 0.75 0.15 0.15 0.25 0.20 0.05 0.41
Cladophora 0.05 0.05 0.15 0.02
Dietyota diehotoma 0.05 0.05 O.oI
Biomass (mg/cm2)
Rhizoclonium sp. 0.060 11.980 12.044 21.472 24.428 6.064 1.016 3.692 9.692 9.0448
Caloglossa lepreurii 0.020 1.092 0.028 0.240 0.336 0.1716
Catenella nipae 51.092 32.172 25.112 17.604 4.660 11.328 3.092 0.068 14.6583
Cladophora
Dietyota diehotoma 0.180 0.440 0.0620
Results occurred in low frequencies on pneumatophores

throughout the transect (Table 1).
The distribution of macro algae along the transect
varied in the three parameters investigated, viz., Macroalgae on tree trunks: Macroalgae were found
frequency of occurrence, percentage cover and growing mostly below the 40 cm mark (Table 2 &
biomass. Changes in the parameters occurred Fig. 2) on tree trunks in the region just above the
gradually. sediment surface. The most dominant macro-
algae species on the lower tree trunks was Rhizo-
Macroalgae on pneumatophores: The most abun- clonium sp., with the highest percentage cover and
dant alga on the pneumatophores was Caloglossa frequency of occurrence, while Catenella nipae had
lepreurii with the highest frequency of occurrence the highest biomass (Table 2).
and percentage cover (Table 1).
Macroalgal distribution on the sediment surface:
Catenella nipae, Caloglossa lepreurii and Bostrichia Dictyota dichotoma and Rhizoclonium sp. in-
radicans were more abundant and frequent at the creased in abundance towards the landward sec-
landward quadrats A, B and C. All these algae tion of the transect while Co/pomenia sp. and
135
2b
::J:
"'::J:C1i
-t
0
Z
-t
"'-t""
"'
""z
c:
a:
w
""
~
>
0
u
""<
0
w "'
3:
l!7 c:
j:! 0
r-
....Z Z
n"'
\oJ
a:
w
Q.
E.
::J:
"'
1.~Z:%~______________--------------------------------------------------------l[4.1 =iC1i
...
-3·0 ~
2a -Z·o "'
- ,..,
- -1-0
~
~
c:
~--~--'I--~---r--~H~--Ir-~G~-'Ir-~F---'--~E~--r-~D~-.r-~(---'-Ir-B~--r-I~Ar-~~o %
QUADRATS [
Fig. 2. (a) Diagram shows the percentage cover of macro algae on the tree trunks upto a height of 40 em above the sediment level
along the 100-metre transect. Key for lowercase letters: a = Caloglossa adnata; b = Cladophora sp.; c = Rhizoclonium sp.;
d = Bostrichia radicans; e = Catenella nipae; f = Caloglossa lepreuii; g = Dictyota dichotoma; h = Colpomenia sp. (b) Diagram indicates
shore profile and location of belt transect sections A to J in relation to diagram (a).
Gracilaria blodgettii showed the opposite trend, Dictyota dichotoma, Catenella nipae, and Graci/aria
increasing seawards (Table 3). This pattern was blodgettii increased in biomass, cover and fre-
persistent for all the parameters measured. quency of occurrence landwards while Colpome-
Gracilaria crassa was found only in tide pools at nia sp., Graci/aria crassa and Graci/aria blodgettii,
the edge of the forest, close to the mudflat. however, showed a decrease in biomass, cover
136
Table 3. The percentage cover, frequency of occurrence and biomass of sediment macroalgae along the belt transect in the
mangrove forest in Sementa, Selangor. Quadrat A is near the high shore.
Quadrat algae A B C D E F G H J Mean
Percentage cover
Dictyota dichotoma 2.2 1.93 0.05 0.1 2.95 1.43 0.09 0.22 0.89
Rhizoclonium sp. 2.0 1.55 0.05 0.36
Gracilaria blodgettii 0.1 0.18 3.5 0.4 0.4 2.52 3.0 1.01
Gracilaria crassa 0.5 2.0 0.25
Colpomenia sp. 0.05 0.04 0.03 0.05 0.18 0.14 1.38 0.18
Bryopsis sp. 0.55 1.03 0.15
Rhizoclonium sp. 0.3 0.5 0.05 0.08
Graci/aria blodgettii 0.05 0.1 0.1 0.1 0.05 0.25 0.4 0.10
Graci/aria crassa 0.1 0.05 0.01
Colpomenia sp. 0.05 0.1 0.05 0.1 0.2 0.2 0.5 0.12
Bryopsis sp.
Biomass (mgjcm2 )
Rhizoclonium sp. 0.0689 0.3051 0.0074 0.0381
Graci/aria blodgettii 0.0133 0.0016 0.6236 0.0427 0.0061 0.2355 9.2526 1.0165
Graci/aria crassa 0.0108 0.0809 12.7657 1.2857
Co/pomenia sp. 0.0041 0.0017 0.0054 0.0062 0.0025 0.0015 0.1957 0.0217
Bryopsis sp. 0.1371 9.0145 0.9151
and frequency landwards. No sediment macroal- Macroalgae do not only compete amongst
gae was found on the sediment in quadrat D themselves for space, but also with barnacles.
which appeared to be the boundary where sea- Barnacles attach to almost any firm structure
ward dominant algae increased in abundance within reach of tidal waters. Some algal species
and landward dominant algae began to decline such as Rhizoclonium sp. are capable of growing
(Table 3). among barnacles while other such as Colpomenia
sp. and Dictyota dichotoma are unable to do so.
Fewer algal species were found on Sonneratia
Discussion and conclusion trunks and pneumatophores as their bark peeled
off easily.
Algal dominance in the mangrove forest was re- The difference in tidal cover at both the ends
lated to their substratum. Algal communities on of the transects caused variations in algal domi-
trees are SUbjected to less tidal inundation com- nance. Sea front quadrats, J and I seemed to have
pared to those on sediment. The morphological promoted macro algal growth which required
structure of the substratum would also affect frequent tidal cover, presence of tidal pools and
macro algal growth. Macroalgae communities on gullies. Landward fringe quadrats on the other
pneumatophores were similar to those on tree hand, supported algal species that are probably
trunks. Similar substratum promoted the growth more adapted to withstand desiccation. They
of similar macroalgae. Macroalgae on the sedi- probably tolerate lower salinities as frequent rains
ment, however, grow on the mud substrate where affected the landward edge of the transect. Bos-
their rhizoids are not easily dislodged by the mild trichia radicans and Rhizoclonium sp. were com-
wave action typical of the mangrove shore. mon species landwardly as they have resilient
137
cell-walls which prevent desiccation (Chapman, algae in the hard-bottom mangal of Sinai. In Por, F. D. &
I. Dor (eds), Hydrobiology of the Mangal. The Ecosystem
1973). The mid quadrats of the transect supported
of the Mangrove Forests. Dr W. Junk Publishers, The
more species compared with those quadrats on Hague: 179-192.
landward and seaward portions of the transect King, R. J. & M. D. Wheeler, 1985. Composition and
(Fig. 2a). Thus, it appears changes in the sur- geographic distribution of mangrove macro algal communi-
rounding environment and the morphological ties in South Wales. Proc. Linn. Soc., N.S.W. 108: 97-
structure of macro algae determine the community 117.
Lambert, G., T. D. Steinke & Y. Naidoo, 1987. Algae asso-
structure. Macroalgae are ubiquitous in the man- ciated with mangroves in southern African estuaries. I.
grove environment and are important as food for Rhodophyceae. S. Afr. J. Bot. 53: 349-361.
deposit feeders and grazers. Lugo, A. E. & S. C. Snedaker, 1974. The Ecology of Man-
groves. Annu. Rev. Ecol. Syst. 5: 39-64.
Post, E., 1963. Zur Verbreitung and Okologie der Bostrychia-
Caloglossa Association. Int. Revue ges. Hydrobiol. Hy-
References drogr. 48: 47-152.
Post, E., 1964a. Bostrychietum aus dem Nationalpark von
Beanland, w. R. & w. J. Woelkering, 1982. Studies on Melbourne. Revue Algol. 3: 242-255.
Australian mangrove algae: II. Composition and geographic Post, E., 1964b. Bostrychia tangatensis Post, synoptische
distribution of communities in Spencer Gulf, South Studie (Anna Weber van Bosse Zum Gedenken) Hydro-
Australia. Proc. R. Soc. Vict. 94: 89-106. bioI. Bull. 24: 584-602.
Beanland, W. R. & W. J. Woelkering, 1983. Avicennia canopy Post, E., 1966. Bostrychietum auf den Philippinen. Hydro-
effects on mangrove algal communities in Spencer Gulf, biologia 27: 344-351.
South Australia. Aquat. Bot. 17: 309-313. Rodelli, M. R., J. N. Gearing, P. J. Gearing, N. Marshall &
Chapman, V. J., 1973. The algae. 2nd edn. Macmillan, A. Sasekumar, 1984. Stable isotope ratio as a tracer of
London. mangrove carbon in Malaysian ecosystem. Oecologia
Chihara, M. & J. Tanaka, 1988. Species composition and (Berlin) 61: 326-333.
ecology of macro algae in mangrove brackish areas of East Tanaka, J. & M. Chihara, 1988. Macroalgal flora in man-
Indonesia. In K.Ogino and M. Chihara (eds), Biological grove brackish areas of east Indonesia. In K. Ogino and
System of Mangroves. Ehime University, Japan: 7-20. M. Chihara (eds), Biological System of Mangroves. Ehime
Davey, A. & W. J. Woeikering, 1985. Studies on Australian University, Japan: 21-34.
mangrove algae. III. Victorian communities: Structure and Tide Tables, 1986. Tide Tables for Malaysia, Singapore and
recolonization in Western Port Bay. J. expo mar. BioI. Ecol. Brunei. The Hydrographer, Royal Malaysian Navy, The
85: 177-190. Ministry of Defence, 50634 Kuala Lumpur, 116 pp.
Dor, I. & I. Levy, 1984. Primary productivity of the benthic
Hydrobiologia 285: 139-150, 1994.
Marine environmental issues of Southeast Asia: state and development
L. M. Chou
Department of Zoology, National University of Singapore, Singapore 0511, Republic of Singapore
Key words: marine, environment, management, resources, development, protection
Abstract
The seas of Southeast Asia play an important role in the economy of the surrounding countries. The
region's constantly expanding coastal population and development has made great demands on marine
resources, with growing evidence seen in the further degradation of the marine environment and con-
tinued exploitation of living as well as non-living resources. Integrated coastal area management has
never been considered in the past while environmental protection measures and policies have largely been
at local or national levels. Implementation of regional study programmes less than 10 years ago and
ratification of international as well as regional agreements aimed at protecting the marine environment
in recent times indicate a more enlightened approach to the problem.
Introduction Physical setting
The seas of Southeast Asia surround land masses Southeast Asian seas stretch over a wide geo-
which support intense as well as rapidly increas- graphical area of 9 million km 2 , or 2.5 percent of
ing coastal development. The expanding popula- the earth's ocean surface (Fig. 1). Filled with is-
tion of the region also places a growing demand lands of varying size, they form the link between
on living and non-living resources, with marine the Pacific and Indian oceans, separate the con-
resources playing a major role. The pace of coastal tinents of Asia and Australia, and are character-
zone development and marine resource utiliza- ised by high diversity habitats favoured by the
tion has in many cases, resulted in the degrada- tropical climate and heavy precipitation that
tion of the marine and coastal environment. Man- transport nutrients from land to sea.
agement plans for such areas were non-existent Indonesia and the Philippines, the two largest
or dismissed by policy makers as impractical and island archipelagos in the world, have more than
a hindrance to development and national eco- 20000 islands combined. Almost all the South-
nomic growth. More recently, there has been a east Asian countries have extensive coastlines and
realisation of the long-term benefits of coastal and numerous offshore islands, most of which are
marine environment management strategies. This coral or volcanic islands. The total length of
paper provides an overview of the state and de- coastline is 92451 km (Table 1) providing a vari-
velopment of the marine environment in South- ety of coastal and nearshore marine ecosystems
east Asia. which is recognised as being greater in Southeast
Asia than in other parts of the Indian Ocean
Table 1. Estimated coastline length of the Southeast Asian The physical oceanographic features of the region
countries (World Resources Institute, 1992).
have been adequately described by Wyrtki (1961)
Country Coastline extent (km) and Soegiarto (1981).
The tropical waters experience little change in
Brunei Darussalam 161 surface temperature and moderate tidal variation.
Cambodia 443 Currents can be strong, however, in many areas
Indonesia 54716
Malaysia 4675
they reverse their direction under the strong in-
Myanmar 3060 fluence of the seasonal monsoons and therefore
Philippines 22540 do not effectively flush pollutants out to the
Singapore 193 oceans. The monsoons give the area definite wet
Thailand 3219 and dry seasons. Soegiarto (1985) describes the
Vietnam 3444
Total 92451
Southeast Asian waters as being ideal for the
study of monsoonal effects on water circulation
and also on the seasonal variation in the physi-
cal, chemical and biological properties.
region (IUCN/UNEP, 1985a). The seas are char- Stratification of the water column occurs in
acterised by both, extensive shallow continental some of the deeper seas but over most of the
shelves and deep basins, trenches and troughs. Sunda and Sahul shelves, the temperature
141
remains uniform throughout the water column. Table 3. Urban population as a percentage of total popula-
tion in Southeast Asian countries (World Resources Institute,
Salinity is variable and lowered in nearshore
1992).
areas, particularly after heavy rainfall. Bays and
channels with restricted circulation also tend to Country Urban population % of total population
have lowered salinity.
Nutrient load in coastal and nearshore waters 1960 1990
is high because of river outflows, and while the
Cambodia 10.3 11.6
average surface primary productivity based on Indonesia 14.6 30.5
limited data indicate low production rates of 1 J-lg Laos 7.9 18.6
C m - 3 h - 1 in the open sea, higher rates are ob- Malaysia 25.2 43.0
tainable in coastal waters (Soegiarto, 1985). The Myanmar 19.3 24.8
Philippines 30.3 42.6
river outflows contribute to the high level of silt
Singapore 100.0 100.0
in coastal waters. Nutrient levels are increased Thailand 12.5 22.6
during the monsoons. In offshore waters, the sur- Vietnam 14.7 21.9
face layers have low nutrient levels (phosphate
content of 0.2 ppm), while the deeper layers have
greater nutrient content (3 ppm). In the tropics, since 1950 and it is projected to grow a further
nutrients remain trapped in the deeper layers un- 63 % (724 million) by the year 2025. Growth rate
like the temperate seas where annual turnover remains high in spite of a slower increase com-
due to seasonal climate variation brings up pared to the 1960s and 1970s. A population shift
bottom nutrients. In tropical open seas, nutrients towards urban centres is apparent (Table 3).
are brought to the surface by upwelling or diver- Most of the region's main cities and towns are
gent water movements at the surface, which are located within the coastal zone, and over 70% of
usually localised or seasonal. the region's popUlation is estimated to be pres-
ently concentrated in coastal settlements. The
continued expansion of the coastal population
Population growth and urbanisation places a heavy strain on the coastal and marine
environment and their resources.
The region's current population of 444 million
(Table 2), represents an increase of over 140%
Living marine resources
Table 2. Population and growth in the Southeast Asian coun-
tries (World Resources Institute, 1992; Kuntjoro, 1987; The wide variation of geomorphological features
UNEP, 1987). together with the physico-chemical, oceano-
graphic and climatic conditions make the region's
Country Population (millions)
seas highly productive and supportive of rich and
1950 1990 2025 extensive marine habitats, and is recognised as
the faunistic center for the entire Indo-Pacific
Brunei Darussalam 0.14 (1971) 0.22 (1985) (IUCNjUNEP, 1985b).
Cambodia 4.4 8.3 14.0
Indonesia 79.5 184.3 285.9
Laos 1.8 4.1 8.6
Malaysia 6.1 17.9 30.1 Coral reefs
Myanmar 17.8 41.7 72.6
Philippines 21.0 62.4 111.5 Southeast Asian seas contain 25 to 30% of the
Singapore 1.0 2.7 3.3 estimated 600000 sq. km of coral reefs worldwide
Thailand 20.0 55.7 80.9
(Smith, 1978). All the morphological reef types
Vietnam 30.0 66.7 117.5
are represented and coral diversity remains high
142
with almost all the known extant coral genera growing and diverse human influence. Excessive
present (Gomez, 1988). The most extensive reefs harvesting of edible fish, shellfish and other reef-
occur in Indonesia and the Philippines. Diversity related organisms for food has led to rapid deple-
of reef fishes and reef-associated organisms is tion of stocks. The marine curio and aquarium
also high. The reef supports both resident and trades have had devasting impacts on the ecosys-
visiting fish species, and is valuable for artisanal tem (Wood & Wells, 1988). The fast removal of
and commercial fisheries. Coral reefs can poten- target species has affected the ecological balance
tially supply 12% of the world's fish catch (Munro of many reefs permanently. Giant clams for ex-
& Williams, 1985). In Sabah, East Malaysia, reef ample, have been depleted from many reefs. De-
fishes make up 25 % of total fish catch (Mathias structive fishing methods such as the muro-ami,
& Langham, 1978). An estimate of 25 % has also blasting, poisoning are detrimental to the ecosys-
been reported for the Philippines (Carpenter, tem. Coral mining weakens the framework of reefs
1977). In Trengganu, West Malaysia, the figure and the activity also destroys live corals and other
reaches 30 % during certain months (De Silva & organisms overlying the limestone foundation.
Rahman, 1982). Mining of buried fossil giant clam shells in Indo-
Apart from fish, other major biological groups nesia is also damaging. Man's activities on land,
of the reef which provide a source offood include such as deforestation, mining and reclamation,
molluscs, crustaceans, echinoderms and seaweed increase the sediment level in the sea which in
(McManus, 1988). Reef resources are also ex- turn affects coral-growth by direct smothering or
ploited for uses other than food. The marine curio reducing light penetration. Sedimentation result-
trade removes shells and coral, while the ing from coastal and marine tin mining in Phuket
aquarium trade demands fish and other inverte- has affected the reefs there (Chansang, 1988).
brates. Most are exported to countries beyond Extensive land reclamation in Singapore since
Southeast Asia. Coral and limestone blocks make 1963 has affected the deeper zones of reef slopes
excellent building and construction materials. In- (Chou, 1988).
creasing attention has been focused in recent Some countries in the region are beginning to
years on the reef ecosystem as a rich source of recognise the importance of this ecosystem and
natural bioactive substances. The region's high have designated reefs as marine reserves in order
biodiversity reefs, with many species still undis- to prevent the further degradation and loss of this
covered, are an enormous potential source of valuable resource.
pharmaceutically important products.
Coral reefs are also valuable in their natural
role as breakwaters while the breakdown of the Fisheries
limestone framework provides a source of sand
for beaches. The visual impact and colourful Fish forms the major source of protein in the
splendor of reefs make them important as a natu- region and there is a heavy dependence on the sea
ral resource for tourism and when properly man- for this resource. Per capita consumption of fish
aged and developed can provide employment and in the Asean countries is high (Table 4), forming
an effective source of revenue. Rates of primary over half of all animal protein consumed. Table 5
production on reefs are high (Lewis, 1981), al- shows annual marine catch amounting to almost
though that of the surrounding waters are low. 8 million tonnes from the region's seas in the late
This is due to the complex but efficient recycling 1980s, representing a significant increase over the
of material within the reef community and to its late 1970s.
ability to fix atmospheric nitrogen (Longhurst & Pauly & Chua (1988) pointed out that annual
Pauly, 1987), thus making them comparable to marine catch in the six Asean countries increased
the tropical rainforest ecosystem. almost fourfold from 1.5 million tons in the early
The region's coral reefs are being impacted by 1960s to 5.5 million tons in the early 1980s. The
143
Table 4. Fisheries consumption in the ASEAN countries degradation, and in some cases loss, of the ma-
(Kent & Valencia, 1985; World Resources Institute, 1992).
rine environment brought about by a rapid in-
Country Per capita consumption (kg(yr) crease in coastal popUlations and development
(Pauly & Chua, 1988). Pauly (1989) points out
Brunei Darussalam 24.1 that classical fisheries development methods
Indonesia 14.0 using incentives such as soft loans, tax rebates,
Malaysia 30.1
etc. can no longer be applied as the fisheries re-
Philippines 33.8
Singapore 39.6 sources of the region have already been over-
Thailand 20.8 fished.
value of fishcatch in Southeast Asia amounts to Mangroves

3 % of the gross national product of these coun-
tries, which is greater than the 1% in most de- Southeast Asian mangroves represent more than
veloped countries throughout the world (Kent & 30% of the world's mangroves and are the most
Valencia, 1985). Most of the catch within the diverse in species composition. Almost half of the
region consists of demersal species, caught by 40 tree species have commercial importance
artisanal as well as commercial fishermen. (Hundloe & Boto, 1990). The areal extent of man-
Indications are that the fisheries stocks in the groves in the Asean countries cover over
region have been overfished. In the Gulf of Thai- 50000 km 2 (Table 6). Their direct value comes
land and west coast of southern Thailand, fish from the traditional use of mangrove wood for
stocks have been exploited at a level of 40 % above fuel and building materials. The bark is used for
the maximum sustainable yield (Arbhabhirama the production of tannin while some species have
et al., 1987). In the Philippines, demersal and edible leaves or fruit (IUCNjUNEP, 1985a & b).
small pelagic fisheries have been declining since Mangrove trees are also used in the production of
the mid 1970s due to excessive effort. In Indo- chipboard and Saenger et al. (1983) identifies
nesia, trawling has been banned since 1980 many other uses of the mangrove ecosystem in
(Sardjono, 1980). The problem appears to have the region.
been aggravated not only by the rapid increase in The indirect value of mangrove ecosystems lies
catch in the 1960s and 1970s, but also by serious in their role of sustaining other natural resources
such as fish, crustacea and shellfish, and they
Table 5. Average annual marine catch in the Southeast Asian
have been shown to support nearshore fish pro-
countries (World Resources Institute, 1992). duction (MacNae, 1974; Unar & Naamin, 1984).
Country Average annual marine catch

Table 6. Areal extent of mangroves in the Asean countries
1987-89 Percentage (Sources: Darsidi 1984, Aksornkoae 1986, Chan 1986, Corlett
(thousand change over 1986, Tech. Staff, Philippine National Mangrove Committee
metric tons) 1977-79 1986, Zamora 1987).
Brunei Darussalam 3 73 Country Area (km2 )

Cambodia 6.5 -34
Indonesia 1971 60 Brunei Darussalam 184
Malaysia 598.6 -10 Indonesia 42510
Myanmar 552.5 40 Malaysia 6288
Philippines 1478.1 25 Philippines 878
Singapore 14.7 -5 Singapore 5
Thailand 2629 35 Thailand 2873
Vietnam 620.8 51 TOTAL 52738
144
In Indonesia alone, the value of mangrove for- 150 years ago to an estimated 1% in 1980 (Chou
estry products both for export and domestic use, et ai., 1980).
amounted to an estimated US$26 million in 1978
while mangrove-linked fisheries amounted to Seagrasses
US$194 million (Salm & Halim, 1984). Chong
et al. (1990) showed that tropical mangroves were The valuable role of the region's seagrass ecosys-
more important as feeding grounds than as nurs- tem is already recognised. Seagrasses thrive in
ery grounds for juveniles of commercially impor- shallow waters and are known to be widespread
tant fish species. Their study also confirmed that and with a high biodiversity (Fortes, 1988). How-
mangroves were important nursery areas for ever, data on the extent of this ecosystem in the
commercially important prawn species. Man- region remain limited (Fortes, 1989).
grove forests are also important in protecting The major economic importance of seagrass
shorelines against erosion and modifying the ef- beds stems from their provision of critical nurs-
fects of typhoons on coastal areas. They trap ery grounds for many commercial species offishes
sediment washed down by rivers and restrict and shrimps (IUCNjUNEP, 1985b). The ecosys-
freshwater runoff from land so the salinity of the tem, with its high primary productivity, supports
coastal area remains stabilised. large invertebrate and fish stocks in surrounding
The region's mangroves support a high diver- areas. Genera of commercial importance which
sity of over 300 plant species and more than 1000 are dependent on seagrass beds for some stage of
marine invertebrate and vertebrate species, in ad- their life history include Penaeus, Lutjanus, Lethri-
dition to 177 bird and 36 mammal species asso- nus, and Siganus. Seagrasses also form the major
ciated with mangroves. Some of these, like the source of food for dugongs, green turtles and
proboscis monkey, Nasalis iarvatus, of Borneo are juvenile hawksbill turtles, all of which are of
of scientific interest. interest from the scientific or conservation point
While a better scientific understanding of the of view.
full potential of the mangrove ecosystem is re- The economic value of the fishery associated
quired to enable a proper valuation of the re- with a seagrass ecosystem in Tarut Bay, Saudi
source, non-sustainable exploitation and destruc- Arabia, has been estimated to be US$8 million.
tion are occurring at an accelerating pace in the If the seagrasses were consumed directly by green
region (Paw & Chua, 1991). Population growth turtles, the turtle yield would be US$46 million
has increased the pressure on mangroves for fuel (Basson et ai., 1977). Fortes (1989) provided a
and building materials. Mangrove related fisher- review of economic valuations of seagrass-
ies are being rapidly exploited. The ever increas- associated fisheries. These values although theo-
ing demand on mangroves for timber, chipboard retical, provide an indication of the importance of
and paper has led to large scale deforestation of the resource. The indirect value of the ecosystem
the resource. Policy makers have long regarded is usually ignored. These communities bind sedi-
mangroves as wasteland that encourages the ments, reduce turbidity, retard erosion and lower
breeding of mosquitoes and have cleared exten- levels of pollution.
sive areas for agriculture, aquaculture, as well as Little attention has been paid to the importance
residential and commercial development. In the of the seagrass ecosystem. A high percentage has
Philippines, the extent of the mangrove ecosystem been affected by industrial, agricultural and sew-
was reduced by about 65% from 418990 ha in age discharge, coastal reclamation and dredging,
1967 to 146139 ha in 1978 (NEPC, 1980). Be- and overfishing. Destruction of this ecosystem
tween 1969 and 1979, 700000 ha in Indonesia has been known to cause the collapse of the
was cleared for agriculture (Soegiarto, 1980). De- shrimp fishery in some parts of the world while
velopment in Singapore has reduced the extent of overfishing leads to changes in the ecosystem,
mangroves from 10 to 12% of total land area often resulting in population explosions of other
145
species such as sea urchins which eventually Rocky shore habitat is widespread in the region
destroy the natural balance (Stauffer, 1937; and range from limestone to volacanic rocks.
Rasmussen, 1977; Stoner, 1980). They support a high diversity of life. The upper
intertidal zone is dominated by barnacles, lim-
pets, nerites and oysters throughout the region.
Soft bottom habitats and rocky shores Corals are usually associated with the lower in-
tertidal and subtidal zones, where abalones and
Soft bottom habitats consist of two categories, spiny lobsters also occur. Commercial fish spe-
open and enclosed. The open habitats include cies include serranids, lutjanids, lethrinids and
inshore shallow seabed areas of sand and silt as breams. This habitat appears to have been less
well as the sandy and sand-mud beaches on ex- exploited.
posed shores. Throughout the region, sand and
clay predominate, influenced by river inputs.
There is a higher biodiversity and greater biom- Non-living marine resources
ass of flora and fauna on these shallow seabeds
than on deeper offshore seafloor. Molluscs and Hydrocarbons
echinoderms are usually associated with these
shallow soft bottom areas which serve as signif- Some areas of the region's seas contain rich
icant fishing grounds suitable for trawling. Stocks deposits of petroleum and related products
of a wide variety of fish species and shrimps are (Valencia, 1983) which contribute to the economy,
available. The sandy shores, themselves with lim- particularly of Brunei Darussalam, Malaysia,
ited flora and fauna, are important as nesting Indonesia and Thailand. A large source of petro-
grounds to turtles. The main threats to these habi- leum and gas reserves along the Sunda shelf re-
tats are beach sand mining and overfishing of mains untapped (Bilal, 1985). This resource in the
stocks. Development of beaches for recreational Asean region currently amounts to 3.5% of total
use which occurs throughout the region is so in- crude oil production and 2.5 % of natural gas
tensive in some areas that it places additional production worldwide. Offshore production of
stress on the environment. crude oil increased from 20% to 50% in 1980,
Enclosed soft bottom habitats include lagoons, and offshore exploration is expected to increase
mudflats, bays and estuaries. These are wide- and extend further into deeper waters. In the ex-
spread in the region, and are extremely productive traction of offshore crude, precautions to prevent
when they support mangrove and seagrass com- pollutive damage to the marine environment are
munities. Mudflats with their covering film of being taken but accidental spills and blowouts
micro algae and are highly productive. They are have occurred from time to time. Fisheries re-
usually associated with river mouths and the sources and ecosystems such as mangroves, coral
nutrient-rich organic sediments support the reefs and estuaries are particularly vulnerable to
growth of benthic organisms. The estuarine areas oil spills. Drilling operations themselves tend to
of the region support major fisheries, and mud- increase the sedimentation levels of the surround-
flats are used throughout the region for the culture ing waters.
ofthe blood cockle, Anadara granosa, which forms
a significant industry. Coastal lagoons are con-
verted for the aquaculture of milkfish (Chanos Minerals
chanos ), tilapia (Oreochromis spp. ) and catfish
(Puntius javanicus). These habitats are often sub- Submarine tin deposits up to depths of 65 m can
jected to heavy pollution because of the natural be exploited but cost considerations and present-
convergence of human populations to the vicin- state technology have so far confined mining ac-
ity of major rivers. tivities to nearshore waters. Silica sand and iron
146
sand deposits occur throughout the region and hydrocarbons, used as pesticides in agriculture
continue to be exploited. Mining of these miner- have also been detected in the coastal environ-
als in the sea involves dredging and disposal of ment but not at alarming levels.
tailings, and results in increased siltation of the Red tides, both toxic and non-toxic, caused by
waters. This in turn decreases marine primary blooms of dinoflagellates have been increasing in
productivity which in turn decreases fishery po- frequency and locality within the region (Maclean,
tential. Coral reefs are particularly susceptible to 1989). Although the causative factors have not
high siltation rates, and the decreased clarity of been positively identified, pollution from land-
water affects marine-related tourism and recre- based sources is strongly suspected. The occur-
ation. Sediment budget changes around the area rence of red tides has an impact on the mari-
being mined can alter coastal geomorphology, as culture industry as it usually results in fishkills of
well as affect power plant cooling. Coastal activi- immense proportions. Paralytic shellfish poison-
ties such as mariculture can also be seriously af- ing caused by ingestion of fish and shellfish dur-
fected. Pollution caused by such mining has oc- ing red tide blooms have resulted in fatalities
curred in various locations of the region, notably throughout the region and is of growing concern.
the Strait of Malacca and around Phuket. The discharge of raw sewage into coastal wa-
ters has raised the coliform count to beyond ac-
ceptable limits. Shellfish in these areas usually
Marine pollution have high coliform counts in their tissues. This
increases the risk of exposure to human patho-
Marine pollution levels are higher in the coastal gens and disease transmission.
waters than the open seas. This issue is of great
environmental concern, particularly with the con-
tinuing trend of increasing coastal population. Degradation of the marine environment
Fast growing coastal cities in the region, many
without adequate sewage treatment plants con- The factors contributing to the degradation of the
tribute to the degradation of coastal waters and marine environment have been identified to
shallow marine habitats. Open drainage canals be mostly human activities (Gomez et aI., 1990;
keep pouring raw effluents and industrial wastes GESAMP, 1990). Rapid development of the
directly into coastal waters. The majority of coastal zone and population growth have resulted
Southeast Asian rivers are so polluted with in the present situation. The uncontrolled exploi-
wastes, including raw sewage, that they are con- tation of living marine resources sometimes using
sidered biologically dead. Case studies of coastal destructive methods has led to the loss of habi-
pollution throughout the region have been high- tats and species. Many of these living resources
lighted by Hinrichsen (1990). are being removed at a rate far exceeding the
One of the problems concerning marine pollu- natural sustainable levels. With non-living re-
tion in the region is the lack of reliable long-term sources, the extraction process itself causes envi-
data (Gomez, 1988). Hungspreugs (1988) gives ronmental degradation. Nearshore mining activi-
an overview of the status of heavy metals and ties have often contributed to erosion of beaches
other non-oil pollutants in the region. Existing and loss of reef habitat due to sedimentation.
information on heavy metals in seawater remains Coastal waters have been sUbjected to pollution
insufficient. Heavy metal levels in benthic sedi- through the discharge of urban and industrial
ments appear to indicate high contamination of wastes. To support the physical growth of
some parameters within localised areas around coastal cities, reclamation of foreshore areas
high density coastal settlements. Hydrocarbon and changes in the coastal geomorphology by
contaminants have also been detected in benthic man-made construction are common. These
sediment of harbours and estuaries. Chlorinated changes have caused further impact on the ma-
147
rine environment by altering current patterns and off the recreational east coast of Singapore in
increasing the sediment load particularly when 1988 cost US$200000 in cleaning up operations
water circulation of the area becomes reduced. alone. Malaysia spent US$1 million to clean up
An economic assessment of logging versus fish- the Diego Silang oil spill in the Strait of Malacca.
eries and tourism in Palawan, Philippines These costs do not take into consideration envi-
(Hodgson & Dixon, 1988), showed that the loss ronmental damage in terms of loss of living re-
in benefits from fisheries and tourism due to sedi- sources and critical habitat, as well as loss of
mentation caused by logging, far outweighed the earnings by fishermen and loss of revenue from
benefits generated from logging. tourism (for recreational areas). Habitat restora-
Southeast Asian seas contain some of the most tion efforts are also costly. The Singapore River
intensively used shipping routes. Marine trans- and Kallang Basin were cleaned and restored to
port plays an important role in the region. A large a non-polluted condition in a 10-year US$150
volume of oil is transported through the region million programme. At Pattaya, Thailand, the un-
each day. Bilal (1985) estimated that 3.8 million restrained growth of beach facilities has resulted
barrels per day leaves the region while another in severe deterioration of coastal water quality
3 million barrels pass through the Strait of which now requires millions of dollars for resto-
Malacca en route to Japan. Such large volumes ration.
being transported daily poses a high risk of acci- It is obvious that restoration of degraded habi-
dental spills. The 1975 grounding of the Showa tats is expensive in terms of direct costs. When
Maru in Singapore Strait resulted in the largest oil seen as a total of other related or implied costs,
spill incident in the region. Mangroves in the sur- it appears appropriate that protective measures
rounding Indonesian islands which were seriously or policies on sustainable development be given
affected showed no sign of recovery or regenera- serious consideration, since it makes more eco-
tion 3 years after the spill (Soegiarto & Polunin, nomic sense in the long term.
1981). Oil discharge also comes from normal
shipping operations such as loading, bunkering
and deballasting. Adding to the problem is the Environmental protection measures and policies
spillage from small craft which discharge oily bilge
waters. The transport of hazardous substances is Many Southeast Asian countries have become
another risk factor that needs to be considered. aware of the environmental cost of marine pollu-
Accidents of this nature at sea however have so tion and habitat degradation and are beginning to
far been uncommon. Gomez et al. (1990) reported focus attention on the management of coastal and
three incidents in the Philippines. In 1977, 350 marine development, as well as on the restoration
tonnes of sulphuric acid were lost at sea and in of degraded habitats and denuded resources. Ex-
two separate incidents in 1978, 500 tonnes of amples demonstrating this trend are the 1989
caustic soda spilled from a sinking barge and Langkawi Declaration on the Environment issued
20000 bags of fertilizer were lost from a vessel hit by the Commonwealth heads of government, the
by a typhoon. 1990 Baguio Resolution on Coastal Resources
Management issued by the Policy Conference on
Managing ASEAN's Coastal Resources for Sus-
Environmental cost tainable Development, and the 1991 Singapore
Resolution on Waste Management issued by the
Environmental degradation has led to loss of Conference on Waste Management in the Coastal
living resources and critical habitats. Although Areas of the ASEAN Region.
difficult to quantify it is evident that the cost can In order to conserve genetic resources and
be enormous. Some indication is provided by the biodiversity, governments are beginning to estab-
cost of cleaning up oil spills. A 50-tonne oil spill lish more nature reserves and protected areas,
148
many of which include the marine environment. on 'Coastal Resources Management' examined
Most of these however, still have a long way to selected coastal sites within the Asean countries
go in terms of effective management and enforce- in order to prepare coastal zone management
ment. Some management plans fail especially plans for them which would serve as models for
when they do not take into consideration the the region. The Asean-Australia project on
needs of traditional users. Others fail because of 'Coastal Living Resources' provided valuable
strong pressure from the commercial sector or data on the state and development of coral reefs,
weak political will. Many of the countries are mangroves and soft-bottom communities, and is
'caught in a trap of debt and development' now examining the inter-relationships between
(Hinrichsen, 1990). More resources have to be them. The Asean-Australia 'Tides and Tidal
exploited to pay interest on growing debts. In Phenomena' project also provided information on
1988, 28% of Philippine's export income went oceanic current patterns and tidal levels and is
towards servicing debts. Many of the countries presently continuing as the 'Regional Ocean
need to balance the growing needs of an increas- Dynamics' project. An Asean-Canada project is
ing population against declining resources. World investigating the establishment of environmental
trade is forcing developing nations to export more criteria for the development and management of
of their natural resources in order to meet na- living marine resources and human health. The
tional development goals. Planners and econo- East Asian Seas Action Plan initiated by the
mists have to face the challenging task of effect- United Nations Environment Programme was
ing sustainable development for long term gain. adopted in 1981 by the governments of Indo-
Some of these measures appear to have worked nesia, Malaysia, Philippines, Singapore and
in small villages where community-based conser- Thailand. The plan provided a means by which
vation measures have been implemented to main- the countries may deal with common marine re-
tain living marine resources at sustainable levels. source problems. A more recent issue fast gain-
At Apo island in the Philippines, the local fishing ing interest is global climate change and the
community was shown that the destructive fish- associated sea-level rise. The implications on the
ing methods which they were using were destroy- marine and coastal environment in the region is
ing the reefs and causing fish stocks to decline. being examined by a task team within the East
They have now implemented certain conservation Asian Seas Action Plan.
measures and are enforcing them as a commu- The information being obtained from the re-
nity. This includes the assertion of their rights sults of these regional projects is useful in the
over the reefs of the island and the prevention of planning of long-term management strategies to
fishing by people of other communities. slow down the further degradation of the marine
More attention is being given to habitat en- environment and irreversible decline of the re-
hancement as a means of increasing productivity sources. An integrated approach will be impor-
and biodiversity. Artificial reefs have been estab- tant as this open-access resource is utilized by
lished in all the Asean countries, some at com- various sectors of the community. Reducing the
munity levels while others at national levels. Re- level of conflict between these sectors will help to
sults appear to be encouraging but more studies achieve the goal of sustainable development.
are required to determine their effectiveness in
contributing to biomass increase, instead of sim- References
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A review of otters (Carnivora: Mustelidae: Lutrinae) in Malaysia and

Singapore
N. Sivasothi 1 & Burhanuddin Hj. Md. Nor2

1 Department of Zoology, National University of Singapore, Kent Ridge, Singapore 0511;
2 Department of Wildlife and National Parks, Km 10, Jln. Cheras, Kuala Lumpur 50664, Malaysia
Key words: otters, Lutrinae, Peninsular Malaysia, East Malaysia, Singapore, review
Abstract
Four species of otters have been recorded from Malaysia and Singapore in the past: Lutra lutra (Common
or Eurasian Otter), Lutra sumatrana (Hairy-nosed Otter), Lutrogale perspicillata (Smooth Otter) and
Amblonyx cinereus (Oriental Small-clawed Otter). All four are listed in the Threatened Species Categories
of the IUCN Red List of Threatened Animals: L. lutra is designated 'Vulnerable' and the status of the
other three Asian species are 'Insufficiently Known' due to lack of information. From a review of the
available literature and collation of museum records from Malaysia and Singapore, the past status of
the four species are examined. Presently, only L. perspicillata and A. cinereus are common in Peninsu-
lar Malaysia where they are widely distributed, but both are rare in Singapore. L. sumatrana is rare but
apparently still found in East Malaysia. It is possible that L. lutra did not range in Malaysia this cen-
tury. Information about the habitat types, group size, range, diet and behaviour for all the species is poorly
known. The available information allows a discussion of the Smooth and Small-clawed Otters only.
This dearth of knowledge has hampered maintenance and breeding efforts of zoos and conservation
activity.
Introduction use as an indicator species for a healthy aquatic

environment. The last is a characteristic of this
At the inaugural session of the First International top predator which is sensitive to poor water
Asian Otter Symposium held at Bangalore, India quality, toxicity in aquatic food chains and dis-
in October 1988, a strong plea was made by turbance of terrestrial habitats adjacent to water-
Dr M. K. Ranjitsinh, Joint Secretary (Wildlife) ways. Thus the disappearance ofthe otter is often
and Director, Wildlife Preservation, Government associated with the degradation of wetland habi-
of India, ' ... to make the otter a symbol of the tats.
wetlands'. The rationale for this, as outlined in In Southeast Asia, little research has focused
the International Union for Conservation of Na- on the otter despite the presence of four species.
ture and Natural Resources (IUCN) Action Plan The authors, who have begun basic research on
for the conservation of otters (Foster-Turley et al., otters recently, surmise the current state of
1990), is three-fold: their worldwide distribution, knowledge about the otters of Malaysia and Sin-
widespread public appeal and in particular, their gapore and highlight gaps in knowledge.
152
Taxonomy taxa. He also pointed out (Van Zyll deJong, 1987:

2537) that the specific epithet of cinereus should
It is perhaps surprising to note that in an animal be used instead of cinerea in accordance with the
as well-known as the otter, the taxonomy has yet gender of the genus.
to be resolved satisfactorily. Many synonyms The species perspicillata Geoffroy, 1826, is
exist in literature and Harris (1968: 290-304) has often attributed to the genus Lutra Brisson, 1762
compiled a useful list of these. Two authors have instead of Lutrogale Gray, 1865. The latter is still
attempted a revision of the Lutrinae since Pohle not recognised at the generic level by many of the
(1919). Davis (1978) did not use the usual char- more recent classification of mammals including
acters such as cranial morphology normally used Ellerman & Morrison-Scott (1951), Honacki
in mammal taxonomy. Although he introduced et al. (1982), Corbet & Hill (1991) and Nowak
the use of vocalizations, baculum shape and (1991). Opinions differ, for Simpson's (1945)
prominence of prepuce in his classification, the classification, and authors such as Van Bree
authors agree with Van Zyll de Jong (1987: 2536; (1968), Willemsen (1980, 1986) and Van Zyll de
1991: 81) that the use of these characters were Jong (1987) use Lutrogale as a full genus.
unsatisfactory. Furthermore, Davis (1978) re- Willemsen's (1980) paper highlights the consid-
duced the number of extant species to nine, dis- erable differences that exist between the cranial
missing, amongst others, L. sumatrana, without and post-cranial anatomy of Lutrogale and Lutra.
examining the type specimens, nor giving reasons Pocock (1921: 543) considered the differences in
for disregarding many of the characters previ- skull structure to be of generic value. The differ-
ously used in defining the species. Unfortunately, ences in skull and external characters (coat, tail
these considerations have been overlooked by and rhinarium) were discussed in detail later (Po-
some authors (e.g. Chanin, 1985) who adopted cock, 1941: 292,293). In addition, there seem to
Davis' classification. A more exhaustive revision be aspects of behaviour which separate the social
was carried out by Van Zyll de Jong (1987) who Lutrogale from the other more solitary Lutra spe-
constructed a phylogenetic relationship within the cies (see Wayre, 1974: 37; Duplaix-Hall, 1975:
family (using 861 skulls and 65 skins from vari- 315,324-326).
ous museums) based on morphological data but The authors thus feel that the weight of the
considers it tentative prior to consideration of evidence favours the use of the genus Amblonyx
other character sets. for the species cine reus, and Lutrogale for perspi-
The current taxonomic problems mainly con- ciliata and use these names as such.
cern the generic status. Amblonyx Rafinesque, There is intraspecific variation of L. lutra
1832 is considered a subgenus of Aonyx Lesson, (Linn., 1758) in Asia (Harris, 1968: 136). The
1827, to which the species cinereus Illiger, 1815 Asian otters are usually smaller, with lighter co-
belongs, by many mammal listings (Ellerman & loured fur (particularly at the throat) and shorter
Morrison-Scott, 1951; Honacki et al., 1982; Cor- hair. Of the subspecies that ranges in Thailand
bet & Hill, 1991; Nowak, 1991). However some and Sumatra, Pocock (1941: 287) states that ' .. .it
authors (e.g. Simpson, 1945; Willemsen, 1980; is, indeed, the smallest known race of Lutra lutra,
Medway, 1983; Van Zyll de Jong, 1987), use Am- apart perhaps from aurobrunnea ... .'. Most authors
blonyx as a full genus. Pocock (1921: 541, 543) accept this form as the subspecies, L. I. barang
considered Amblonyx sufficiently different from F. Cuvier, 1823. There are even suspicions that
Aonyx based on the webbing of the fore feet and the Indo-Malayan form of the Eurasian otter is a
the facial vibrissae. When Van Zyll de J ong (1987) distinct species (see section on distribution of
subjected quantitative and qualitative characters Lutra lutra). Unfortunately, the diagnoses of
of extant species of otters to morphometric and lutra/barang and Lutra sumatrana (Gray, 1865)
cladistic analyses respectively, the results indi- are far from complete such that the possibility
cated that Amblonyx and Aonyx are divergent that they belong to the same species cannot be
153
excluded (P. J. H. van Bree, in litt.). Until further whose feet are incompletely webbed and bear ru-
clarification of the taxonomy, the authors use the dimentary claws which do not project beyond the
names Lutra lutra and Lutra sumatrana. toes. Wayre (1976: 118, 120) however, noticed
well-developed, curved claws on the paws of a
cub. Claws of the forepaws withered and dropped
Identification of otters in Malaysia and Singapore off in about five weeks. Pocock (1941: 295) ob-
served the same in feet of young specimens, sug-
Four species of otters have been recorded from gesting that this occurred when the cubs began to
this region (Figs 1 & 2). The smallest of these is hunt independently. Both authors observed that
the Asian Small-clawed Otter (Amblonyx cine reus ) claws of the forepaws dropped offfirst. The three
Fig. 1. Otters reported from Malaysia and Singapore. Schematic drawings of animal (side view) and rhinarium (front view).
a. Lutrogale perspicillata (Smooth Otter); b. Lutra sumatrana (Hairy-nosed Otter); c. Lutra lutra (Common Otter); d. Amblonyx
cinereus (Small-clawed Otter). Rhinarium drawings after CITES, 1983.
154
directly, such as interviews, should be treated with

caution.
Distribution
The assessment of the distribution of the four

otter species is based totally on an examination of
Fig. 2. Sketches of otter skulls (side view). a. Lutrogale per- literature and specimens from relevant museums
spicillata (Smooth Otter); b. Lutra sumatrana (Hairy-nosed in the region, namely the Zoological Reference
Otter); c. Lutra lutra (Common Otter); d. Amblonyx cinereus
(Small-clawed Otter). After Payne et al., 1985 and Lekagu1 &
Collection in the Department of Zoology, Na-
McNeely, 1988. tional University of Singapore, the Sarawak Mu-
seum at Kuching, Sarawak, and Muzium Negara
in Kuala Lumpur (see Lim, 1985). Material in the
larger species, however, possess feet with well- British Museum (Natural History), London, was
developed webbing and claws. The short fur of also considered. No specimens from the Malay
the Smooth Otter (L. perspicillata) gives its coat Peninsula were found amongst the collection of
a smooth, velvety appearance. It has the most the Bogor Museum (Museum Zoologicum
massive head of the four species and the terminal Bogoriense), Indonesia. The collated list oflitera-
half of its tail is flattened. Ansell (1947: 381) even ture and museum records is presented chrono-
suggested that the species be given the common logically in Appendix I. Prior to this century, the
name of 'flat-tailed otter'. The widely distributed records are not consistent and specific references
Common or Eurasian Otter (Lutra lutra) has to mammals of the Malay Peninsula besides Can-
denser, coarser fur which appears grizzled or tor (1846) are few and far between. There is also
frosty due to the paler tips of its guard hairs. The some confusion with the identification of species.
teeth are smaller and the muzzle is longer than Most of the early literature do not specifically
that of L. perspicillata. The Hairy-nosed Otter address the Lutrinae but are general accounts of
(L. sumatrana) is similar to the Eurasian Otter mammals. Thus few accounts before 1900 are
but is distinguished by its longer, flatter skull and included and the period considered covers this
a hair-covered rhinarium, which is bare in the century only. Different synonyms which were
other species. The colour of the upper lip, chin used in the course of the century had to be clari-
and forethroat is whitish or yellowish and strongly fied. While the record presented here may not be
contrasted with the rest of its coat. exhaustive, the material examined thus far is suf-
Concise, detailed descriptions exist in literature ficient for an assessment of the distribution of
(Pocock, 1941; Harrison, 1966; CITES, 1983; otter populations past and present. Duplication
Medway, 1983; Payne et a!., 1985; Lekagul & of records amongst the literature has been avoided
McNeely, 1988) but the characteristics outlined as far as possible, for some accounts of mamma-
above suffice to distinguish the four species. It lian fauna are based on the same material e.g.
must be noted, however, that although size is rea- Davis (1962) is not included for his material has
sonably useful in distinguishing the Small-clawed been accounted for elsewhere.
Otter from the other species within this region, it
is not always reliable. Identification of the larger
three species is difficult without a close examina- Amblonyx cine reus
tion of the nose, fur or skull. The Hairy-nosed
Otter in particular, is the ' ... most difficult to iden- This was a common species in Malaysia and Sin-
tify in the field' (Foster-Turley & Santiapillai, gapore as far as old records suggest. Both writ-
1990). Thus information of sightings obtained inten accounts and specimens attest to its presence.
155
Kloss (1909) and Robinson & Kloss (1909) con- bode well for this animal. Harrison's (1966: 223)
sidered it the most abundant otter in the Malay claim that they were ' ... abundant in the sea off
Peninsula and Singapore and this seems to hold Penang' is challenged by Wayre (1974: 26) who
true for the rest of the century for they are still the considered the statement to be ' ... clearly a case of
most common today. The same may be said of misidentification', as that location was and is well
East Malaysia as well. In Singapore, otters have known for Smooth Otters. It was apparently less
always been elusive and Harrison (1966) felt that common in Singapore during this period for Har-
during his time, they were visitors rather than a rison (1966) felt their status to be more of visitors
permanent fixture of the Singapore coast. Cur- rather than residents. Or perhaps Chasen (1924:
rently, it would seem to be only this species which 84) was correct in saying that the otters were ' ...
is sighted infrequently amongst the least disturbed adepts at concealment'. At any rate, there seems
areas of the island, present probably as visitors. no reason to doubt their presence during the ear-
There is insufficient evidence at present to suggest lier part of the century. Surveys conducted since
otherwise. An exception could perhaps be Pulau (Wayre, 1974; Osman & Shariff, 1988; Nor,
Tekong Besar, an island in the northeast of the 1990a) did not encounter this species. Misiden-
main island. In 1989, two adults with not less tification cannot be ruled out due to the superfi-
than three cubs (probably this species) were cial similarity in its appearance to the Smooth
sighted, raising the possibility of a breeding resi- Otter. Few people are familiar with the form of
dent population (Wang, pers. comm.). the Hairy-nosed Otter, and currently there is little
in the way of feral or captive populations, or mu-
seum specimens to examine. This is aggravated
Lutrogale perspiciliata by the fact that the species' more obvious feature,
the 'hairy nose', is apparently not reliable in
The record indicates that the Smooth Otter adults. Both Cantor (1846: 195) and Blanford
ranged throughout Malaysia during the early (1888: 187) concur that hairs of the nose become
1900's. The early record for Borneo is quite poor, partially worn off in adults. Anderson (1878: 205),
while in Singapore, a single specimen was col- commenting on the nose of an adult specimen of
lected in 1938, from Lazarus Island in the south. Cantor's in the Indian Museum, said that it is
This could have been a visitor. Currently, this ' ... not so thickly clothed as in the young'. Pocock
species can be found with ease in Peninsular Ma- (1941: 289) reports that while hairs may be rubbed
laysia and a recent sighting of an otter in Sin- off from dried museum skins, the follicles leave a
gapore was probably of this species. It is still finely-pitted rhinarium surface in sumatrana, as
regarded to be rare in East Malaysia. This may opposed to a coriaceous surface in other bare-
be an underestimate, for many parts of Borneo nosed oriental species. Wayre (1974: 37) sug-
remain poorly explored. gested that the Hairy-nosed Otter would be found
in torrential streams at elevations above 300 me-
tres in the Peninsula. Recently, however, a skull
Lutra sumatrana from a road kill in Seberang, Perak in 1991, was
identified by Bishop of the BMNH as
The Hairy-nosed Otter does not seem to have L. sumatrana (A. C. Sebastian, pers. comm.) and
been rare in Malaysia or Singapore during the three individuals observed in Ulu Lepar, Pahang
early half of the century. All the early works and were recorded as this species (Sebastian, in prep.).
lists of mammals in the region include this species The presence of the Hairy-nosed Otter in Borneo
but it is not known if these lists were based on has been recorded during the earlier half of this
fresh sightings. It was present throughout Borneo century. Recent records suggest that this otter still
and the record for Sarawak is particularly good. ranges in Borneo and is less rare there than in the
But the latter half of the century did not seem to Peninsula.
156
Lutra lutra 1924, 1925a, 1940; Tate, 1947; Harrison, 1964,

1966) did not include the Eurasian Otter. Kloss
The range in which the Eurasian Otter is said to (1909: 33) aptly points out: 'Although Lutra vul-
have occurred stretches from continental Asia garis, the common otter, has been included at
through the Malay Peninsula to Sumatra in the times in the Peninsular fauna, there is no satis-
south. However, only two papers mention the factory proof of its occurrence'. It has not been
existence of this otter in Malaysia and Singapore encountered during recent surveys either (Wayre,
this century, and both are dated 1900. Flower's 1974; Osman & Shariff, 1988; Nor, 1990a). Hence
(1900) record is unconfirmed for he refers to there is still 'no satisfactory proof of its
specimens labelled as Lutra vulgaris (which he did occurrence' in Malaysia or Singapore this cen-
not examine) in the Raffles Museum purportedly tury. As such, labelling the Common Otter as
from Singapore and Malacca. It is prudent to 'Maybe extinct' in Malaysia and Singapore
consider Chasen's (1925a: 87) statement that it (Foster-Turley & Santiapillai, 1990: Fig. 3) is per-
' ... must be remembered that very large numbers haps not accurate for it assumes a previous dis-
of mammals are imported into Singapore each tribution here. Concluding that this species never
year for trade purposes. Individuals very fre- occurred here would be, however, overlooking the
quently escape and are just as often captured or fact that Southeast Asia was relatively little stud-
shot and brought to the Museum with the laconic ied and otters are in general, shy creatures which
statement that they were obtained in Singapore'. make encounters with man infrequent. The soli-
Thus the accuracy of the specimens' identity and tary nature and the more inaccessible habitats
locality cannot be guaranteed. There are no speci- (mountain streams and lakes) which this species
mens of the Eurasian Otter in the Zoological Ref- supposedly occupies (see Ewer, 1973: 266; Lim,
erence Collection (formerly the Raffles Museum). 1991) adds to the difficulty of its accurate iden-
The only set of specimens from this collection tification.
sent to the British Museum (Natural History) for The single record for Malaysia in Pulau
which there exists a record is the Robinson's Col- Langkawi, its northernmost island, is also the
lection (see Hill, 1960: 37), but this contains no southernmost record of Lutra lutra for this part of
Eurasian Otters. No other records seem available continental Asia as well. This suggests that
in the British Museum (J. E. Hill, pers. comm. to L. lutra in Sumatra has been geographically iso-
C. M. Yang). Recently, Daphne M. Hills of the lated from the rest of its range. Three possibilities
Mammal Section there compiled a list of speci- remain: that this species did occur in the penin-
mens of the four species collected from this re- sular but was never detected, or that sumatrana/
gion. Despite looking for specimens of Lutra lutra lutra/barang are the same, or that the Indonesian
from the Malay Peninsula in particular, none were L. I. barang is a distinct species from the Palae-
found (D. M. Hills, in litt). arctic L. lutra. The last possibility was raised by
The second reference is Miller (1900), who P. J. H. van Bree (in litt.) who pointed out the
records an adult female caught on Pulau case of Mustela sibirica/M. lutreolina (see Van
Langkawi by Abbott on 10th December, 1899. It Bree & Boeadi, 1978).
is to this record which Medway (1969, 1978, 1983) It is emphasized that otters are shy and hence
refers when he includes the Eurasian Otter as a a precise appreciation of the distribution of any of
member of the mammalian fauna of this region. the otter species can not be known to any great
Unfortunately, many e.g. Inskipp & Barzdo degree without thorough field work specifically
(1988: 66) have assumed this to be Medway's aimed at surveying these animals. This is reflected
record and thus recent (i.e. 1969, 1978 or 1983 by F. N. Chasen's (1925a) comment that 'All the
instead of 1900). Significantly, several lists and local inhabitants seem agreed that Otters are nu-
general accounts of the mammalian fauna in the merous thereabouts but we did not meet with any
region (Flower, 1900; Kloss, 1908, 1909; Chasen, on the present occasion'.
157
Ecology and biology of Smooth and Small-clawed viable otter populations. These have been identi-
otters fied as: an ample food supply, a freshwater source,
unpolluted water, protection from man, adequate
This section summarises the information avail- cover and accessible dry substrate (Foster-Turley,
able from Malaysia and Singapore. A reasonable 1989). The last two factors are important to the
amount of literature is available only for A. ci- animal for maintaining a healthy coat condition
nereus and L. perspicillata. The few comments that which is known to be a vital consideration in the
are available about the Hairy-nosed Otter are in- maintenance of captive otters (Duplaix-Hall,
cluded but are inconsequential. In East Malaysia 1972, 1975). In the wild, covered dens and dry
where it still ranges, no ecological studies have resting sites are found in a variety of locations.
been attempted (Davies & Payne, 1982). The Dens are found in earth tunnels below bamboo
Eurasian Otter has not been studied in this part thickets, amongst cavities in exposed Ficus tree
of the world. Hence only the Smooth and Small- roots, openings in boulder piles, amongst Acros-
clawed Otters are considered here. For an ac- tichum ferns in mangroves, and in dense shrubs
count of adaptations to an aquatic existence by adjacent to ricefields in which the Small-clawed
the Lutrinae, see Estes (1989). Otter shelters during dry seasons. The Smooth
Otter is a powerful burrower and able to dig into
banks to make dens whereas the Small-clawed
Habitat types has a more limited digging ability. The otters have
been observed resting on bushy scrubs in rice-
The Small-clawed and Smooth Otters adapt well fields, bare sand or non-woody vegetation on
to a variety of habitats. In Malaysia these otters shoulders of river banks, mud banks in man-
are found in freshwater and peat swamp forests, groves, and on sandy banks or beaches of lakes
ricefields, and other freshwater (lakes, streams, and rivers (Wayre, 1974; Shariff, 1984, 1985;
reservoirs, canals, flooded fields), brackish (man- Foster-Turley, 1989; Nor, 1990a).
grove) and marine (ocean shores) habitats
(Osman & Shariff, 1988; Nor, 1989). Nor (1990a)
noted some degree of habitat partitioning: the Group size, and range
Smooth Otter is found amongst the large lakes
and rivers whereas the Small-clawed seemed con- The two species of otters are found in groups in
fined to smaller rivers and streams. These obser- the wild. The Small-clawed Otter is said to ' ... live
vations concur with those ofWayre's (1974). Both in loose family groups of about a dozen
species were, however, equally common in areas individuals' (Timmis, 1971), while a typical group
of mangrove and ricefields - habitats in which the of Smooth Otters observed by Wayre (1974) con-
highest populations of otters could be found. In sisted of two parents with up to six young. Both
particular, ricefields appear to be one of the most species have been recorded in larger numbers in
suitable habitats in supporting viable populations India (Pocock, 1941: 302).
of otters. It is suggested that the Smooth Otter Foster-Turley (1989) elaborates another char-
seems more tolerant of the presence of man acteristic that the two species share with other
(Shariff, 1984), and that the Small-clawed Otter otters, namely, the larger populations and more
is more adept at exploiting the variable fish popu- gregarious nature of otters in coastal areas or
lations in the ricefields which peak prior to plant- marshes than in rivers. Wayre (1974) suggested
ing and harvesting (Shariff, 1985). In Singapore, that a group of Smooth Otters requires 7 to 12
recent sightings were all in or near mangroves kilometres of river for their territory and an even
(Yang eta!., 1990). longer stretch of coastline ifliving along the shore.
Despite the variety of habitats which otters oc- Furuyu (1977: 41) observed 'presumably' Small-
cupy, a few prerequisites appear necessary for clawed Otters often in groups of four to eight at
158
sites in Padas Bay, Sabah. Shariff(1984) detected been reported in other parts of Asia of the Small-
the presence of large groups but low incidence of clawed, Smooth and Common Otters (Blanford,
solitary sightings (6) in coastal mangrove, and at 1888: 184; Pocock, 1941: 302,313).
a freshwater system, reported a high incidence of No comprehensive work had been attempted
solitary sightings (65) but absence oflarge groups on the diet of these animals in the wild until
(> = 4). Foster-Turley's (1989) observations of Foster-Turley initiated her investigation on the
the species in Malaysia and Thailand are in agree- Small-clawed Otter. Her results will be published
ment. soon (in litt.). The available literature consists of
Melquist & Homocker (1983) correlated the rather brief analysis of scats (or spraints). Only
home range of Lutra canadensis inversely with hard parts from the diet remain and identification
food abundance. This could explain the greater of skeleton fragments or fish scales to species
numbers of otters on the coast which enjoy a level has not been attempted. This will remain one
relatively constant food supply unlike riverine of the major problems in diet studies. However,
systems which suffer seasonal changes in water there is sufficient information in the available lit-
level and possibly other factors affecting prey erature to highlight two points for consideration.
availability. The first observation is that availability of prey
affects diet. Ewer (1973) considers a successful
carnivore as one which practises flexibility in prey
Feeding and diet utilisation. Foster-Turley (1989) mentions such
diet variation in three populations of Smooth Ot-
Medway was reported by Wayre (1976: 167) to ters. One mangrove population fed on only man-
believe that L. sumatrana inhabits torrent streams grove fish while another fed on both fish and ma-
and feeds on small fish, frogs and crabs. A rine crabs. A third population in nearby
collector's note accompanying specimens of this freshwater ricefields with adjoining canals fed on
species from Sarawak said, 'Food: crabs and freshwater fish and rats. Likewise, Shariff (1985)
small fish'. Davis (1958: 138). Cantor (1846)notes found only fish scales in scats collected from rice-
that 'Its food is not confined to fishes and crus- fields (Kampung Pandak Putih, Perak) while
tacea; birds and insects are equally relished'. scats collected from two coastal islands
Lutrogale perspicillata subscribes to the feeding (Langkawi and Anak Gua Cerita) also contained
manner of members of the genus Lutra which crab and crayfish. Scats of the same species from
tend to locate their prey visually, catching it in the east coast of Malaysia examined by Wayre
their mouths. Shariff (1984) observed them for- (1974) consisted entirely of crabs.
aging among fallen tree trunks, rapids, fishing nets The second observation about diet concerns
and other obstacles in the water. They held their the Small-clawed Otter. Lekagul & McNeely
heads above the water surface, moving slowly (1988) state that they eat less fish and rely more
with sudden glides into water to catch fish. A. ci- on molluscs and crabs. But their diet too still
nereus has developed considerable digital move- depends on prey abundance. The main prey item
ment and very sensitive forepaws which it uses as in scats examined by Nor (1989) of both species
hands to feel for prey and conduct other manipu- from ricefields of Tanjung Piandang, Perak, was
lative behaviour (Timmis, 1971; Duplaix-Hall, fish. However, crustacean remains were found
1972; Wayre, 1976). It will even search under only among scats of A. cinereus but not L. perspi-
boulders and in crannies and has less trouble in cillata. Foster-Turley who also collected scats
murkier waters than L. perspicillata (Medway, from four sites at Tanjung Piandang observed
1983; Nowak, 1991). Timmis (1971) observed a that while crabs were infrequently found in scats
group of ten digging up shell fish which they of Smooth Otters, they formed the major com-
opened by laying them out in the sun. There are ponent of the Small-clawed Otter's diet (Foster-
no accounts of co-operative fishing, which have Turley, in litt.). Examination of her samples indi-
159
cate that the crabs were mainly Sesarmines, which (1984) observed otters at Kuala Gula in Perak
seems true of the species at Kuala Selangor N a- repeatedly belly-sliding down banks for long pe-
ture Park (first author, pers. obsv.) as well. Lim riods of time and classified such behaviour as
Boo Liat who examined stomach contents offour play. Similarly in Singapore mangroves, Hiscock
individuals (A. cinereus) from Bukit Lagong and (1990) sighted an otter repeatedly wriggling on all
Ulu Gombak forest reserves, Selangor, found fours over a slight tidal incline to gain momentum
mainly crab particles (in litt.). The other material before releasing its forelimbs to belly slide. He is
included scales of fish and parts of giant scorpi- of the opinion that 'It appeared to be doing this
ons (Heterometrus longimanus) and millipedes for amusement rather than a means of travel.' It
(Julidea sp.). Many accounts of the Small-clawed was still engaged in this activity when the observ-
Otter also comment on its reliance on other ers left the scene. The purpose of such activity, if
aquatic organisms such as crabs and molluscs it is not play, has yet to be determined.
rather than fish (Medway, 1983; Lim, 1990; Little is mentioned about calls. Cantor (1846)
Nowak, 1991). Such variation in diet is of much writes of L. sumatrana, 'Its voice is a short shrill
importance to captive breeders in their efforts to whistling, not unlike the sound of the cricket, but
maintain healthy popUlations of these animals. stronger'. Timmis (1971: 109) writes ofA. cinereus
A possible reason for the importance of rice- that apart from basic instinctive calls of alarm,
fields as a habitat to otters in Malaysia could be greeting and mating, up to 12 calls are recognised
the availability of food. Both vertebrate (fish and and Medway (1983: 87) reports a 'variety of yelps
rats) and invertebrate (molluscs, crustaceans and and whimpers and, when disturbed, high pitched
insects) sources are abundant in this habitat (Nor, ullulating screams'. It is to be expected of more
1990a). The synchronous incidence of A. cinereus social species that an extensive vocal range exists
in ricefields with fish populations may be associ- (Ewer, 1973: 255). The second author has noted
ated with Trichogaster pectoralis, the snakeskin that distress call of A. cinereus served to rally the
gouramy (sepat siam), a food resource which help of other otters: on one occasion when he was
makes this habitat attractive. This fish occurs in carrying a captured otter in a sack, it gave out
vegetated open areas and is a prolific breeder. It distress calls. Soon, a pack of otters appeared
is abundant during rice growing season, and de- and an individual amongst them lunged for the
clines in density at the end of growing season. sack, in an apparent attempt to rip it open. Play-
Other fish found in the ricefields which are likely back of such alarm calls also attracted the ani-
prey species include the common snakehead or mals.
aruan (Channa striata), keli kayu (Clarias batra- Bipedalism is a gait not uncommon to otters.
chus) and the swamp eel or belut (Monopterus Duplaix-Hall (1972: 179; 1975: 326) mentions ot-
albus). ters walking biped ally when carrying cubs with
their forepaws back to their nest under their chins,
or when carrying bedding material to the nest.
Behaviour Many authors mention this posture adopted by
otters when they chanced upon the animal in the
No comprehensive studies exist on the behaviour wild. Notes by N. Annandale & H. C. Robinson
of otters in their natural environment; in fact, in Bonhote (1903: 11) relate of A. cinereus: ' ... a
there are few published articles of observations of party of four ... sat up on their hind legs and
Asian otters in the wild (F oster-Turley, in litt.). watched us, rubbing their faces with their paws',
The playful nature of the otter has been ob- as did Banks (1931: 61): 'When surprised, they sit
served in captivity (Leslie, 1971; Duplaix-Hall, upright on the hind legs and tail, the short fore-
1972) and in the wild. Although Chanin (1985: 23, paws hanging down free .. .'. Banks (1949: 40) also
24) is of the view that otters slide for practical reports bipedalism as a common trait of Lutra
reasons of locomotion rather than play, Shariff sumatrana, and Wayre (1974: 32) writes of a group
160
of six Smooth Otters which ' ... ran to and fro (1991) and Wayre (1976, 1989). Observations in
pausing to stand up on their hind legs' as his boat these literature point to significant social and be-
approached them too closely. This response is havioural differences between the Asian (A. ci-
reflected by Hiscock (1990) who reports: 'The nereus and L. perspicillata) and European
otter seemed to be aware of our presence for it (L. lutra) otters. The Asian species are social, di-
would often lift itself up on its front legs and look urnal and vocal with the male involved in raising
over at us'. Both A. cinereus and L. perspicillata the cubs. In the more solitary, nocturnal, quieter
may occasionally be observed on their hind legs Lutra !utra, however, the male is generally kept
at Zoo Negara, Kuala Lumpur (pers. obsv.). away from cubs by the female. During a brief
Duplaix-Hall (1972) warns of the problems this study of a family (1 male, 2 females, 1 cub) of
poses in zoos with such sunken enclosures. The A. cinereus in the Melaka Zoo by Nor (1990b),
curiosity which prompts the animals to go on the male spent more time on the maintenance of
their hindlimbs to peer over walls is often misin- the nest while the two females raised the cub.
terpreted as begging, thus encouraging feeding by Pellis (1984) gives an account of play-fighting in
the public who throw in food or dangerous for- this species. Studies on the Smooth Otter all give
eign objects - both of which prove harmful to the mention to its fossorial nature prior to breeding
animals. (Yadav, 1967; Badham, 1973; Desai, 1974;
Shariff (1984) observed that urination and de- Duplaix-Hall, 1975; Markowitz, 1982). Many
faecation occupied a significant part of the zoos in the world keep A. cinereus but L. perspi-
Smooth Otter's active period. Such conspicuous cillata is not commonly kept, e.g. 15 of the 31
scent post toilets of scats deposited on dry ground 'good zoos' in Britain display the former but none
are essential markers for other otters to locate display the latter (Ironmonger, 1992).
(Foster-Turley, 1989). These fixed defaecation There are two roles that a zoo can serve by its
spots are true of other otter species and may be maintenance of captive population of otters: pub-
observed of captive otter populations as well. This lic education and breeding. Public education is
habit provides reliable signs and is normally used aided by the fact that otters have a high exhibit
by researchers as an indication of the presence of value due to the water acrobatics of the Lutra spp.
otters (Mason & Macdonald, 1986: 47-53). and the fascinating land behaviour of the clawless
otters (Timmis, 1971; Duplaix-Hall, 1972). Its
role in environmental education as defined by the
Otters in captivity IUCN is discussed by Janf3en (1991). The role of
captive breeding in the conservation of species is
Not all wild animals are easily tamed but there is an important one (Flesness & Foose, 1990). The
sufficient evidence to indicate that otters, when breeding record of otters remains poor except for
caught young, are easily tamed. Ansell (1947: 382) the Otter Trust which successfully raised Eur-
comments that captive Smooth Otters in asian and Small-clawed Otter cubs (Chanin,
Rangoon Zoo quickly became accustomed to hu- 1985: 165; Wayre, 1989: 39, 47). The studies
mans. They have been kept as pets or employed mentioned so far tend to be based on observation
for their natural predatory skills by fishermen of sole collections in individual zoos. Recently,
(Cantor, 1846; Gudger, 1927; Maxwell, 1960: the American Association of Zoological Parks
158; Wayre, 1976: 103-169). L. sumatrana has and Aquariums (AAZPA) has taken the initiative
been kept as a pet as well (Cantor, 1846; Banks, to include the Asian Small-clawed Otter as part
1931: 60; Wayre, 1976: 167). of its Species Survival Plan (SSP) programme. It
A summary of otters in captivity is unneces- intends to manage viable captive populations of
sary, the reader being referred to the accounts by A. cinereus in order to assist conservation in the
Duplaix-Hall (1972, 1975), Foster-Turley (1990) wild (Foster-Turley & Engfer, 1988). Such coor-
(who also refers to Wright, in press), Reuther dinated work effectively increases the population
161
size of otters observed, thus allowing a significant certain instances, the otters are killed, despite the
contribution from zoos. The SSP identified renal prohibition by law. The extent of such incidents
calculi as a significant health problem amongst is not known and are difficult to estimate. How-
A. cinereus in North American zoos (Foster- ever, this is not considered the major threat to
Turley & Engfer, 1988; see also Nelson, 1983; populations. Otters are totally protected in Ma-
Calle & Robinson, 1985). This has also been laysia by the Wildlife Act of 1972 and in Sin-
noted at the Otter Trust (Wayre, 1989: 82). Other gapore, by the Wild Animals and Birds Act, 1985.
health problems amongst captive otters (Lan- The contribution of these laws to the eventual
caster, 1975: 65; Rogoschik & Brandes, 1991) survival of the otter species in this region is at
that might exist have to be highlighted in order to best, minimal, for their habitats are not protected.
initiate preventative measures for successful Pollution of the wetlands and the rapid conver-
maintenance by zoos. sion of their habitats to other land uses will re-
Locally, emphasis should be placed on docu- main the bone of contention in the discussion of
menting and publishing work based on captive the otter's survival in Malaysia and Singapore.
studies. Local zoos possess the advantage of Perhaps many feel that the current situation
natural climatic conditions which can only benefit does not justify concern. Mterall, the two species
any breeding plan they conduct. As of now, no currently present in the Peninsula are widespread.
major work on captive otters native to the Indo- However, with the increasing threat offered by
Malayan region originate there although the Asian industrialisation and development, conservation
Small-clawed Otter has been bred in the Sin- plans which include public education as well as
gapore (Calle & Robinson, 1985: 1149), Melaka the many aspects of research, have to begin well
(Nor, 1990b) and Kuala Lumpur (pers. obsv.) in advance. Complacency over the current situa-
zoos. tion could result in significant problems later.
The case of the European otter L. I. lutra is
worth considering. The reasons for the decline of
Conservation this species ' ... were the usual ones' (Lang, 1977).
River bank improvement, intensive use of the
The four species recorded from Malaysia and river, pollution and hunting all contributed to the
Singapore are all listed under the Threatened pressures that suppressed its popUlation. Chris-
Species Categories of the 1990 IV CN Red List of tian Schmidt of the Zoologischer Garten Zurich
Threatened Animals. Lutra lutra is listed as (1972) summarised the decline in Switzerland
'Vulnerable' while Lutrogale perspicillata, Lutra where it was formerly abundant: the otter was
sumatrana and Amblonyx cinereus are designated exterminated many years ago because it was
the category, 'Insufficiently Known', assigned to wrongly considered to be the main competitor of
taxa of which little is known. This highlights the fishermen. The Swiss government even paid a
appalling state of knowledge about the Asian ot- bounty for every otter killed. In 1938, it was still
ters which is lacking in as fundamental an aspect possible to record free-living otters in the centre
as distribution. of Berne. By 1952, the otter was protected but
Although otters have never constituted a food only 40-60 individuals were alive at that time. By
source nor been hunted for its pelt, it is often 1972, the European Otter had vanished in Swit-
construed to be a pest by fishermen. A problem zerland; the protective measures taken had been
created by otters in the ricefields is the damage too late. A decision was then made to build a
caused by their play. They also feed on the fish compound with the aim of keeping and breeding
which are drained into sump ponds after the har- otters on a large scale and under natural condi-
vest of ricefields. This fish constitute an additions. The authorities hoped to educate the pub-
tional income for the ricefield farmers who usually lic that the species is not a pest and eventually
do not want to invest in otter-proof fencing. In reintroduce surplus specimens into suitable wild
162
Swiss habitats. Its breeding record in captivity is presence of otters strikes an optimistic note in an
unfortunately poor (Lang, 1977) partly due to otherwise gloomy chapter of her disappearing
conditions in zoos (see Harris, 1968: 119). The fauna. It is possible that the establishment of a
current status of the animal in Switzerland (fide bird reserve in a patch of mangroves on the north
Macdonald & Mason, 1990) is that it is almost coast, the Sungei Buloh Nature Park, may even-
extirpated; restricted to one small lowland popu- tually attract the otter back.
lation only - the remnant of the reintroduction In his foreword to IUCN's Otters - An Action
programme of 1975 which failed to re-establish Plan for their Conservation (Foster-Turley et al.,
the otter (Macdonald & Duplaix, 1983). The 1990), George Rabb, Chairman of the IUCN
threats remain the same: habitat destruction and Species Survival Commission, identifies informa-
fragmentation of suitable habitats and water pol- tion gathering as a major part of the task ahead
lution (for a review of the latter, see Mason, 1989). in the conservation of otters. In the case of the
Switzerland is not the exception. The Action Plan Asian otters, details concerning distribution,
for European Otters warns that ' ...there are now habitat preference, diet and feeding patterns, in-
many western countries where the Eurasian otter terspecific interaction, field and captive aspects of
is very rare or extinct' (Mason & Macdonald, reproductive biology are still poorly if not known.
1990). This lack of knowledge has hampered the main-
There is serious cause for concern for Malay- tenance and breeding efforts of captive zoo popu-
sian otters too. Since the 1960's, apart from the lations as well as conservation programmes.
road kill mentioned earlier and the apparent Hence, research in the near future should be co-
sighting of three individuals in Ulu Lepar, the ordinated to answer the immediate questions of
Hairy-nosed Otter appears not to have been seen relevance to the long term survival of the otter in
in either Peninsular Malaysia or Singapore. Malaysia and Singapore.
Wayre's (1974) conclusion that it is isolated in the Certain lines of research pertinent to the even-
higher elevations of the Peninsular is perhaps the tual conservation of the otter species are being
first hint of a threatened species. Any animal that addressed. The second author is currently con-
favours forested streams would have been seri- ducting a detailed survey to determine the distri-
ously affected by changes in land use over the bution of otters in Peninsular Malaysia which is
past thirty years. If sizeable populations of this expected to be completed by 1992. It will also
species are not found soon, and its habitat not identify important areas and threats that these
protected, the Hairy-nosed Otter, which is en- species may face in the future. Other studies in-
demic to Southeast Asia, may well become ex- clude the feeding patterns of otters in ricefields of
tinct in Peninsular Malaysia. Scott & Poole's Tanjung Piandang, Perak and habitat use of ot-
(1989), Priority Species listing (covering mam- ters in Taman Negara. A project is about to begin
mals, birds, reptiles and amphibians) for Malay- on the spacing patterns and territorial behaviour
sia includes L. sumatrana amongst the nine spe- of the Small-clawed Otter with the radio-
cies identified as a special responsibility of the telemetry. The first author will be undertaking
country. That the list of site accounts does not basic research into the biology and ecology of the
include Peninsular Malaysia is itself an indication Smooth Otter which will include studies on habi-
of the threat to this species. A concerted effort tat, diet, behaviour and its interspecific interac-
will have to be made soon to ensure that the tions with the Small-clawed Otter.
continued survival of the Hairy-nosed Otter is The research to be undertaken is necessarily
given the concern that it deserves. basic for little is really known about this animal
Singapore's thriving development has reduced for which there is call to recognise as a symbol of
most of the mainland mangroves to small areas in 'the wetlands. It is ironical to compare the extent
the north. Despite its highly disturbed state, the of the literature available in Europe on their single
continued though infrequent indications of the species, Lutra lutra and the dearth of literature
163
that exists on the Southeast Asian otters, of which museum records and loan of specimens and
there are four species! The authors have generally Zainal A. Jamaludin of Muzium Negara. The au-
refrained from discussing the situation in other thors thank P. J. H. van Bree for advise on tax-
Southeast Asian countries for besides the Action onomy and loan of books, Mohd Khan Bin
Plan for Asian Otters (Foster-Turley & Santiapil- Momin Khan of Perhilitan, Kelvin K. P. Lim of
lai, 1990), not much information is available. the ZRC for the drawings and help with the search
Consider Lutra l. barang, for example. It is pos- for distribution records and Peter Ng for advise
sibly a distinct species and yet very little is known on taxonomy and for checking the draft.
about its ecology in Southeast Asia where it is
elusive and rare. The numerous undiscovered
habitats and the size of the Indonesian Archi- Appendix I
pelago only poses challenges and a lot of ques-
tions. It is possible that some work has been done Record of otter observations, 1900-1991
but remain inaccessible in publications such as
departmental journals. An important element that Relevant references made of otters in the literature is com-
piled. Prior to 1900, little information was available; however,
has thus to be considered in the conservation of the few that were found are included for the record. Synonyms
otters will be the establishment of a common da- that were used in the original literature appear in square
tabase of information. This is easily managed and brackets. The museum records include all material from the
yet will accelerate the pace of research while respective collections even if dated prior to this century. All
avoiding overlaps or loss of any useful work. Such specimens are skins, unless otherwise stated. ZRC: Zoologi-
cal Reference Collection (formerly the Raffles Museum) - the
a regional outlook of the conservation problems specimens in this collection were examined by P. J. H. van
which face the otter and its wetlands habitat must Bree; SM: Sarawak Museum - specimens were examined by
remain a priority for researchers in Southeast K. Lim of the ZRC, identity on the specimen labels, if differ-
Asia. ent from that diagnosed, is in square brackets; some speci-
There still remain populations of the four spe- mens have two catalogue numbers; MN: National Museum,
Malaysia - specimen examined by senior author; BMNH:
cies within this region. There is time as yet to British Museum (Natural History) - specimens examined and
avoid the fate that has befallen this animal over list compiled by Daphne M. Hills. Question marks (?) are
much of western Europe which has seen the sig- inserted where doubts exist about the specimens, spelling,
nificant decline of the Eurasian Otter together locality or species.
with the healthy wetlands it once used to inhabit.
Records of Amblonyx cinereus.
Published in literature:
Acknowledgements Cantor, 1846: [as Aonyx leptonyx, Gray] "Anjing Ayer' of the
Malays of the Peninsula.' 'This, as well as the two preceding
species (L. perspicillata & L. sumatrana), inhabits numer-
The first author would like to thank Dr Peter ously the banks of the Malayan rivers, and all are at times
K. L. Ng of the National University of Singapore used by the Malays in river fishing.'; Distr. - Malayan
for supervising this project. Many people encour- Peninsula, Singapore (p. 196).
aged the first author, offered useful advice and Lydekker, 1894: [as L. cinerea] Ranges through the Malay
responded with literature, for which he is very Peninsula and islands. (p. 96).
Ridley, 1895: Two species of otter have been met with in
grateful - Lim Boo Liat, Pat Foster-Turley, Singapore, viz., Lutra sumatrana and L. leptonyx; but they
P. J. H. van Bree, Gordon B. Corbet, Sheila seem to be rare, and little is known about them. The Malays
Macdonald, Padma de Silva, Charles Santiapil- often call them 'Anjing Ayer' (water dogs) (p. 94).
lai, Claus Reuther, Nicole Duplaix-Hall and Flower, 1900: [as Lutra cinerea Hliger.] Recorded from the
Wayne E. Melquist. Thanks to Daphne M. Hills Malay Peninsula; Malacca (1891); Tahan River, Pahang
(1894); Batu Pahat, lohore (1894); Singapore (1895, 1898)
of the British Museum (Natural History) for the by various authors. ' ... and there is a specimen caught in
compilation of their otters specimens, Charles Selangor in the Museum at Kuala Lumpur.'; Distr. -
Leh of the Sarawak Museum for help with the South-eastern Asia (p. 335).
164
Bonhote, 1903: [as Lutra cinerea IIliger.] 'Otters, probably of Banks, 1949: listed as a mammal of Borneo (p. 41).
more than one species, are common in Patani States, both Timmis, 1971: 'During February 1962, I was able to observe
high up in the rivers, in estuarine waters, and even in Patani from a small boat a group of ten Amblonyx at work on
Bay, the coast form attaining a very large size. The people a large sandbank 40 miles above Sibu on the Rajang
of the fishing village of Tanjong Budi, on Patani Bay, told river, Sarawak. ... On another occasion, several miles
us that the species was polygamous, and that the old dog inland between Balikpapan and Samarinda in Indonesian
otter always endeavoured to destroy the male pups, the Borneo, I heard the bird-like cries of Amblonyx without
usual number of the litter being four. It was very abundant actually seeing the animals; this was in quite heavy jungle
in this locality, and was often to be seen along the edge of with numerous small streams and swamp areas.' (p. 110).
the mangroves at low tide, or swimming in the waters of the Harrison, 1964: recorded from North Borneo (Sabah) (p. 22).
bay. Travelling down the Patani River, above Biserat, in Harrison, 1966: It appears in Singapore from time to time;
very rainy weather, we surprised a party offour on a shingle perhaps as a visitor, or perhaps as an escape - for it is a
bank, who sat up on their hind legs and watched us, rub- popular pet (p. 224); ' ... have been recorded from time to
bing their faces with their paws.' (p. 11). time, and one suspect that they visit here from the mainland.'
Kloss, 1908: [as Lutra cinerea, IIliger] included in the provi- (p. 6); present in Penang (p. 7); known from Malaya
sionallist of mammals of the peninsular region. (p.332).
Kloss, 1909: [as Lutra cinerea, IIliger] 'Throughout the Pen- Harrison, 1969: 5 animals collected by IMR between 1947-
insula and Singapore where it is the commonest of otters.' 1957 (p. 175).
(p. 33). Wayre, 1974: common in Rompin area and Kuala Trengganu.
Robinson & Kloss, 1909: [A onyx cinereus, IU.] 'Quite the Furuyu, 1977: observed families of otters at Padas Bay, Sabah.
most abundant of the otters of the Malay Peninsula and Medway, 1977: 'Records cover all mainland Borneo .. .'
found equally in salt, brackish and fresh water, and even in (p. 133).
mountain streams.' (p. 112). Davies & Payne, 1982: most information come from chance
Chasen, 1924: [as Lutra cinerea, Illiger.] listed in checklist of sightings and published works; A. cinerea is the most com-
mammals of Singapore (p. 83); 'Otters of any species are mon species, occuring wherever, there are streams or for-
either not common in Singapore or adepts at concealment, est cover. The otters are not regarded as sufficiently rare or
possibly the latter.' (p. 84). threatened by development to warrant special conservation
Chasen, 1925a: included in the true Singapore land fauna methods now.
(p.88). Shariff, 1984: Smooth Otter more abundant than Small-
Banks, 1931: occur in Sarawak (p. 60). clawed Otter in Kuala Gula, Perak, and Taman Negara,
Allen & Coolidge, 1940. Reporting The Asiatic Primate Ex- Pahang.
pedition, June-August, 1937, specimens in Museum of Shariff, 1985: Smooth Otter in Langkawi Island; Smooth
Comparative Zoology, Harvard: [as Micraonyx cinereus (1I- Otter and Small-clawed Otter in and around rice fields at
liger)] MCZ.36766, female, Sarawak, Borneo; MCZ.36627, Kampung Pandak Putih Baru, Perak.
female, Talibang near Tuaran, North Borneo (Sabah); Davison & Heang, 1987: Sighting of A. cinerea amongst ob-
MCZ.36726, female, Kalabakang River, Tawau (Sabah). servations made on mammals in Ulu Endau, 1985-1986.
(p. 150). (p.438).
Hill, 1960. Reporting on the Robinson collection (before Kemper, 1988: traces and tracks and sighting of A. cinerea in
1926): BMNH.55.1598, male, Taiping, Malaysia; primary lowland dipterocarp forest. (p. 14).
BMNH.SS.1599, male, Kuala Lumpur, Malaysia. (p. 37). Osman & Shariff, 1988: Results of preliminary survey of Pen-
Chasen & Kloss, 1931: [as Lutra cinerea, Ill.] 'Hose records insular Malaysia - only two species present, Small-clawed
the species as very rare in Borneo but it was very common Otter and Smooth Otter. Highest numbers of otters re-
at Bettotan [near Sandakan, Sabah] and a much larger ported from the state of Kedah.
series could have been obtained with ease.' (6 males, 3 fe- Nor, 1989: the Smooth Otter and Small-clawed Otter inhabit
males coli.) (p. 15). ricefields of Tanjung Piandang, Perak.
Chasen, 1940: Distr. - Malay Peninsula, Sumatra, Rhio- Foster-Turley & Santiapillai, 1990: Common in Peninsular
Lingga Arch., Borneo, Java (p. 93). and East Malaysia; believed extirpated in Singapore.
Davis, 1958. Reporting on Harrisson's collection from the Yang et aI., 1990: In Singapore, recent unconfirmed reports
Kelabit Plateau, northeastern Sarawak, between Sept. 1945 from Choa Chu Kang and Pulau Tekong and Central
and Dec. 1949: One female, two juveniles, Pa Main, 3500 Catchment Area (all amongst the least disturbed areas of
feet; one male, Bario, 3800 feet; one male, one female, Singapore); status = visitor straying from Peninsular
Bario. Collector's note. - 'In small shingle and sand stream Malaysia/Uncommon, occurs in small numbers in various
on fiats two miles from village.' 'In stream in jungle. Many localities and endangered.
others. Food: fish' (p. 134). Nor, 1990a: Only the Smooth Otter and the Small-clawed
Tate, 1947: ' ... typically of Java, but also found in Borneo, Otter found during survey of northern Malaysian states. No
Sumatra, and the Malay Peninsula' (p. 157). other species observed.
165
Museum records Lim, B. L., 1958: BMNH.1961.1266, female, with skull, Ulu
Gombak For. Res., Selangor.
Borneo:
Anon., 1905: ZRC.4.1156, male, Larut, Perak, Malaysia.
(?): Haviland, G., 1892: SM.49.22, male, Kuching, Sarawak.
Anon., 15.I1I.1920: ZRC.4.1157, male, Jeram, Selangor, Ma-
(?): Bartlett, E., 1892: SM.49.23, male, Kuching, Sarawak.
laysia.
(?): Saintbois, A., 5.VI.l900: SM.49.25, female, mounted
Neubromer, C. B. A., F. Ederma & J. Hausen, 25.x.1925:
skeleton, Kuching, Sarawak.
ZRC.4.1166-7, two juv. males, Pasir Panjang (?Bintan
Lewis & Hose: BMNH.1890.1.17.1, female, young, with skull,
Island in the Riau Archipelago or Singapore).
Sarawak.
Anon., 23.I1I.1892: SM.A49.21 (SM.0174/1), female, Kuch- Records of Lutrogale perspicillata.
ing, Sarawak.
Everett: BMNH.1893.3.4.7, female, with skull, Poeh River, Published in literature:
Poeh Mt., Sarawak. Burton, 1987: [examined libraries of Linn. Soc. Lond., Zoo\.
Scott, H. W. V., 1O.xU.1894: SM.49.13 (SM.0174/2) [un- Soc. Lond. and Mamm. Sect. of BMNH, also coli. of
iden.], juv. male, Kuching, Sarawak. latter] A single specimen was collected in Borneo last
Hose, 1894: BMNH.1899.12.9.29, female, with skull, Baram century (p. 143).
District, Sarawak. Cantor, 1846: [as L. nair F. Cuvier] "Anjing Ayer' of the
Anon., 7.lV.1902: SM.A49.26 (SM.0174/3), male, Kuching, Malays of the Peninsula.' (p. 195).
Sarawak. Lydekker, 1894: [as L. macrodus] ' ... also extends to Burma,
Barker, A. J. G., 1903: SM.A49.17 [Lutrasumatra],juv. male, the Malay Peninsula, and Sumatra.' (p. 96).
wet- preserved, Kuching, Sarawak. Flower, 1900: [as Lutra macrodous Gray] 'Recorded from the
Lalay (native?), 25.VI.l926: SM.A49.18 [Lutra sumatrana], Malay Peninsula by Cantor. A specimen caught in Selangor
juv. male, Tanjong Datu. is in the Museum at Kuala Lumpur.'; Distr. - India, Burma,
Liew, Moses, 6.VI.l956: SM.0174/4 [uniden.], juv. female, Malay Peninsula (p. 334).
died in captivity. Kloss, 1908: [as Lutra macrodous, Gray] included in the pro-
Liew, Moses, 6.VI.l956: SM.0174/5 [uniden.],juv. male, died visional list of mammals of the peninsular region.
in captivity. Kloss, 1909: [as Lutra macrodous, Gray] distr. - The Penin-
Seal, J., 29.VIII.1957: SM.0174/6, [Lutraperspicillata], male, sula (p. 33).
'bought ... off garbage collectors, 27.8.57, died 29.8'. Smith, 1919: [as Lutra tarayensis Hodgs.] 'Other specimens
Chua, Ah Bar, 23.lX.1963: SM.528 [uniden.], 'caught by his of this otter in the Raffles Museum are from Pulau Rumbia
son in the ditch. The boy was seriously bitten in the hand. in the Straits of Malacca off the Selangor Coast .. .' (p. 46).
He was paid $5 as compensation.', Satok Road, Kuching, Hill, 1960. Reporting on the Robinson collection (before
Sarawak. 1926): [as Lutrogale perspicillata, ( Geoffrey)]
Anon., 6.V.1978: MN.630, male, Santubong, Sarawak. BMNH.S5.1596, male, Port Weld, Larut, Perak;
Kloss, C. B. & F. N. Chasen, 2-25.VIII.1927: ZRC.4.1158- BMNH.55.1597, male, Kg. Padang, Tembeling, Pahang
65,5 males & 3 females, Bettotan, near Sandakan, Sabah. (p.37).
Referred to in Chasen & Kloss, 1931. Chasen, 1940: Distr. - Malay Peninsula, Sumatra (p. 93).
Anon., 14.VII.1962: ZRC.4.1534, female, ZRC.4.1535, male, Tate, 1947: ' ... occur in the Malay Peninsula' (p. 157).
Cocoa Research Stn., 700ft, Tawau, Sabah. Harrison, 1964: ' ... is not recorded from Borneo' (p. 182).
De Garcia & Harrison: BMNH.1971.3066, male, with skull, Harrison, 1966: 'It does not appear to be very common, and
Kalabakan, North Borneo (Sabah). is not recorded from Singapore.' (p. 224); known from Ma-
laya (p. 332).
Malaysia and Singapore Wayre, 1974: common wherever there was a suitable habitat
Anon.: BMNH.1839.11.11.10, sex uniden., purchased in Penang, Langkawi, Taman Negara, Kerteh and Rompin
(Stevens), with skull, Singapore Island. rivers, and Pangkhor Island.
Anon.: BMNH.1842.4.12.15, sex uniden., young, purchased Medway, 1977: Lists three records, near Sandakan; Darvel
(Warwick), skull only, Singapore.
Bay; Badang, S. Bahau, East Kalimantan (p. 133).
Cantor/Indian Mus.: BMNH.1879.11.21.568, male, with Davies & Payne, 1982: most information come from chance
skull, Malacca.
sightings and published works; large otters (L. sumatrana &
Int. Fisheries Ex.: BMNH.1883.11.20.1, female, with skull,
L. perspicillata) are seen along the Kinabatangan river. The
Singapore Straits Settlements.
otters are not regarded as sufficiently rare or threatened by
Robinson & Annandale: BMNH.1903.2.6.30, male, with
development to warrent special conservation methods now
skull, Ban Sai Kan, ?Malaya. (p. 141-146).
Butler & Robinson: BMNH.1955.1598, male with skull, Kuala
Shariff, 1984: Smooth Otter more abundant than Small-
Lumpur, Selangor.
clawed Otter in Kuala Gula, Perak, and Taman Negara,
Robinson, 1912: BMNH.1955.1599, male, with skull, Taiping
Pahang.
Gardens, Taiping, Perak.
Shariff, 1985: Smooth Otter in Langkawi Island; Smooth
166
Otter and Small-clawed Otter in and around rice fields at Lydekker, 1894: 'The hairy-nosed otter (L. sumatrana) is a
Kampung Pandak Putih Baru, Perak. very well-marked species from the Malayan region .. .'
Osman & Shariff, 1988: Results of preliminary survey of Pen- (p.96).
insular Malaysia - only two species present, Small-clawed Ridley, 1895: Two species of otter have been met with in
Otter and Smooth Otter. Highest numbers of otters re- Singapore, viz., Lutra sumatrana and L. leptonyx; but they
ported from the state of Kedah. seem to be rare, and little is known about them. The Malays
Nor, 1989: the Smooth Otter and Small-clawed Otter inhabit often call them 'Anjing Ayer' (water dogs) (p. 94).
ricefields of Tanjung Piandang, Perak. Flower, 1900: 'Recorded from the Malay Peninsula by Can-
Hiscock, R., 1990: large otter sighted, possibly Smooth Otter, tor, and from Singapore by Ridley. A specimen caught in
but could be Hairy-nosed Otter as well (see also Lim, 1990). Selangor is in the Museum at Kuala Lumpur.'; Distr. -
Foster-Turley & Santiapillai, 1990: Common throughout Pen- Malay Peninswa and Islands (p. 334).
insular Malaysia, more rare in East Malaysia; believed ex- Kloss, 1908: included in the provisional list of mammals of the
tirpated in Singapore. peninsular region.
Yang et al., 1990: In Singapore, Smooth Otter - status doubt- Kloss, 1909: Distr. - The Peninsula, Singapore and Langkawi
fw. (p. 13). Island. (p. 33).
Nor, 1990a: Only the Smooth Otter and the Small-clawed Chasen, 1924: listed in checklist of mammals of Singapore;
Otter found during survey of northern Malaysian states. No 'Otters of any species are either not common in Singapore
other species observed. or adepts at concealment, possibly the latter.' (p. 84).
Chasen, 1925a: included in the true Singapore land fauna
(p.88).
Museum records: Banks, 1931: occurs in Sarawak (p. 60).
Chasen, 1940: Distr. - Malay States, Sumatra, Banka, Borneo
Borneo (p.93).
Anon., 20.lX.1895: SM.A49.12 (SM.0174/8) [Lutra Davis, 1958. Reporting on Harrisson's collection between
sumatrana], female, Kuching, Sarawak. Sept. 1945 and Dec. 1949: One male, Pa Vmur, 3900 feet.
Malaysia and Singapore Collector's note. - 'In small rocky brook. The animal also
goes into jungle. Food: crabs and small fish.' (p. 133).
Cantor/East India Co.: BMNH.1860.5.4.62, female, with
Tate, 1947: 'The range includes the whole of the Malay Pen-
skull, Malacca, Malay Penin.
insula ... It also occurs on Sumatra ... and on Borneo'
Cantor/East India Co.: BMNH.1860.5.4.63, female, with
(p. 157).
skull, Province of Wellesley (Malacca).
Banks, 1949: listed as a mammal of Borneo (p. 40).
Cantor/Indian Mus.: BMNH.1879.11.21.567, female, Pro-
Harrison, 1969: 5 animals collected by IMR between 1947-
vince of Wellesley (Malacca).
1957 (p. 175); family in river seen in an area of disturbed
Linnaea & Frankfurt: BMNH.1884.3.14.1, sex uniden., Sa-
langa Island, Malacca. rainforest at Sg. Buloh, Selangor (p. 176).
Harrison, 1964: recorded from North Borneo (Sabah) (p. 22).
Anon./Fed. Malay Sts Mus., 1919: BMNH.1955.1596, male,
Port Weld, Larut, Perak. Wayre, 1974: 'Medway told us that in 1964 a Hairy-nosed
Anon./Fed. Malay Sts Mus., 1922: BMNH.1955.1597, male, Otter had been caught by Aborigines near Janda Baik in the
Kg. Padang, Tembeling, Pahang. Bentong division of Pahang at approximately 550 metres in
a torrent stream' (p. 37).
Mus., G. T., 2.III.1922: ZRC.4.1183, female, Kg. Tembeling,
Pahang, Malaysia. Harrison, 1966: 'It is abundant in the sea off Penang Island.
It is also recorded from Singapore, but not from the sea
Nicholes, D. W. A., 28.VII.1938: ZRC.4.1175, male, Lazarus
Island, Singapore. there.' (p. 223); ' ... have been recorded from time to time,
Anon., 12.VII.1955: SM.OI73/3, female(?), Sungai Mugang, and one suspect that they visit here from the mainland.'
(locality unknown). (p. 6); present in Penang (p. 7); known from Malaya
Ng, P. K. L. et al., V1.1991: Tanjung Karang Road, Kuala (p.332).
Medway, 1977: ' ... specimens indicate that this otter occurs
Selangor, Malaysia.
throughout mainland Borneo, from the coast to small
streams of the far interior' (p. 133).
Records of Lutra sumatrana. Davies & Payne, 1982: most information come from chance
sightings and published works; large otters (L. sumatrana &
Published in literature: L. perspicillata) are seen along the Kinabatangan river. The
Cantor, 1846: [as L. barang RafHes] "Mumrang' or 'Amrang' otters are not regarded as sufficiently rare or threatened by
of the Malays of the Peninsula.' 'The Malayan individuals development to warrent special conservation methods now
appear to attain to a greater size than the Sumatran, de- (p. 141-146).
scribed by RafHes'; Distr. - Malayan Peninsula, Borneo Foster-Turley & Santiapillai, 1990: ' ... the current existence of
(p. 195). ... the Hairy-nosed otter ... is unconfirmed', in Peninsular
167
Malaysia. 'In Sabah and Sarawak, ... at least three species Foster-Turley & Santiapillai, 1990: The current existence of
of otter occur', including the Hairy-nosed otter (p. 58-9). the Eurasian otter on Peninsular Malaysia is unconfirmed.
Foster-Turley & Santiapillai, 1990: Not reported in Peninsu- (p. 58); Listed in distribution map as 'maybe extinct' in
lar recently, possibly still present in remote areas, in scat- Peninsular Malaysia and Singapore.
tered localities in East Malaysia; believed extirpated in Sin-
Museum records (sp.?): see Payne et al. (1985: 280).
gapore.
Anon, 18X1.l959: SM.0173/4 [Lutra sumatrana], male, Bai
Yang eta/., 1990: In Singapore, current status unknown.
Rio, Kelabit, Sarawak.
Nor, 1990a: Only the Smooth Otter and the Small-clawed
Anon, 29.III.1961: SM.0173/2 [uniden.], male, Bai Rio,
Otter found during survey of northern Malaysian states. No
Kelabit, Sarawak.
other species observed.
Museum records:
Borneo References
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Hydrobiologia 285: 171-175, 1994.
Hatch rates of green turtle eggs in Sara wak
Charles M. U. Leh
Curator of Natural History, Sarawak Museum, Jalan Tun Hj. Openg, 93655 Kuchong, Sarawak,
Malaysia
Abstract
Hatch rates of green turtle Chelone mydas (L.) eggs in the beach hatchery of the turtle islands of Sarawak
varied between 53 % and 96 % during the period 1970-1990. The hatch rates of natural or in situ nests
were 71.8 + 18.4 % and 65.3 + 5.9% at Talang-Talang Besar and Talang-Talang Kecil respectively. There
was no significant difference between the hatch rates of in situ nests and replanted hatchery nests in both
the Talang-Talang islands.
Introduction landing is due in part to water temperature related

to the equatorial climate, currents and rainfall. In
The green turtle is an endangered and protected 1961, only 1 to 3% of the eggs laid occurred in
species in Sarawak with primary nesting sites at the months of November to April (Harrison,
the three turtle islands, Satang Besar, Talang- 1962). Elsewhere there are no published records
TalangBesar and Talang-Talang Kecil (Leh et al., of the effects of the monsoon on turtle egg hatch-
1985; Leh, 1989). Conservation efforts at the ability (Brahim et aI., 1987).
turtle islands including the replanting of turtle eggs The present study attempts to analyse the hatch
began in the early 1950s. Earlier Banks (1937) rates of green turtle eggs during the last two de-
observed that the local rainy season or landas cades in Sarawak. Hatch rates of eggs were com-
(from November to April) drove the turtles away pared between the in situ nests and hatchery nests.
from the feeding grounds and prevented mating The hatch rates of turtle eggs during the wet mon-
on the water surface thus reducing the fertility of soon months are also presented.
the subsequent egg clutches laid. The rough sea
disperses the sand on the beaches which become
unsuitable for egg laying (Harrison, 1951). The Materials and methods
heavy rainfall during this period wets the sand
and drowns the laid eggs. It is necessary for the The green turtle egg replanting data at the
turtles to lay their eggs in pits above the high Sarawak Turtle Islands were obtained from the
water mark in order for the eggs to hatch (Banks, annual turtle conservation records maintained by
1937). Harrison (1951, 1961, 1962) did not the Turtles Board office at the Sarawak Museum.
attempt to hatch turtle eggs in the wet months of Eggs were replanted in the beach hatchery at
the Northeast monsoon. depths recommended by Leh et al. (1985) and
Harrison (1962) observed the monthly egg depths recorded from the natural nests. Natural
laying cycles of green turtles in S arawak and noted nest depth varied greatly.
that the basic factor causing the seasonal trend of
172
Results mean hatch rate of 55.5 % and 65.3 % at Talang-

Talang Besar and Talang-Talang Kecil respec-
The mean hatch rate of green turtle eggs has fluc- tively. Nests replanted in the beach hatchery of
tuated over the years since 1970 (Fig. 1). Between Talang-Talang islands had a lower range of hatch
1970 and 1980, the mean hatch rate was gener- rate of 55.3 to 65.0%. There were no significant
ally over 70%, however, between 1980 and 1990 difference between in situ and hatchery hatch rates
it dropped to around 60 %. on Talang-Talang islands during the May/June
During the 1990 replanting season, the overall season (Table. 1). On Satang Besar island where
mean hatch rate of green turtle eggs on the islands all the eggs laid were replanted, the hatch rate was
was 54.7% (Table 1). Natural in situ nests have a low (45.2%). The hatch rates of in situ nests at
100
----- REPLANTED
0--0 HATCH RATE
80
~ 60
L4J
l!)
~
Z
L4J
W
0::
L4J
a.. 40
20
Fig. 1. Hatch rates of green turtle eggs and the annual percentage of eggs replanted between 1970 and 1990 in turtle islands of
Sarawak.
173
Table 1. Green Turtle egg replanting at the turtle islands of Talang-Talang Besar and Talang-Talang Kecil-
Sarawak in 1990.
ranged from 43.7 to 89.7% and 57.8 to 74.4%
Depth No. No. No. respectively (Tables 2 and 3).
%
of of hatched hatched Hatching results in the hatchery during the
nest eggs monsoon season from November to April the fol-
lowing year showed that there is a high percent-
Talang-Talang Besar
age of loss due to unhatched eggs (Table 4). In
Natural nest (in situ) 9 859 477 55.53
Normal (52 cm-60 cm) 417 39534 22449 56.78 general, the rate of loss of eggs on the three turtle
Natural (64 cm-94 cm) 120 11825 6539 55.29 islands can be summarized as follows:
Sub-total 546 52218 29465 55.87
Talang-Talang Kecil Month Mean percentage Range Sample

Natural nest (in situ) 7 675 439 65.31 unhatched eggs (%) size (nests)
Normal (52 cm-60 cm) 315 30353 19716 64.96
Natural (67 cm-74 cm) 12 1141 677 59.33 January 82.0 1
April 69.0 28.0-100.0 10
Sub-total 334 32169 20832 63.11
May 59.0 55.0-63.0 2
June to 45.4 19.0-48.0 20
Pulau Satang Besar
September
Normal depth 51 4482 2028 45.24 35.0-64.0
October 60.5 10
(52 cm-60 cm) 43.0-88.0
November 59.0 16
Total 931 88869 52325 54.74
Table2. Hatch rates of in situ Green Turtle nests on Talang-Talang Besar in 1990.
Nest Depth of Date Incubation No. of eggs No. of young Hatch

no. nest (cm) replanted period (days) replanted hatched rate (%)
ttb3 73 7.5.90 52 73 45 61.64

ttb4 nest destroyed by landing turtles 0
ttb5 82 7.5.90 53 97 87 89.69
ttb6 nest destroyed by landing turtles 0
ttb7 94 7.5.90 51 95 74 77.87
ttb86 64 19.6.90 59 135 96 71.11
ttb87 82 19.6.90 55 81 79 97.53
ttb9i 74 20.6.90 59 99 58 58.58
ttb92 73 20.6.90 61 87 38 43.67
Mean hatch rate = 71.87 + 18.42 omitting the 2 lost nests.
Table3. Hatch rates of in situ Green Turtle nests on Talang-Talang Kecil in 1990.
Nest Depth of Date Incubation No. of eggs No. of young Hatch

nest (cm) replanted period (days) replanted hatched rate (%)
ttk12 70 10.6.90 49 82 61 74.39

ttk16 67 12.6.90 57 107 67 62.62
ttk17 74 12.6.90 57 116 67 57.76
ttk19 73 13.6.90 55 84 51 60.71
ttk23 74 19.6.90 58 96 67 69.79
ttk24 69 19.6.90 51 105 73 69.52
ttk25 70 20.6.90 51 85 53 62.35
Mean hatch rate = 65.31 + 5.98.

174
Table 4. Records of hatchery replanting programme of Green Discussion

Turtle eggs in 1989 and 1990 in Sarawak.
Date Eggs Nest Eggs Percentage There are numerous studies on the status of ma-
replanted depth hatched infertile rine turtles in Sabah (de Silva, 1986, 1987), Sa-
(cm) rawak (Leh et al., 1985; Leh, 1989) and Peninsu-
lar Malaysia (Chua & Furtado, 1988a & b;
Pulau Satang Besar
Mortimer, 1991). There are few published record
Year 1989
24.4.89 10952 55 50 of hatch rates of green turtle eggs in Malaysia
26.4.89 100 52 0 100 although turtle hatcheries operate in several
26.4.89 86 52 0 100 states. Leh et al., (1985) indicated the hatch rates
28.4.89 96 52 20 79 of green turtle eggs at different replanting depths
9.11.89 120 52 67 44
in the hatchery in Sarawak. At a nest depth of
16.11.89 85 52 22 74
26.11.89 92 52 41 55 60 cm, the hatch rates of nests ranged from 60.0
to 72.7%, while hatch rates for those at depths of
Year 1990
10.10.90 76 60 43 43
56 cm and 52 cm were 47.2 to 76.9% and 37.1 to
17.10.90 85 60 0 100 84.3 % respectively.
23.10.90 94 60 49 48 The present study indicated that the mean
5.11.90 to Total of 608 hatch rate of green turtle eggs in the hatchery was
28.11.90 eggs in 7 nests similar to that in in situ nests. Hatch rates of in-
Year 1991 situ nests also showed a large variation ranging
20.1.91 79 60 15 82 from 43.7% to 97.5% on Talang-Talang Besar
island in 1990. Two in situ nests were lost on the
Pulau Talang Talang Kecil
Year 1989 island in 1990 when landing turtles dug up the
5.4.89 93 52 39 58 nests completely. The low hatch rates in 1990
5.4.89 88 60 47 46 could be due to the weather conditions that year.
3.11.89 92 52 51 44 Poor hatch rates were observed during the wet
15.11.89 91 52 52 43
monsoon months of November to April. The per-
16.11.89 106 60 59 44
30.11.89 97 60 31 68 centage of unhatched eggs was as high as 82 % in
January when heavy rainfall fell on the beach.
Year 1990
16.4.90 107 52 0 100
During the peak egg laying season in September,
16.4.90 100 52 0 100 when there was no rainfall, the hatch rates of eggs
7.5.90 82 52 30 63 were high.
8.5.90 91 52 41 55 Research has to be conducted to look into fac-
17.11.90 116 52 63 46 tors that affect the hatchability of green turtle eggs
27.11.90 106 52 58 45
in Sarawak in order to enhance the conservation
Pulau Talang Talang Besar programme. The use of in situ nests as a conser-
Year 1990 vation strategy is recommended. Complete loss of
15.4.90 93 52 66 29 eggs can occur in a natural nest when another
15.4.90 101 52 73 28 landing turtle invades the old nest site.
7.5.90 98 in situ 0 100
8.5.90 94 in situ 0 100
1.11.90 112 52 58 48
2.11.90 103 52 51 50 Acknowledgements
16.11.90 91 52 15 84
19.11.90 92 60 11 88 The author would like to thank the staff of the
29.11.90 111 52 59 47
30.11.90 91 60 37 59
Turtles Board office and the Natural History Sec-
tion, Sarawak Museum for assistance in obtain-
ing the egg replanting data.
175
References Borneo. 1. Breeding season. Sarawak Mus. J. 5: 593-

596.
Banks, E., 1937. The breeding of the edible turtle (Chelone Harrison, T., 1961. Some new hatching observations. Sa-
mydas). Sarawak Mus. J. 35: 273-277. rawak Mus. J. 10: 293-299.
Brahim, S., E. H. Chan & A. K. Rahman, 1987. An update Harrison, T., 1962. Monthly egg laying cycles. Sarawak Mus.
on the population status and conservation of leatherback J. 10: 624-630.
turtles in Trengganu. In A. Sasekumar, S. M. Phang and Leh, C. M. U., S. K. Poon & Y. C. Siew, 1985. Temperature
E. L. Chong (eds), Proc. 10th Annual Seminar, Malaysian related phenomena affecting the sex of green turtle (Chelone
Society of Marine Sciences, University of Malaya, Kuala mydas) hatchlings in the Sarawak Turtle islands. Sarawak
Lumpur: 69-77. Mus. J. 55: 183-193.
Chan, E. H., 1989. A bibliography on Malaysian sea turtles. Leh, C. M. U., 1989. The green turtle, Chelone mydas (L.) in
Universiti Pertanian Malaysia, Trengganu, Malaysia, 8 pp. Sarawak: Is there a future? In S. M. Phang, A. Sasekumar
de Silva, G. S., 1986. Turtle tagging and international tag & S. Vikneswary (eds), Proc. 12th Annual Seminar, Ma-
returns for Sabah, E. Malaysia. Sarawak Mus. J. 57: 263- laysian Soc. Marine Sciences, University of Malaya, Kuala
272. Lumpur: 219-228.
de Silva, G. S., 1987. The Leatherback and Olive Ridley in Mortimer,J., 1991. Marine turtles. In H. Benard & M. Brooke
Sabah waters. Sarawak Mus. J. 58: 115-122. (eds), Insight guide, Southeast Asia Wildlife, APA Publi-
Harrison, T., 1951. The edible turtle (Chelone mydas) in cations, Hong Kong: 168-171.
Hydrobiologia 285: 177-187, 1994.
The use of artificial reefs in enhancing fish communities in Singapore
Christopher Y. Y. Chua & L. M. Chou

Department of Zoology, National University of Singapore, 10 Kent Ridge Crescent, Singapore 0511
Key words: artificial reefs, fish communities, resource enhancement
Abstract
Intense development of the coastal zone in Singapore has resulted in the degradation of much of the
marine ecosystem. In order to restore and enhance fish communities of denuded areas, an artificial reef
consisting of a tyre reef and a concrete reef, was established in the vicinity of the southern islands of
Singapore. Results from fish visual censuses after the establishment of the artificial reef indicated an
increase in numbers of juveniles and adults. A total of 37 and 32 fish species were recorded over a period
of 11/2 years at the concrete and tyre reefs respectively. The dominant fish families were Pomacentridae,
Labridae, Chaetodontidae, Apogonidae, Gobiidae and Nemipteridae. The artificial reefs also serve as
a nursery ground for some species (e.g. Neopomacentrus sp.) which are important primary consumers
of algae on natural reefs. Greater numbers of 'target' (food-important) fishes were observed at the
concrete reef while the tyre reef harboured more juveniles and smallersized adults. The results indicate
that the concrete modules were more effective than the tyre reef in terms of fish abundance per unit
volume. Such structures can enhance the biological resources of relatively unproductive areas.
Introduction culture (Chia et al., 1988). Fish catch has declined

from its peak in the 1940s down through the 1970s
Artificial reefs can rejuvenate the fish communi- and 1980s. This is also reflected in the present low
ties of previously barren stretches of the sea floor. number of licensed fishermen. Though it may not
Experiences in countries like the United States of be practical to revive the local fishing industry to
America, Japan, Taiwan and in recent times, its previous state, in view oflimited seaspace and
some Southeast Asian countries (White et al., intensive use of port waters for other marine-
1990), have shown the varying successes of these related activities, recreational fishing could be
structures in increasing the biomass offish in the promoted as an alternate activity.
surrounding waters. Over-exploitation of existing In the United States, marine recreational fish-
living resources and the urgent requirement to ing is a multimillion dollar business with esti-
increase fish catch to feed rapidly expanding mates of over 20 million fishermen landing 700
populations have resulted in the implementation million kilogrammes of fish annually (Stone,
of extensive artificial reef programmes in these 1978). Artificial reefs there serve a public interest
countries. for the 12 million recreational saltwater fishermen
Much of Singapore's annual fish supply is im- who spent US$ 2.4 million pursuing their hobby
ported, with only 12 % coming from local capture (Sport Fishing Institute, 1983). When well-
fisheries along the coastal waters and from aqua- planned and well-managed, artificial reefs are
178
useful in developing and enhancing fishery cept for those occurring in large numbers which
resources. were estimated using a Log 4 Abundance cat-
It was with this aim that between 1989 and egory. In addition, size estimates (in cm) were
1990, artificial reef modules of different materials made of the 'target' species.
were constructed and established on the sea floor At the artificial reef, fish visual censuses were
adjacent to a patch reef in the southern islands of also used within the vicinity of the modules. This
Singapore. This pilot project was conducted to involved the actual count of fish within the arti-
find out if previously denuded and barren sea ficial reef framework and the seaspace 3 m be-
floor areas can be enhanced and made more pro- yond the perimeter of the modules. The area es-
ductive. timated was 60 m2 and 112 m2 for the tyre and
Recent initiatives propose setting aside certain concrete reef respectively.
areas in the southern islands of Singapore for Surveys at the artificial reef site were conducted
conservation purposes. The findings from this in two phases:
pilot project may serve to accelerate the process
i. Pre-establishment surveys consisting of 150 m
of establishing a national marine park for proper
line transects on the 3 m and 10 m depths of
and well-managed utilisation of the limited ma-
the natural reef slope. Two visual censuses
rine resources.
were carried out at both depths for the natu-
ral reef adjacent to the potential artificial reef
site; one in 1988 and another in 1989. At a
control site 600 m south of the above, a single
census was conducted in 1989.
An artificial reef was established on the sea floor
Exploratory censuses were also carried out on
adjacent to a natural patch reef (Terumbu Pem-
the sea floor on which the artificial reef mod-
pang Tengah) west of Pulau Hantu (Fig. 1). This
ules were to be laid. However, the paucity of
pilot study involved the use of two different ma-
fish observed terminated further investigation
terials (Fig. 2): 1 m3 hollowed concrete blocks,
in this phase.
and tyre pyramids each consisting of 42 disused
tyres weighed down with 50 kg concrete cubes. ii. Post-establishment surveys using similar
These two different materials were placed ap- methods were employed. At the adjacent natu-
proximately 10 m apart on the sea floor at the ral reef, two censuses were carried out in 1990
same depth to determine which was the better at both depths. Similar replicates were per-
material for recruiting fish and encouraging de- formed at the control site.
velopment of sessile communities. Fifty hollow In addition, a total of 17 censuses were carried
concrete 1 m3 blocks were used, with ten double out between 1989 and 1991 at the concrete
stacked above the rest. The 20 tyre pyramids were reef. Similarly, 14 censuses were completed at
placed side-by-side and secured to prevent shift- the tyre reef during this period.
ing and movement.
Fish transect surveys (Dartnall & Jones, 1986)
were used to assess fish communities on the ad- Results
jacent natural reef. A 150 m transect tape was
laid on the reef substratum or sea floor along From Table 1, the fish data collected from the
which a diver swam, performing fish counts 3 m pre- and post-establishment surveys at the adja-
to the left, right and above the line, correspond- cent natural reef showed similarity with only slight
ing to 2700 m3 per transect. Fish were categorised seasonal differences. The preestablishment sur-
into 'indicator' species (Chaetodontidae), 'target' veys showed that at the 10 m depth, poorer fish
species (food-important) and 'other families'. Ac- populations were observed. Two chaetodontid
tual counts were made for all species of fish ex- (,indicator') species (Chaetodon octofasciatus and
SINGAPORE
o Skm
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\ ... ./f ".
I
D'- / 1] . vQj-D~
Ayer Chawm (' . :.-'..... ~~~!I_,
~) ___
,.-,
-"'"'-'-6 · • - ,", Cyrene reefs
a. ~..J
0:.
Pulau DIISing~
Sultan Shoal P.Dukom
Ughthouao . Terumbu ,~-- 0 .. O
Pempang L-':>(')~ ... \ .,.~ P. Seringat
Laut ' ...0 , -t:..........
T P ...,' '--' Q.{)D ... , .
... St. John'. ""~
Tengah ~~~t P.l\anto _ Tenllllbll "'~Sollth Dukom Island ~~'-.) \)J Kusu Island
P. Salo ':2 ....semakao
Terumbo... _"\{~', (": P. Jong
O"~ " • Raya \$:) P. Sakeng ~
U,\ Q~ '\.l Lazarus Island
'\ l~·. The Sisters
P. Sudoal '. •• '.\ \
~ ~"'{-" \ .. "
\ ) ~'i\':J ~P. Sebarok
• -, Det- .....-:" \. .,.; V
tng ,_...... u
....~) Dembnn P Semakau
"._.....,.... Desar· , ...""
c-:-,
\ -...'
'••'
Sechirit Deacon
p. pawal. (, .../ "
'" .... ~.,.,.,..
a," r~'~-''''~~'
",
, ,...,I P. Senang
......... {J P. Diola
Coral reefs
o o Artificial reef
I:
Raffies Ughlhouse
Fig. 1. Map showing the artificial reef site located at Terumbu Pempang Tengah in the southern islands of Singapore.
......
--.l
\0
180
polyc:hyl~t TOpt
diun.6-7mm
(27 + 12 + 3 = 42 tyres)
A. Subunit (3 tyres) of a tyre-pyramid module B. Plan view of a tyre-pyramid module
1m
c. Side "iew of a tyre-pyramid module D. Hollow concrete module

Fig. 2. Tyre pyramids (A, B & C) and 1 m 3 concrete modules (D) used to construct the artificial reef.
Chelmon rostratus) were common at 3 m depths angelfish, Chaetodontoplus mesoleucus, was also
while only C. rostratus was recorded during the common at both depths here. The data collected
10 m transects. 'Target' species were lacking with on the sea floor of the potential artificial reef site
only a school of Caesio teres in the 1988 survey during the preestablishment period indicated an
(3 m depth) and a single individual of Lutjanus absence of 'target' (foodimportant) and 'indicator'
carponotatus (10 m) in the latter survey. Poma- species. An unidentified nemipterid species and
centridae and Labridae (,other families') were the some gobiids were the only fish types observed
dominant families in terms of diversity and abun- here.
dance while at the 3 m depth, the common fish Table 1 also shows the results from surveys
genera were Neopomacentrus, Paraglyhidodon, conducted after the establishment of the artificial
Pomacentrus, Halichoeres and Choerodon. Other reef. These were carried out in February and June
families common along this depth included 1990. The abundance of the chaetodontid species
Apogonidae, Gobiidae and Nemipteridae. The remained unchanged except for Chaetodon octo-
181
Table 1 Pre- and post-estabhshment fish transect (150 m) data at Terumbu Pempang Tengah (actual counts except * Log 4
abundance categones)
Famdy Taxon Artificial reef SIte Control SIte (east)

3 m depth 10 m depth 3 m depth 10 m depth
Pre-E Post-E Pre-E Post-E Pre-E Post-E Pre-E Post-E
1988 1989 1990 1990 1988 1989 1990 1990 1989 1990 1990 1989 1990 1990
5/8 22/6 28/2 516 7/7 15/6 28/2 516 17/5 28/3 20/8 17/5 28/3 20/8
'IndIcator' speCIes
ChaetodontIdae Chaelodon octofasctalUs 11 4 3 26 1 17 3
Che/mon roslralUs 5 3 6 8 5 7 9 3 3 7 2 2
CoradlOn chrsozonus 3 1 1 1
Target' speCIes
CaeslOmdae CaeslO teres 3* 4* 3* 5' 4* 4* 5' 4*
CarangIdae SelarOldes leptolepsIs 3* 5"
A/ectls sp (clltarls?) 4*
CentropomIdae Psammoperca watgtenslS 1
Lutjamdae Luljanus carponotalUs 29 25 1
Serramdae Cephalopholts pachycentron 1 4 2 2
Plectropomus maculatus 2
SIgamdae Siganus guttatus 4 1
Other famIlIes
Pomacentndae Abudefduf benga/ensls
A saxattils
Amb/yg/yphldodon curacao 2"
A leucogasler 1
AoguoeU/b frenatus 2*
A coe/laTls 2*
Dascyllus trtmacu/alUs 1
DlschlStodus prosopotaemae 7 3
Neopomacentrus nemurus 4* 4* 5* 2* 4* 5' 4* 4* 3 3* 4*
N taemurus 4* 4* 2* 4* 4* 4*
Parag/yphldodon mgrorts 4* 2* 2* 3* 2 4'
Pomacentrus alblmaculus 4" 4' 2* 5* 3* 4* 5' 5* 4' 4' 4* 5* 4* 3
P a/exanderae 2* 2
P Itttora/is 4* 3" 4" 3* 3' 3" 2"
P moluccensls 2'
P rhodonotus 2*
Pomacenlrus sp 1
Labndae Che/mus fasclatus 2 2 2 2
Choeodon ancorago 2* 3* 1 5 3' 2
C shoen/emll 2" 2 4
H altchores ch/oropterus 2* 4* 5 3 2 2
H dussumleTl 2' 4* 4* 4' 4 4 4" 4" 4' 11 1 4' 3*
H hoevem 1 3" 2" 3* 3* 1 2 6 2 2
H me/anochtr 2*
H scapu/aTls
Haltchoeres sp 1 3
GrammIsudae Dip/opTIon btfasctatus 4* 3" 3* 3 11 16 9 4 4 29 8
PomacanthIdae Chaetodontop/us meso/eucus 3* 3* 3' 3* 5 13 10 25 27 3 7 5
Pomacanthus sexstrtatus 1 1 1 1 I
Apogomdae Apogon compressus 1 10
A sea/el 3 10
Chellodlpterus macrodon
Blenmdae Melacanthus grammlslts 2
Gobudae Goby sp 1 1 2 2
MogdOldIdae Parapercis sp 1
Nemlptendae Pentapodus canmus 2 6
P setosus 4
Seoiopsis bl/meatus 2 2
S Cillatus
S dublOSUS 2
S vosmeTl 1 2
Pemphendae PempheTls sp 4
Scarldae Scarus sp 6 2
Total 220 262 189 624 33 120 611 451 203 195 296 464 83 94
182
Jasciatus which increased along the 3 m transect. and Pomacanthus sextriatus, apogonid Apogon
Another chaetodontid, Coradion chrysozonus, was compressus, soapfish Diploprion bifasciatus, blenny
also noted. The 10 m depth also saw an increase Meiacanthus grammistes, cavefish Pempheris, ne-
in chaetodontid numbers. Species recorded in this mipterids Pentapodus setosus and Seolopsis vos-
post-establishment period included fusilier Cae- meri, pomacentrids Neopomacentrus nemurus,
sio, scad Selaroides leptolepis, seabass Psammo- Pomacentrus albimaculus and P. alexanderae and
perca waigiensis, snapper Lutjanus carponotatus, labrids Choerodon ancorago, C. schoenleinii and
groupers Cephalopholis and Plectropomus and si- Halichoeres spp. Also seen were Arothron sp. and
ganids Siganus guttatus. Snapper abundance also Upeneus tragula but nine were exclusive to the
increased significantly with 29 and 25 individuals artificial reef. These were two Acanthurus spp.,
in the February and June surveys respectively. two Platax spp., Pterogogus sp., Monacanthus
'Target' species remained low at the 3 m depth spp., Scarus ghobban, Plotosus sp. and Sphyraena
while pomacentrids and labrids remained high. flavicauda?
At the 10 m depth, an increase in fish abundance A higher diversity (five) offish species was ob-
was noted although the overall diversity still did served at the concrete reef and 26 species were
not surpass that at the 3 m depth. Slight increases common to both tyre and concrete reefs. On the
in Diploprion bifasciatus and Chaetodontoplus me- whole, greater n1;lmbers of 'target' species like
soleucus were also observed. haemulids, snappers, grouper Cephalopholis
The transects carried out at the adjacent natu- pachycentron and siganid Siganus guttatus were
ral reef after the artificial reef was established observed at the concrete than the tyre reef. On the
showed an increase in 'target' species at the 10 m other hand, more juveniles of Selaroides leptolepis
depth of the patch reef. This included snappers and Plectropomus maculatus were recorded from
Lutjanus carponotatus, rabbitfish Siganus guttatus, the tyre reef. Of significance is the presence of two
groupers Cephalopholis, Plectropomus maculatus species of batfishes, Platax orbicularis and P. pin-
and scads Selaroides sp. Also found here were natus. These were consistently observed at the
increased abundances of chaetodontids, soapfish concrete reef though not at the tyre reef. Also
Diploprion bifasciatus, small angelfish Chaetodon- present at the concrete reef were two large lut-
toplus and some pomacentrid species. At the 3 m janids each measuring approximately 0.9 m.
depth, pomacentrid and labrid diversity and These were not observed at the tyre reef though
abundance remained high while that of 'target' a smaller lutjanid (0.4 m) was seen here on one
species was depressed, with a record of only one occasion. Monacanthids were also larger in size
species, Caesio teres. at the concrete reef.
With the establishment of the artificial reef, a Juveniles and smaller-sized adults were more
total of 17 and 14 fish visual censuses were car- numerous at the tyre reef. Examples of these were
ried out at the concrete modules and tyre pyra- Diploprion bifasciatus, Chaetodonoplus mesoleucus,
mids respectively within a period of 16 months. Neopomacentrus nemurus, Apogon and Sphyraena.
Tables 2 & 3 and Fig. 3 show the abundance of
fish species recorded at both the concrete and tyre
reefs from all surveys (1989,1990 and 1991). Fig- Discussion
ure 4 shows the increase in the number of fish
species at both the tyre and concrete reefs. The The total of 37 species at the concrete reef and 32
total number of species was 37 at the concrete at the tyre reef within the short time of I' /2 years
reef and 32 at the tyre reef. The 'target' species may be considered good in view of the small di-
comprised 24.3 % at the concrete reef while a mensions of the artificial reef. The increase in fish
higher percentage of 31.25 % was recorded at the abundances after the installation of the artificial
tyre reef. Some of the fishes in the 'other families' reef has shown the effectiveness of artificial reefs
included angelfishes Chaetodontoplus mesoleucus in aggregating fish and improving the productiv-
Table 2. Fish Visual census data at the concrete reef (actual counts)
Fannly Taxon Census No 4 5 6 10 11 12 13 14 IS 16 17 Remarks
Year 1989 1990 1991
Ddte 14/9 13/10 25/10 26/10 27/10 7/11 24/11 22/12 12/2 13/2 19/2 12/3 16/3 4/6 17/9 15/10 22/2
~]ndlcator'
specIes
Chaetodontldae ehe/mon rostralus 2 2 2 2
CoradlOn chrysozonus I I
'Target' specIes
CarangIdae Caranx sp 1 19
SelarOides fepto/epis 30J 30 20
Haemuhdae Plectorhynchus P'Ctus I 2 I 3 4 4 3 TL 25-60 em
LutJanldae Lutjanus carponotatus 5 2 2 2 2 2 TL 25-40 em
Luyanus sp 2 2 2 2 3 2 2 TLlm,40em
Serramdae Cephalophoils pachycentron 4 2 I I TL 9-15 em
Plectoropomus maculatus I
Sigamdae Slganus guttatus 4 4 3 2 TL 30-40 em
S Javus TL 20,30 em
Number of adult target fishes I 32 32 0 0 11 14 12 12 12 10 17
Other fanuhes
Acanthundae Acanthurus sp
ApogoOldae Apogon compressus -70
A cyanosoma 2 -50
Apogon JUl1emles +(2) +(3) + +
Blenmdae Melscanthuf grammlftes 2
Ephlppldldae Platax orbrcu/afls 4 5 3 10
P pmnatus 2(70) 9 12
Goblldae small spp + + + + + + + + + + + + + + +
GrammlStldae DIp/oprlOn bifasclatus I I 5J 3J I I I 2
Labndae Choerodon ancorago I I
C schoenlemll 4 2 I TL 15-45 em
Habchoeres dussumlen + + + I 2 4
H scapu[ans'J +
Pterogogus sp 1(25)
Juvemles (umdennfied) +
Monacanthldae Monacanthus chmensis 2(15) I 6
M macrurus 1(15) 4(14) IJ 3
Mulhdae Upeneus tragula
Nemlptendae Pentapodus setosus + + + + +
SeQ/apsIs vosmen 2(18)
Plotosldae PlotOfiUS sp -20
Pomacanthldae Chaelodontopius meso/eucus 2 4
Pomacanthus sextrmtus I 2 TL 40-45 em
Pomacentndae Neopomacemrus nemurus +J +J +J +J +J + + + + + + + + + + +
Pomacentrus alblmacu/us I 4J I 2
P alexanderae
Scandae Scarus ghobban 1(15)
Tetraodontldae Arothron sp 1(60)
Sphyraemdae Sphyraena flavrcauda? +J +J +J +J +J +J +J 7(20) 6 12 9 7(25)
Total number of adult fishes (abundanee)6 IS II 16 45 43 28 28 49 39 32 37 36 51

Total number offish speeIes4 10 10 II 14 19 16 14 15 17 15 II 17 19 22
Other mobIle mvertebrates
cowne Cypraea talpa
holothunan Stlchopus vanegatus
Hatworm Tvsanozoon sp .....
00
(TL Totd! lengths (em) of selected fishes 1D parentheses •• + 4 presence, 'j' JuveDlles) W
00
.j::>.
-
Table 3. Fish ViSUal census data at the tyre reef (actual counts)
Fanuly Taxon Census No 4 10 II 12 13 14 Remarks
Year 1989 1990 1991
Date 13/10 25/10 26/10 27/10 7/11 22/12 13/2 19/2 12/3 16/3 4/6 17/9 15/10 22/2
·Indlcator' specIes
ChaetodontIdae ehe/mon rostratus 2 2
Corad,on chrysozonus 2
Target' speCIes
CaraogIdae Selaroldes lepta/eptS - 100) - 100 )
CaesJODIdae CaeslO teres 6(14)
Haemuhdae Plectorhynchus p,ctus 2(25) 1(20) 2(30) 4 TL 20-42 em
Centropomldae Psammoperca va/glens,s I TL 20em
Lut)amdae LUljanus carponotalu~ 4 10 I TL 15-35 em
LUljanus sp 2 1(40)
Serramdae Cepha/ophoils psycentron 2(15) 2(15) 2(20)
Pleclropomus maculatus 1(25) 4(40)
Smgamdae Slganus gutlalUS 2 TL 22-35 em
S Javus 3(18)
Number of adult target fishes 4 4 10 13
Other famdles
Acanthundae Aeonthurus sp 2
Apogomdae Apogon cyanosoma -20
Apogon Juveniles - 100) +) +) +) +) +)
EphlPPldldae Platax orblculans 2 3
Grammlstldae Dlp/opr/on bifasclatus I I 3 4 6 I
Labndae Choerodon ancorago 2 I 2 TL 13-25 em
C schoen/emil 5 3 6 2 TL 15-45 em
Hallchoeres dussumle" 4 4 4 3
H scapulans' +
Pterogogus sp 4 TL 10-25 em
Monacanthldae Monacanthus chmensls 5(10) 2 TL 6-15 em
H rnacrurus 1(18)
Nemlptendae Pentapodus setotus + +
SeolopSls vosmen I TL 10-18 em
Pemphendae Pemphens sp
Pomacanthldae Chaetodontoplus mesoleueus 4 4 4
Pomaeanthus sexstnatus 2 I 2 TL 15-40 em
Pomacentndcle Neopomaeentrus nemurus +) +) +J +) +) +) + + + + + + + +
Pomacentrus alblmaculus I) I 2
Scandae Scarus ghobban 2(40) 1(40) 3 TL 15-40 em
Sphyraemdae Sphyraena flaVleauda:;' -100) - 50) -20 7(15) 17(15) 15 19 30 II TL 20 em
Total number of adult fishes (abundance) 16 10 4 17 25 28 30 49 19 37 55 53 30 60

Total number offish speCIes 12 6 14 16 II 12 15 \0 10 18 9 16 17
Other mobIle mvertebrates

sea urchm Dwdema setosum
sea-anemone Radlanthus:;' sp
(TL Total lengths (cm) of selecteffishes In parentheses, • +' presence, 'J' Juvemles)
185
70
60
50
40
30
20
10
o ; concrete reef
+ ; tyre reef
o
Sep Oct Nov Dec Jan Feb Mar Apr!May Jun Jul Aug Sep Oct Nov Dec Jan Feb
1989 1990 1991

TIME
Fig. 3. Total fish abundances at the concrete and tyre reefs between 1989 and 1991. Bars show means ± SE for replicates dur-
ing that month.
ity of the oncebarren sea floor. Studies on the low only juveniles and smaller adults were observed.
profile tyre reef in Brunei Darussalam showed 55 Such small dimensions excluded larger fishes
species offish after five years (Chou et al., 1991). while at the same time, provided more protection
In the Philippines, 81 species were recorded at the for smaller-sized fishes. The darker interior of the
then two-year old but larger tyre reef at Dum- tyre reef also provided suitable cover for some
aguete (Alcala, 1979). Common reef fishes made cryptic species such as the seabass Psammoperca
up 70% and 63% of the total species richness at waigiensis and the grouper Plectropomus sp.
the concrete and tyre reefs respectively, the re- The large dimensions of the more open con-
maining being the 'target' species. crete modules gave better shelter for larger-sized
The physical differences in size and configura- fish, such as the snapper Lutjanus sp., batfish
tion of the two artificial reef modules contributed Platax sp., angelfish Pomacanthus sextriatus and
to the differences in fish counts. It can be seen sweetlips Plectorhynchus pictus and more room for
that the smaller cavities within the tyre pyramids them to swim and manoeuvre within the modules.
restricted the size of fishes sheltering there, thus The investigators were able to approach to within
186
26
24
22
20
j 18
u
~
.CI 16
~
....
Q
14
""
~
l 12
i
(I)
10
8
~
6
~
4
o : concrete reef
+ : tyre reef
2
0
Sep oct Nov Dec Jan Feb Mar Apr May Jun Jul Aug SepOct NOV'~ Dec Jan Feb
1989 1990 1991

TIME
Fig. 4. Number offish species (richness) recorded at the concrete and tyre reefs between 1989 and 1991. Bars show means ± SE
for replicates during that month.
1 m or less of these large fishes without alarming attributed the attraction of juveniles to artificial
them. reef modules to the reduced competition for un-
Concrete modules were more effective (based claimed territories provided by the newly-created
on a higher fish abundance per unit volume) than habitats of the artificial reef modules. While some
the tyre reef. In terms of species richness though, species stayed on in the modules themselves, oth-
there was little difference (five species). However, ers were noted to be absent after having been
the presence of the tyre pyramids are also impor- initially recorded. This suggests a third role for
tant in enhancing fish communities. First, the re- the artificial reef, as a nursery for more pelagic
cruitment of juveniles to the artificial reef is es- species.
sential in establishing a viable community. These In recent years, efforts have been mounted by
initially serve as prey for larger predators seeking volunteer organisations to set aside some areas in
shelter in the artificial reef. Second, survivors the southern islands of Singapore for the purpose
continue to grow and subsequently build an iso- of conservation. Artificial reefs, if established in
lated community at the artificial reef. Stone (1979) these areas, can serve as a resource enhancement
187
tool because oftheir ability to aggregate adult and Coastal Resources Management Project funded
juvenile fishes, and in time serve as recreational by the United States Agency for International
fishing grounds once the fish communities have Development. Overall management was provided
stabilised. Close proximity to natural reefs will by the International Center for Living Aquatic
double the carrying capacity and biomass of the Resources Management. The study was under-
fish community in that area. In Japan, Kakimoto taken by the Reef Ecology Study Team of the
(1979) recorded the same fish type caught at an Department of Zoology, National University of
artificial reef compared to a natural fishing ground Singapore.
nearby. It was also estimated (Stone et al., 1979)
that by providing 'rough bottom' habitats on the
sea floor and thus creating artificial underwater
rocky features, the productivity of the once bar- References
ren sea floor will be increased. The installation of
more artificial reefs in such conserved areas will Alcala, A. C., 1979. Fish standing stock and yield of an ar-
then meet the aims of resource enhancement. tificial reef off Bantayan, Dumaguete City, Philippines. Sil-
liman Journal 26: 253-262.
Should such areas be made available for recre- Chia, L. S., K. Habibullah & L. M. Chou, 1988. The coastal
ational fishing, they must be effectively managed environmental profile of Singapore. ICLARM Technical
to ensure long-term gains. Without such manage- Reports 21. International Center for Living Aquatic Re-
ment, artificial reefs only serve to deplete existing sources Management, Manila, Philippines, 92 pp.
Chou, L. M., G. S. Y. Lim & C. B. Leng, 1991. An assess-
species from the area (White et at., 1990), creat-
ment of the fish communities of artificial reef structures in
ing negative consequences. Brunei Darussalam, with recommendations for manage-
It is too early to say that this artificial reef has ment and development. Resource Management and Opti-
stabilised because of its close proximity to the mization, 9: 15-31.
natural reef which encourages movement of fish Dartnall, A. J. & M. Jones (eds), 1986. A manual of survey
methods for living resources in coastal areas. Australian
populations between them instead of isolating a
Institute of Marine Science, Townsville, 160 pp.
community at the artificial reef. However, these Kakimoto, H., 1979. Artificial fish reefs in Japan Sea coastal
preliminary findings highlight the effectiveness of regions. Proc. 7th Japan-Soviet Joint Symp. Aquaculture,
artificial reefs, even in the turbid waters of Sin- Sept 1978. Tokyo, Japan.
gapore. Sport Fishing Institute, 1983. The marine recreational fishing
industry and opportunities for development. Final Report
Further monitoring is being carried out over a
to the National Marine Fisheries Service. Washington, D.C.
long term to determine whether the use of artifi- Saltonstall-Kennedy Contract no. NA82AA-H-00054 -
cial reefs as a resource enhancement tool is most Phase 2. 83 pp.
cost-effective in areas of low productivity and as Stone, R. B., 1978. Artificial reefs and fishery management.
such, to advocate the use of well-planned artifi- Fisheries. 3: 2-4.
Stone, R. B., H. L. Pratt, P. O. Parker, Jr. & G. E. Davis,
cial reefs to change relatively unproductive habi-
1979. A comparison of fish populations on an artificial
tats into more dynamic and productive ones. and natural reef in the Florida keys. Mar. Fish. Rev. 41:
1-11.
White, A. T., L. M. Chou, M. W. R. N. De Silva &
Acknowledgements F. Y. Guarin, 1990. Artificial reefs for marine habitat en-
hancement in Southeast Asia. ICLARM Education Se-
The artificial reef project was carried out as the ries 11. International Center for Living Aquatic Resources
Singapore component of the ASEAN-US Management, Philippines, 45 pp.
Hydrobiologia 285: 189-201, 1994.
A. Sasekumar. N. Marshall & D. J. Macintosh (eds). Ecology and Conservation of Southeast Asian Marine and 189
Threats to the indigenous freshwater fishes of Sri Lanka and remarks on

their conservation
Rohan Pethiyagoda
Wildlife Heritage Trust, 95 Cotta Road, Colombo 8, Sri Lanka
Key words: Sri Lanka, conservation, freshwater fish
Abstract
Sixty-two freshwater dispersant (26 endemic), 26 saltwater dispersant and 20 exotic fishes have been
recorded from Sri Lanka's inland waters. The discovery of nine new species of freshwater fish during
the past decade suggests that Sri Lanka's fish fauna is not yet well known and that further discoveries
can be expected. All the new species, and almost all the endemic ones, have been recorded from for-
ested areas in the foothills of the southwestern wet zone and the Knuckles hills. The known ranges of
many species are exceedingly small; almost all of them are habitat specialists. 25 species have at least
two strong habitat preferences (in almost all case, shade being one), though a majority (32 species) have
none. Habitat alteration is therefore the greatest threat to their survival.
Eight freshwater fish species are considered endangered and a further five, vulnerable. Three species
are rare, 18 common and 13 abundant. All but two of the endangered species and all the vulnerable
species are endemic in Sri Lanka. Nine species are each restricted to only one drainage basin, and seven
have a known range < 50 km 2 • Three species of diadromous gobies known only from small populations
are considered vulnerable.
Examination of the existing threats suggest that deforestation, the widespread dispersion of exotics,
pollution caused by agricultural chemicals and increasing pressure from the food fishery present the
greatest threats to fish populations. While the endemic fishes are restricted to or most abundant in the
wet zone, the overwhelming extent (> 95 %) of Sri Lanka's nature reserve areas are in the dry zone. A
programme to maintain self-sustaining captive populations of the endangered species is urgently nec-
essary.
Introduction and three diadromous gobiids are restricted to the

island (cf. 27% estimated by Erdelen, 1988). The
Recent surveys by Senanayake (1980) and taxonomy of the fishes of South Asia is very con-
Pethiyagoda (1991) suggest that populations of fused (Kottelat, 1989) and future revisions could
several species of freshwater fishes in the island well demonstrate a much higher degree of ende-
of Sri Lanka are in decline. I have shown else- mism.
where (Pethiyagoda, op. cit.) that the degree of Although IUeN (1990) lists two Sri Lankan
endemicity of the Sri Lankan fishes is much higher fishes as endangered, six as vulnerable and two as
than was previously supposed. At least 26 fresh- rare, my observations show that the status of sev-
water dispersants (42 % of a total of 62 species) eral other taxa is critical, and that unless early
190
remedial action is taken, extinctions may follow. haweli River drainage (see Fig. 1). The remaining
This paper seeks to identify the causative factors 'dry zone' province of the island contains mostly
of this decline and the species and biotopes most fish species also shared with India and the wet
at risk, and propose relevant conservation mea- zone.
sures.
Methods
Geography
Methods of sampling and collection are discussed
Sri Lanka is a tropical island of 65 610 lan 2 situ- in Pethiyagoda (1991: 16). All major basins were
ated approximately 6 0 -10 0 N, at the southern tip sampled, those of the wet zone (which contain a
of India. It is separated from the mainland by the greater diversity of species and biotopes) more
Palk Strait, in places less than 20 km wide. Fre- intensively than those of the dry zone. Sampling
quent and prolonged connections have occurred was on the basis of catches made by seine and
between the two land masses in the past, the most hand net, visual observations with snorkel and
recent one probably having been during the Upper mask, and the examination of commercial fish
Pleistocene (Jacob, 1949). Kortmulder et al. catches of both the food and aquarium fisheries.
(1990) and Pethiyagoda op. cit., compare aspects
of the ichthyofaunas of southern India and Sri
Lanka.
The central area of the island is mountainous, "".
rising to a maximum 2524 m above sea level. One
hundred and three river basins radiate from these
mountains, less than half these rivers being pe-
rennial. Although Sri Lanka has no natural lakes,
about 2% of the land surface is occupied by sev-
eral thousand irrigation reservoirs. Much of the .' .'
island is dry (precipitation of 750-2000 mm p.a.)
and depends for water on the rains resulting from
the north-east monsoon (November to February)
and the discharge of rivers originating in the cen-
e'
tral massif. The south-western quarter of the is- e'
land, known as its wet zone, enjoys an additional

monsoon from May to September, and obtains
an annual precipitation of 2000 to 6000 mm.
Pethiyagoda op. cit. showed that the endemic ,.
fishes of the island are, by and large, restricted to
its wet zone. This endemicity is attributed to the
south western wet zone being a 'land-island'
biotope rather than insularity caused by the pe-
riodic severance of the land connection with
India. The wet zone itself can be subdivided into
two ichthyofaunal provinces, firstly the more or
less contiguous southwest-flowing drainages of eo'
Motorn
.,. Iun
.,.
the Kelani, Kalu, Gin and Nilwala Rivers, and Fig. 1. Sri Lanka: major river basins and areas of high eleva-
secondly the 'land-island' province formed by the tion and rainfall (only the five most important basins are la-
Knuckles hills within the northeast-flowing Ma- belled).
191
Results Habitat alteration and degradation
The endangered and vulnerable (sensu IUCN, The secondary effects of deforestation, urbaniza-
1990: xxiii) freshwater fishes are listed in Appen- tion and gem mining are partially interrelated:
dix 1. Only the most immediately threatened taxa pollution, the loss of shade (and therefore exo-
are treated here; fishes considered rare (sensu genous food sources by way of insects and lar-
IUCN, 1990) but not endangered or vulnerable vae), increased erosion, substrate alteration (in-
have been omitted. cluding the reduction of leaf debris caused by
shade loss), changes in water quality and tem-
Discussion perature, and desiccation due to reduced ground
water detention (Garg and Saxena & Dharwan,
The most recent published list of threatened Sri in Agrawal and Chaturvedi, 1988, review the ef-
Lankan fishes is contained in IU CN (1990). Here, fects of industrial pollution on fishes in India). It
Lepidocephalichthys jonklaasi and Labeo fisheri are is evident from Appendix 1 that the greatest
classified as endangered. Puntius cumingii, P. ni- short-term threat to the endangered fishes is habi-
grofasciatus, P. pleurotaenia, P. titteya, Rasbora tat alteration by deforestation and siltage. Be-
vaterifloris, and Malpulutta kretseri are considered cause distribution and microhabitats suggest that
vulnerable. These 'vulnerable' species are popular many species, particularly the endemic ones are
aquarium fishes and may be under some pressure specialists (Senanayake, 1980; Wikramanayake
for that reason. However, they do not appear to & Moyle, 1989; Wikramanayake, 1990; Pethiya-
warrant classification in this category or even as goda, op. cit.), significant habitat degradation can
rare species as they all have relatively large, widely lead to changes in community structure and de-
distributed populations (each in at least four creased diversity. Such altered habitats tend to be
river basins) and are, with the exception of better suited to habitat generalists and introduced
R. vaterifloris, reasonably tolerant of habitat exotic species (Lowe-McConnell, 1987: 302).
modification. Given the present causal factors, In addition to the advantages shade confers on
none of them appears to be in danger of entering fishes (Helfman, 1981), detrivores and inverti-
the 'vulnerable' category in the near future. vores have obvious feeding advantages in forest
The highest incidence of biodiversity has al- streams, and deforestation would pose an imme-
ready been identified with the lowland wet zone diate threat to these species, especially those in-
of Sri Lanka (> 1000 m. a.s.l., > 2000 mm pre- habiting headwaters (Lowe-McConnell, 1987:
cipitation p.a.), particularly the southwestern 167).
quarter of the island (i.e. the Kelani, Kalu, Gin
and Nilwala River basins) (Senanayake et aI.,
1977; Gunatilleke & Gunatilleke, 1983; Erdelen, Degradation of wet zone forests
1988, 1989; Pethiyagoda, op. cit.). Of the 29 en-
demic fishes, 20 are restricted to this area. A fur- Sri Lanka's historical civilizations had been based
ther four are restricted to the lowlands of the mainly in the dry zone. Until 1815 there was little
Knuckles hills. The remaining five endemic spe- agriculture and only small pockets of population
cies enjoy a wider distribution, but are neverthe- in the wet zone. Large-scale deforestation was
less restricted to the wetter parts of the island. necessary however, for the planting of coffee
Senanayake & Moyle (1982) identified a num- (Coffea), tea (Camellia), rubber (Hevea) and Cin-
ber of factors contributing to the decline of native chona by the British up to 1948. The Central Pro-
fish populations in Sri Lanka: (1) deforestation, vince (5674 km2 ), which comprised almost en-
(2) urbanization, (3) water diversion, (4) gem tirely primary wet zone montane forest,
mining, (5) pesticides, (6) exploitation and (7) the experienced a population increase of 1340 % be-
introduction of exotic species. tween 1850 and 1981. Rudran (1990) however,
192
concluded that 'the human population's numeri- desiccation of headwater streams. Steadily de-
cal increase per se was not the reason for the clining precipitation in the headwater catchments
rapid decline of forested areas'. Much of the de- (a decline of perhaps as much as 25% in the past
forestation had been sponsored, and almost all of century: Abeywickrema et al., 1991: 149) will also
it acquiesced to, by the State. By 1850, 18% of worsen this situation.
the Central Province had been deforested for the While tea plantations occupy much of the cen-
planting of coffee (De Silva, 1981: 294). Natural tral highlands and southern wet zone, much (ea
forests today account for less than 5 % of the area 200000 ha) of the western and southwestern wet
of this headwater catchment from which several zone is planted with rubber. Despite the pollution
major rivers, accounting for 38 % of the total flu- resulting from the processing of latex, rubber is
viatile runoff of the island, originate. Since 1948, tolerant of ground cover (obviating the need for
deforestation has been largely for the harvesting herbicides) and appears to have been a more be-
of timber, fuel-wood or vegetable and tobacco nign crop insofar as the fishes are concerned. The
(Nieotiana) cultivation. presence of ground cover also contributes to re-
Peeris (1975) showed that of the 830 endemic ducing erosion, and the relatively tall (10 m) trees
flowering plants in Sri Lanka, 326 (39%) were confer a high degree of shade to waterways in
restricted to the moist lowlands of the island. rubber plantations.
Gunatilleke & Gunatilleke (1983) noted, however, Habitat degradation is all the more significant
that only 9.1 % of the lowland wet zone was still since 25 of the 62 Sri Lankan freshwater species
forested (i.e. 1.45% of the land area of the is- have at least two strong habitat preferences (shade
land), and estimated that less than 22% of this usually being one). 32 species however (mostly
forest was (as at 1983) still undisturbed. non-endemics), are habitat generalists.
Much of the central and southwestern hills
(which are also headwater drainages) are planted
with tea, this crop now accounting for more than Degradation of wet zone swamps
200000 ha (Land use maps, Sri Lanka Survey
Dept., 1980-84). The immature brown loam soil Although Sri Lanka has no natural lakes (and
of the lowland wet zone hills on which tea is therefore no specialized lacustrine fishes), a few
grown is prone to heavy erosion, a situation ex- endemic fishes have a strong association with
acerbated by high rainfall and the deliberate in- still-water habitats. Some of these are essentially
hibition and removal of ground cover. Abeywick- marshland species, e.g. P. eumingii (Kelani basin),
rema et al. (1991: 105) estimate that as much as P. vittatus, Horadandia atukorali (possibly not en-
30 cm of top soil has been lost from upland areas demic: see Rema Devi & Menon, 1992), Heterop-
during the past century since tea was introduced, neustes mierops, Pseudosphromenus eupanus, while
equivalent to 40 MT ha - 1 yr - 1. The consequent others are adapted also to slow-flowing waters,
loss of soil fertility has resulted in this agriculture e.g. P. titteya, M. kretseri, Maerognathus aral. It is
becoming largely dependent on fertilizers. Some regrettable, therefore, that the lowland wet zone
40000 MT offertilizer (NPK) are utilized by the marshes, identified also as a floral habitat of great
tea industry each year. The montane ichthyofauna interest (e.g. Kostermans, 1979) have almost
of Sri Lanka is noticeably depauperate, compris- completely given way to rice cultivation. The
ing only five species, all of them also found in the compulsory seasonal desiccation of rice fields and
lowland wetlands. I have elsewhere commented their high agrochemical usage (Abeywickrema
(Pethiyagoda, 1991: 26) on the possibility of ex- et al., 1991: 106; Mubarak, 1987: 93) result in this
tinctions having occurred in this fauna as a result habitat now being of little value to fishes, even as
of large-scale habitat alteration. a drought refuge.
Deforestation has also resulted in reduced
ground water retention and therefore the periodic
193
Degradation of water and substrate quality 'successfully' to capture fishes by poisoning in

shallow-water lagoons and mangroves (pers.
The surface water draining from the granitic cen- obs.). Although toxicity to aquatic fauna has not
tral hills of Sri Lanka is normally clear, soft (2- been assessed for the pesticides in use, damage to
5 DH, conductivity 18-3 j.1S at ca 22°C) and aquatic biota should be expected.
more or less neutral (5.8-7.0 pH) (Costa & Star-
milhlner, 1972; Weninger, 1972; Starmilhlner, Introduction of exotics
1984). Lowland standing-water bodies have more
dissolved solids (conductivity 400-1100 j.1S at ca Some 20 species exotic fishes are found in Sri
27°C) and are generally alkaline (7.4-8.4 pH) Lanka's inland waters. Four species ofpoeciliids
(De Silva, 1988: 37-38). While no recent com- introduced to Sri Lanka for malaria control and
parative analysis has been made, there is little accidentally in the period 1930-1960 are known
doubt that continuing deforestation and poor only from small, localized populations.
slope agricultural practices are causing increasing Several species of carps have been introduced
silt age of rivers (J ayewardene, 1992). Using for the food fishery, and of these only Cyprinus
samples taken during the relatively dry period carpio is known to have a self-sustaining popula-
from August to December 1983 however, Dissan- tion. This species is the only one now occurring
ayake & Weerasooriya (1986) concluded that the in many headwater streams above 1500 m. a.s.l.
general chemical quality of the water of the Ma- elevation. while L. rohita possibly breeds in Sri
haweli River was satisfactory for most purposes. Lankan waters, its frequency appears to be de-
In the present survey, comparative sampling clining rapidly (pers. obs.) after the suspension of
(with seine and cast-net) was done in several sets stocking in 1990. The other species, Catla catla,
of otherwise similar, proximal streams, one being Cirrhinus mrigala, Hypophthalmichthys nobilis,
undisturbed while the other was silted and made H. molitrix and Ctenopharyngodon idella do not
turbid by upstream rice fields or mineral extrac- breed in Sri Lanka, and are therefore control-
tion. This showed that several species normally lable. Carassius auratus, was regularly cultured
associated only with clear water and pebble or and introduced until the 1980s, but no longer ap-
gravel stream beds, e.g. P. srilankensis, L. fisheri, pears in inland catches (pers. obs.).
Garra spp., R. vaterifl,oris, M. kretseri, are far less Of the four anabantoid species introduced, only
common in turbid water compared to clear water. Trichogaster pectoralis has a popUlation high
enough to warrant a fishery, but its restriction to
Pesticides standing waters (mostly in coastal marshes) ex-
cludes it from the most fragile wet zone habitats.
The effects of pesticides on Sri Lanka's environ- While Osphronemus goramy, is known from sev-
ment, and their toxicity to the aquatic fauna, have eral small, localized populations, its large size
not yet been assessed. Sim (in Fernando, 1989) and breeding requirements restrict it to the larger
estimated that approximately 1350 MT of insec- water bodies.
ticides, 2180 MT of herbicides and 720 MT of Oncorhynchus mykiss originally introduced as a
fungicides were used by Sri Lanka's agricultural sport fish in 1882, survives in numbers only in the
sector in 1986. The Anti-malaria Campaign is Horton Plains at an elevation ca 2000 m. a.s.l.,
also a major user of insecticide. Somaratne (in where it is now strictly protected by the State. The
Fernando, 1989) states that the tea industry, destructive nature of this fish, which may already
which is located largely in headwater catchments, have been responsible for species extinctions in
utilizes some 400000 I of Paraquat (a herbicide Sri Lanka, is well documented (Day, 1989;
with an LD50 of 150 mg kg- I for rats and fatal to Pethiyagoda, 1991: 26, 33).
adult O. mossambicus at concentrations of Five species of tilapiine cichlids (tilapia, s.l.)
20 ppm) per annum. Pesticides are also used have been introduced to Sri Lanka since 1952,
194
together with a number of hybrids. There is little in pockets; Kottelat (pers. comm.) reported ob-
doubt that Oreochromis mossambicus has, by far, serving til apia in Sulawesi in high-gradient head-
been the most successful of these (De Silva, 1988: water torrents with rocky substrates.
97). Of the annual freshwater fish catch of ca Oreochromis mossambicus breeds throughout
40000 MT, til apia accounts for 56-99%, depend- the year, but more frequently during rains (De
ing on the location (Fernando & De Silva, 1984; Silva, 1985) thereby coinciding with the breeding
De Silva, 1985). However, in the 40 years since of most of the indigenous fishes. The population
its introduction to Sri Lanka's inland waters, no oftilapia in Sri Lankan standing waters also out-
critical assessment has yet been made of its im- numbers and outweighs that of all the other spe-
pact on the indigenous fishes. Instead, the uncon- cies combined. While Costa and Abeysiri (1978)
trolled dispersion of tilapia in Sri Lanka has found that 58-78 % of the diet (excluding sand
probably been viewed with complacence by fish- and detritus) of o. mossambicus 25 mm long com-
eries personnel as it is commonly thought of as prised zooplankton, this fish is also known to be
being a herbivore, and therefore not a threat by an opportunistic predator of small fish (Schuster,
predation to the indigenous fishes. The validity of 1952; Jayaram, 1981: 340; pers. obs.) and when
this assumption is questionable (Arthington & food is scarce, even its own young (Neil, 1966).
Mitchell, 1986), and both fish and fishery may These aspects of til apia biology are usually ig-
pose a threat to the aquatic fauna and flora of Sri nored by most literature dealing with its 'success'.
Lanka. The reason usually attributed to the success of
The success of o. mossambicus in the fishery til apia in major perennial reservoirs in Sri Lanka
has been attributed to many factors. It is a spe- is the lack of 'specialized' lacustrine species in the
cies of high resilience and fecundity. It is able to indigenous fauna (Fernando & Indrasena, 1969;
breed throughout the year; to withstand a high De Silva, 1988). While this may strictly be cor-
degree of pollution; to tolerate, breed in and even rect, there being no lakes in the island, it does not
disperse via sea water; and utilize almost all avail- mean there are no specialized stillwater species -
able sources of food (Maitipe & De Silva, 1985; see comments on wet zone swamps above. De
De Silva, 1988 and references therein). The Silva (1988: 46) listed 25 indigenous species as
present range of o. mossambicus in Sri Lanka occurring in reservoirs, 13 of them in all five res-
includes the entire dry zone, where it is the most ervoirs sampled. Another direct consequence of
abundant species, most (brackish water) estuar- the introduction of tilapia and its success has
ies and lagoons in both the dry and wet zones (De been a dramatic increase in the fishery effort and
Silva & De Silva, 1987) and the larger streams, the introduction of new, more intensive fishery
reservoirs and rivers of the wet zone lowlands up technologies. Almost all reservoirs are now inten-
to 600 m elevation (De Silva & De Silva, 1991; sively fished, often with gill nets of small mesh
Costa & Fernando, 1967; pers. obs.). size (3-5 cm).
Tilapia is however, absent from rapids and Although o. mossambicus is not one of the
streams where the substrate does not consist of ca 800 cichlids native to the African lakes, it is
sand, mud or silt (pers. obs.), but increasing silt- often treated as a specialized lacustrine species
ation of lowland streams could result in this spe- (e.g. the views of Fernando & Indrasena, 1969,
cies invading wet zone forests: o. mossambicus above), probably for the reason that most fresh-
has already been recorded from the borders of the water fish culture takes place in lakes and reser-
knuckles hills at Rattota, the Kanneliya Forest voirs. While it is undoubtedly very successful in
Reserve, the Peak Reserve at Kitulgala and these environments, its natural habitat in the
Opatha, close to the southern margin of the Sin- Zambezi Basin is fluviatile (Trewavas, 1983: 311).
haraja Forest (pers. obs.). In principle there is Breeding takes place in pools, but the fish can
nothing to restrict the dispersion oftilapia into the withstand fast-flowing water (pers. obs.).
central hills, where it has already established itself In the longer term, extensive invasion of the
195
lowland wet zone and the Knuckles hills by ex- Fernando (1991: 27) argued that despite the
otics, most notably til apia, would almost certainly introduction of til apia to Sri Lanka's lowland res-
result in extinctions. Environmental disturbance ervoirs, 'Indigenous fishes have not been nega-
caused by on-going colonization is resulting in tively affected. On the contrary, it appears that
these areas increasingly developing the character- they have benefitted from the presence of tilapias
istics of invadable ecosystems (Muller, 1991; Ar- (De Silva & Fernando, 1980) ... Tilapias probably
mantrout, 1981). More exotics are likely to be reduce predation pressure on indigenous
added to the fishery and informal proposals for cyprinids'. However, while it is arguable that the
the introduction of the nile perch Lates niloticus production of indigenous species taken as a whole
(Centropomidae) and largemouth bass Mi- has increased, possibly as a result of the presence
cropterus salmoides (Centrarchidae) have already of tilapia, there is no evidence to suggest that the
been made (Jonklaas, 1989, in !itt.). production of every species has increased. For
The casual approach to introductions and the instance, the population (and therefore the pro-
citing of a lack of evidence of pressure to indig- duction) of L. porcellus, a cyprinid which was until
enous Asian fishes by exotics (e.g. De Silva, 1989) the 1970s sufficiently abundant in Sri Lanka's dry
is in part because few Asian countries have made zone reservoirs to warrant a fishery (Senanayake,
systematic assessments of their native fish re- 1980), has become critical (Pethiyagoda, 1991:
sources (or faunal diversity in general). Even re- 87).
cent publications offaunas in many Asian coun- There is little doubt that almost all exotics do
tries are based more on dated literature or old exert pressure on the host fauna. Such pressures
museum collections than on recent surveys. In need not necessarily be the result of direct influ-
these circumstances, it is unlikely that extinctions ences such as predation or competition, but could
will become known until long after they have oc- be the result of indirect influences such as in-
curred. In fact, the state of taxonomic knowledge creased fishery pressure or potentially harmful
of the fishes of the Asian region in general is so fishery technologies (e.g. small-meshed gill nets).
poor that many extinctions may never be known. Moreover, the taxa under greatest pressure need
The only assessment of the effects of exotics in not be (and seldom are) the target species of the
the less developed countries of Asia have fishery. For example, Sri Lanka's freshwater
appeared in fisheries literature, and it is not sur- turtles Lissemys punctata (Trionychidae) and Mel-
prising that these generally favor the exotics, anochelys trijuga (Testudinidae) have almost dis-
which have undoubtedly contributed to an in- appeared from dry zone reservoirs, in which they
crease in production. The deleterious effects of were previously abundant, evidently as a result of
exotics have not been examined on the basis of the gill net fishery (pers. obs.). Assessment of the
comprehensive surveys but have usually been effects of individual pressure factors on fish popu-
based mainly on fisheries catch data (e.g. De lations is not always simple, especially when sev-
Silva, 1989). Such surveys often mention only eral factors exist. The presence of exotics is usu-
those species captured and omit or ignore those ally only one of a number of pressure factors on
indigenous species previously present but now native fishes and population declines could be the
not captured or rarely recorded. In a regional result also of other factors, the effects of which
assessment, De Silva (1989: 141) considered that are not easily evaluated separately (Moyle et al.,
O. mossambicus has not endangered any of the 1986).
indigenous fish species or has not had any Problems arise primarily from fisheries manag-
detrimental/deleterious effects on any waters in ers attempting to fill what they perceive as 'vacant'
India, Malaysia, the Philippines, Sri Lanka, Tai- ecological niches with more or less benign exot-
wan or Thailand, and gave positive responses ics. Many of the trophic, ecological and ethologi-
with regard to these two criteria for all 14 exot- cal labels in use are oversimplified, the 'niches'
ics listed. being artificial, and there clearly are dangers in
196
using these labels to justify introductions (see also and more colorful specimens are selected, and
Ribbink & Eccles in Leveque et al., 1988: 296-7). fishing becomes very difficult during the rains,
Many species are also capable of utilizing differ- giving the popUlation time to recover (the rains
ent niches in different localities. also coincide with the breeding period of many
While there is no lack of native species inhab- species). At the present time, pressure from the
iting the reservoirs, what is lacking are lacustrine aquarium fishery is controllable by voluntary re-
fishes of size comparable to tilapia and suscep- straint on the part of collectors and exporters, as
tible to the gill net fishery. Of the larger indigenous is evidenced by attractive but rare fishes such as
secondary consumer (invertivorous and piscivo- P. srilankensis, P. bandula or D. pathirana, not
rous) fishes, Anguilla spp., Wallago attu and being commercially collected despite the exist-
Channa marulius are excluded from the commer- ence of an obvious market (pers. obs.). There
cial fishery by their low frequency and elusive appears to be no evidence to support the state-
habits (they are best captured by angling). Except ment made by Hoffmann (1990) that 'Endemic
for being expensive to feed, Channa striata would forms of sweet water fishes have suffered greatly
appear to be the best candidate for a food fish- from collection as aquarium fish for export; it is
ery. In reservoirs however, C. striata is not a pe- claimed that as a result some species have be-
lagic species, preferring shallow (1 m), marshy, come nearly extinct'.
marginal waters, and is therefore excluded from The possibility of indigenous fish species being
the main reservoir fishery. accidentally translocated with fish fry cultured for
With the exception of Etroplus suratensis which the fishery has been insufficiently examined in Sri
normally feeds on macrophytes but is capable of Lanka (Pethiyagoda, 1991: 36), although evidence
subsisting also on molluscs, all the other Sri Lan- of such incidents have been documented in other
kan primary consumer fishes are cyprinids. With countries (e.g. Armantrout, 1981).
the exception of L. fisheri, a very rare species in-
habiting montane rapids, the largest of these fishes
( < 25 cm standard length) are T. khudree, L. dus- Hydrological alterations
sumieri and L. porcellus. Tor khudree is primarily
a fluviatile species, but larger specimens tend to Diversions and impoundments of rivers are
descend to the floodplain reservoirs (Pertwee, thought not to have seriously affected any fishes
1913; pers. obs.). It is, however, now very rarely as yet. With the exception of the Mahaweli River
found in standing waters. Labeo dussumieri is es- Project, most diversion and impoundment
sentially a benthic detrivore, and well adapted to schemes up to now have affected only the upper
life in rivers, and still has a minor place in the regions of major basins and relatively minor dry
reservoir fishery. Labeo porcellus however, had zone basins. None of the diadromous or vicari-
previously been recorded only from still waters, ous species appears to have been affected by these
and appears to have suffered from direct compe- disturbances as yet. The endemic gobies Sicy-
tition with tilapia spp. A fishery did, however, opterus halei, Sicyopus jonklaasi and Schismatogo-
exist for it until recently (Senanayake, 1980). bius deraniyagalai are restricted to the lower
( < 500 m elevation) regions of the southwestern
wet zone rivers. With the exception of two widely
Exploitation distributed species of Anguilla, significant obliga-
tory intra-basin migration in the Sri Lankan
The only significant fishery for the rain forest freshwater fishes is unknown, although Silva &
fishes in Sri Lanka is by aquarium fish exporters. Davies (1986) observed upstream spawning mi-
I have noted elsewhere (Pethiyagoda, 1991: 32) grations from lowland standing waters in nine
that in comparison with other threats, the threat fish species, none of them considered endangered
from the existing fishery is small: only the large or vulnerable.
197
Conservation damomum), both to State and private agencies,

resulting in severe stresses on habitats (Gurus-
The Department of Wildlife Conservation is re- inghe, 1988).
sponsible for all faunal and most floral conserva- It is noteworthy that while Sri Lanka has a
tion activities in Sri Lanka (legislation is discussed relatively high population density (ca 270 km 2 ),
by Crusz, 1973). Conservation activity is pres- deforestation of the wet zone has had little to do
ently restricted to the enactment oflegislation and with increasing population pressure. These for-
regulations against the exploitation and destruc- ests have been felled or otherwise severely ex-
tion of protected species. Sri Lanka's nature re- ploited, largely by or under the patronage of the
serves were originally established as game re- State, for short-term economic gains.
serves during the early part of this century. Large Despite being committed officially to a progres-
game is often more abundant and can no doubt sive national conservation strategy (IVCN, 1991)
be shot with greater ease in the dry zone scrub Sri Lanka's present conservation programme is
than in rain forest. A curious situation therefore orientated towards awarding protection to a list
exists where 96.4 % of the protected areas of Sri of taxa which are themselves resident largely in
Lanka are in the dry zone, only 3.6 % being in the unprotected habitats. The State's conservation
wet zone (Green, 1990: 195). A much-quoted sta- effort is targeted almost entirely at the larger 'zoo'
tistic (Green, 1990: 195) is that 12.2 % of the land mammals: elephant, bear, leopard and deer. El-
area of Sri Lanka receives some measure of pro- ephants however, presumably for the symbolic
tection (i.e. not used for agriculture, cf. 3.7% glo- and cultural reasons, have become the 'flagship'
bal average estimated by IVCN, 1990). This ex- species and receive the greatest emphasis, and
ceeds, by proportion, the extent of protected areas therefore a grossly disproportionate share of the
in neighboring countries, but it is ironic that hardly resources available for conservation. Meaningful
any of the endemic taxa benefit from this. Erdelen conservation can take place only by eliminating
(1989) showed that the lowland rain forests of Sri this 'Bambi syndrome' from the conservation
Lanka have the highest incidence of endemic ver- strategy and moving away from the concept of
tebrates - and the dry zone the lowest incidence. protected species lists toward more objective, ho-
No endemic vertebrates are restricted to the dry listic approach to conservation (i.e. the conser-
zone. It is relevant that only 74940 ha. (3.3 % of vation of ecosystems, habitats and communities
the wet zone's 22500 km 2 ) is now forested, less rather than individual taxa). It is important that
than half of this being primary forest. the remaining wet zone forests and associated
Such areas as did receive some protection in biotopes be strictly protected, even if it is at the
the wet zone were reserved for timber (e.g. the partial expense of dry zone reserves. It is also
Sinharaja and Kanneliya forests). Much of the important that reserves be designed taking into
Kanneliya forest has been felled by the State account the distribution of the biotas they are
for the manufacture of plywood. Logging in Sin- intended to protect and minimizing the problems
haraja (with ca 47370 ha. of primary rain forest associated with isolating plant and animal popu-
remaining) ceased in 1977. It is estimated that lations (Shafer, 1990).
60% of the flowering plants endemic in Sri Lanka As for the threatened and endangered freshwa-
occur in this forest, with 40 % of these being re- ter fishes of Sri Lanka, given existing trends, no
stricted to it (Zoysa & Raheem, 1990). The means of protection is likely to ensure their
Knuckles Range of hills is another region of spe- survival in the wild in the medium term. The
cial interest. According to currently available dis- strategy most likely to succeed is the maintenance
tribution data, four species offish are restricted to of a sufficiently large number of captive popula-
it. Much of it however, has yet to be surveyed. tions with a view to reintroduction once the pres-
Vast tracts (50%) of this forest have been leased sures on wild populations have been controlled or
out for the cultivation of cardamom (Eletteria car- their survival assured. While this is not an ideal
198
solution, it represents a means of securing the thyofauna can be considered to be in trouble.

survival of at least part of the genetic material. In Overall, the native fishes and fish assemblages
the case of the endangered taxa, the present popu- remaining in Sri Lanka enjoy no protection what-
lations are so small that genetic erosion soever, none of the species with restricted distri-
may already have commenced. Moyle & Williams bution being located in a protected area.
(1990) state that even abundant species can
undergo sudden crashes in their populations over
short periods, as has been seen in Sri Lanka Acknowledgements
in the cases of L. porcellus and M. aral (Pethiya-
god a, op. cit.). The degree of monitoring in coun- I thank Prof. C. H. Fernando, Dr Maurice Kot-
tries such as Sri Lanka is so small that such telat, Dr Peter K. L. Ng, Prof. Mangala de Silva
crashes could easily go undetected; this is also and Dr Eric Wikramanayake for reviewing the
probably the reason for the very small number text and making valuable suggestions for its im-
of extinctions reported from developing coun- provement, and Mr Cedric Martenstyn for assist-
tries. ing with collections. I also acknowledge with
A further possibility is the translocation of gratitude the support given to this project and my
threatened fishes, particularly those with local- work by Mr Asanka Goonewardena.
ized distribution, into less stressed habitats. This
has already been attempted successfully in the
case of four endemic Sri Lankan cyprinids Appendix I. Threatened freshwater fishes of Sri
(Senanayake & Moyle, 1982; Wikramanayake, Lanka.
1990). Several dangers, however, are contingent The terms 'endangered' and 'vulnerable' are used in the sense
on such translocations (Pethiyagoda, 1991: 34- oflUCN (1990: xxiii).
36; Wikramanayake, 1990) which should be un-
dertaken only with a good knowledge of the host Endangered species
biotope and the likely range of dispersion of the Heteropneustes microps (GUnther, 1864). Known from Sri
translocated species. Lanka from only a few specimens. Endemic? (Talwar &
Jhingran, 1991: 690, record the species from India). Habi-
tat: ponds and ditches in a single, heavily man-modified
locality in the south-western wet zone. Range < 10 km 2.
Conclusion Threats: agricultural effluents.
Labeofisheri Jordan & Starks, 1917. Restricted to deep, fast-

The overall conservation status of Sri Lanka's flowing, clear-water mountain streams around the Knuck-
freshwater fish fauna is poor. The wet zone spe- les hills (Mahaweli Basin). Not recorded from un shaded,
cies are threatened mainly by deforestation and silted or turbid waters. Known only from small, discon-
its consequences, while those in the dry zone ap- tinuous populations. Threats: shade loss, siltage. Range
pear to be subject to increasing competition from <500km2.
exotics and fishery pressure. Fishes in both zones Labeo porcellus (Heckel, 1838) s.l. Probably a misidentifica-
are threatened by the increasing use of pesticides. tion and possibly an undescribed endemic (Pethiyagoda,
All nine recently-discovered (last decade) species 1991: 87). Restricted to the dry zone. Population was large
have very small distribution, and are therefore enough to sustain a fishery in 1980 (Senanayake, 1980) but
no specimens were collected in extensive sampling in 1989-
under threat from even relatively minor local dis- 91. Habitat: lowland reservoirs and still waters. Threats:
turbances. Although it is likely that several other competition/predation by exotic spp.? Population declining
fish species await discovery, the rate of increase rapidly.
of pressure on this fauna is so high that extinc-
Lepidocephalichthys jonklaasi (Deraniyagala, 1956). Three
tions are expected. small, widely separated populations known from the south-
In the long term, unless adequate steps are western wet zone. Habitat: shallow, slow-flowing rivulets,
taken to protect aquatic habitats, Sri Lanka's ich- heavily shaded, with leaf debris. Threats: deforestation.
199
Macrognathus aral (Bloch & Schneider, 1801). Formerly a les Range (Mahaweli Basin). Range: 200 km2 • Threats:
common, widely distributed species (Senanayake, 1980: agrochemicals, food fishery, siltage.
319) but not collected since. Habitat: still, muddy water
with vegetation (swampland, rice fields). Range (formerly) Schismatogobius deraniyagalai KotteIat & Pethiyagoda, 1989.
> 25 000 km2 • Threats: agrochemicals and possibly pres- Known from two populations in foot-hill tributaries of the
sure from exotics. Kelani R. Captive spawning achieved (Horsthemke, pers.
comm.), but not reared to maturity. Habitat: unshaded,
Puntius bandula KotteIat & Pethiyagoda, 1991. Known from shallow-water areas of rivers with gravel substrate. Range
one very small population in the Kelani Basin. Habitat: < 50 km2 • Threats: downstream impoundment and
small, rocky stream flowing through rice field and rubber competition/predation by O. mossambicus.
plantation. Captive breeding possible. Range < < 1 km 2 •
Threats: aquarium fishery, agrochemicals. Sicyopterus halei (Day, 1888). Known from four small popu-
lations over a wide range (KeIani-Gin Rivers). Locally very
Puntius srilankensis (Senanayake, 1985). Only one very small restricted. Habitat: unshaded, clear-water, foot-hill rock
population known, from the Mahaweli Basin. Population pools with moderate to fast flow. Threats: down stream
declining rapidly, probably because of siltage of river bed impoundment, siltage.
(Senanayake, 1980: 401; 1985; Pethiyagoda, 1991: 115).
Habitat: shaded, mid-hill clearwaters with pebble/gravel Sicyopus jonklaasi Klausewitz & Henrich, 1986. Known from
substrate. Range < 1 km 2 • Threats: (on going) deforesta- a few small populations from foot-hill streams over a wide
tion and siltage caused by upstream mining. range (KeIani-Nilwala Rivers). Captive breeding possible
(Beyer, 1989). Habitat: clear, unshaded, rocky, sub-
Rasbora wilpita Kottelat & Pethiyagoda, 1991. Two wildely-
montane rapids. Threats: downstream impoundment.
separated populations known from a range < 1 km2, both
in the south-western wet zone. Captive breeding possible.
Habitat: shallow, slow-flowing rivulets, heavily shaded, with
leaf debris. Range (each population) < 1 km 2 • Threats: (on
going) deforestation.
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Diversity and conservation of blackwater fishes in Peninsular Malaysia,

particularly in the North Selangor peat swamp forest
Peter K. L. Ng, J. B. Tay & Kelvin K. P. Lim

Key words: blackwater fish, Peninsular Malaysia, stenotype, rare taxa, ecology, conservation
Abstract
One of the most extreme freshwater habitats in Peninsular Malaysia is the peat swamp forest, with
dark-coloured and highly acidic waters. Surprisingly, little is known about blackwater fishes in Penin-
sular Malaysia. Until 1968, only 26 fish species were known from blackwaters throughout Peninsular
Malaysia, of which only one can be regarded as stenotopic. A recent intensive survey of part of the North
Selangor peat swamp forest yielded 47 species, of which 14 are probably stenotopic taxa. These include
four undescribed species and several new records for western Peninsular Malaysia. These discoveries
are significant in that they include the family Chaudhuriidae which until 1985, was not reported from
Sundaic Southeast Asia, and the rare genus Encheloclarias which had not been encountered for over 50
years. The rapid rate of destruction of the peat swamp forest owing to development, forestry and agri-
cultural activities must be halted or slowed significantly to enable the proper zoological surveys and
studies to be conducted. Conservation plans and environmental impact assessments based on inadequate
sampling and knowledge of species present is acutely dangerous. There are no longer substantial un-
disturbed blackwater peat swamp forests left in most of Peninsular Malaysia. Conservation of the re-
maining blackwater biotopes is critically important if extinction of many species, here regarded as
economically valuable renewable resources, is to be prevented.
Introduction Wyatt-Smith (1959, 1964), Johnson (1967a,

1968), Anderson (1983) and Whitmore (1984,
Peat swamp forests are one of the most threat- 1988) have been converted to padi, pineapple or
ened, yet least studied and most poorly under- oil palm plantations, or developed for some other
stood biotopes in Peninsular Malaysia. One of use. The blackwater swamp forests in Johor, pre-
the most extreme peat swamp water habitat is viously the most extensive in Peninsular Malay-
what is termed 'blackwaters'. Although Johnson sia, are nearly gone. What remains of the black-
(1967a, b, 1968) recognised several kinds of water peat swamps of any substantial size in
blackwaters in Peninsular Malaysia, the most ex- Peninsular Malaysia appear to be restricted to
treme in conditions are those which have dark, northern Selangor, central Terengganu and parts
tea-coloured waters and low pH. of Pahang. In Johor, only pockets of the old peat
Many of the blackwater peat swamp areas in swamps remain. There is thus at present, an
Singapore and Peninsular Malaysia listed by urgent need to conduct more studies on the
204
remaining blackwater peat swamp forests before level and high acidity of blackwaters, which may
they are also lost through reclamation for agri- have given rise to the misconception that such
cultural development. biotopes are 'inhospitable', and thus sustain a
The freshwater fishes that inhabit blackwaters poor faunal diversity. The results of Johnson
are poorly known. There have been few direct (1967a, b, 1968) show the presence of only 26
studies on blackwater species. Studies on acid species in blackwaters, of which only one is a
blackwater fishes were made by Johnson (1967b, stenotopic blackwater taxon.
1968), who compared species composition with Here, we review what is known about black-
that of other freshwater habitats. He (1968) com- water fishes in Peninsular Malaysia, and elabo-
mented that blackwaters were generally species- rate on the importance of this knowledge for fish
poor in fishes. Even less is mentioned in the re- conservation. The possible reasons for previous
cent book on Malaysian fishes by Mohsin & studies having underestimated the fish diversity
Ambak (1983). Mizuno & Furtado (1982) re- and resource are discussed. We draw mainly from
ported on an extensive ecological study of fishes our experiences with the North Selangor peat
in the swamp-lake ofTasik Bera in Pahang, which swamp forest (Fig. 1) and the remaining patches
includes peat swamp forests with blackwaters (see in Johor.
also Lim et al., 1982; Shiraishi et aI., 1972) but
did not specify which species actually line in
blackwaters (see discussion later). In recent years, The chemistry and definition of blackwaters
several anabantoids have been described from or
near Malaysian blackwaters by aquarists - Betta Johnson (1967a, b, 1968) defined blackwater as
tussyae Schaller, 1985, Betta waseri Krummen- waters originating from peat swamp forests,
acher, 1986, Betta persephone Schaller, 1986, highly acidic with pH ranging from 3.6 to 5.9,
Parosphromenus nagyi Schaller, 1985, and Paros- tea-coloured when seen against transmitted light,
phromenus harveyi Brown, 1987. Several new and black when seen en masse via reflected light.
records have also been reported. Vierke (1988) Johnson (1967b) recognised three types of black-
mentioned Betta coccina from near acid waters waters: Gelam (i.e. Melaleuca-associated), dilute
near Batu Arang, Rawang, while Zakaria-Ismail and polluted. The blackwaters discussed in the
(1991) and Ng & Lim (1991) recorded Mystus present study generally follow Johnson's (1967a,
bimaculatus (Bagridae) and Channa bankanensis 1968) chemical definition, but the type is not one
(Channidae), respectively, from or near blackwa- of the three he recognised. A fourth type should
ters of the North Selangor peat swamp forest. be recognised - concentrated blackwaters. This
There has only been one concerted effort in is the habitat on which many of our studies have
Malaysia in recent years to study blackwater spe- been based.
cies, and that is the survey by Davies & Abdullah Johnson (1967a, 1968) noted that blackwaters
(1989) of the North Selangor peat swamp forest. in Peninsular Malaysia are rich in sulphate as
They obtained 16 species from blackwater sites. compared to those in Amazonia, which are rich
In four repeated surveys of the same forest over in chloride (see also Junk & Furch, 1985). It is the
a period of 12 months, the authors obtained 47 presence of high concentrations of sulphates that
blackwater species, the highest number of black- probably accounts for the very low pH. Peat de-
water taxa recorded so far from Peninsular Ma- rived from plant detritus is found at blackwater
laysia. sites and is usually about 0.5 metres thick (see
Considering that the fish fauna of Peninsular Whitmore, 1984, 1988).
Malaysia is relatively well studied compared to There have been no detailed limnological stud-
neighbouring areas, this lapsus in sampling and ies on blackwaters since Johnson's papers, al-
knowledge is surprising. This might well be due to though the work of Lim et al. (1982) for Tasik
the extreme dark colour, low dissolved oxygen Bera (which includes peat swamps) should be
205
t
N
Fig. I. Map of Peninsular Malaysia, showing the North Selangor peat swamp forest (blackened area).
206
consulted. Johnson's work was based primarily necessarily by pH alone. Categorisation of the
on the blackwater swamp forests of southern Ma- various species is based partly on the scarce pub-
laysia (mainly Johor). Although the physico- lished literature and more on our collecting expe-
chemical characteristics of the blackwaters in the riences with these fishes.
swamps of north Selangor (being on the west Habitat preferences of some species, especially
coast) are probably very similar to those of Johor, the rarer or newly discovered ones, are probably
no detailed studies have been done to validate not very accurate. Some species however, nota-
this. Soil conditions on the east coasts of Malay- bly Channa bankanensis, appear to be strict,
sia are different, and some differences in the stenotopic acid blackwater taxon. We obtained
physico-chemical characteristics of the blackwa- large numbers of this species from Selangor and
ters can be expected, e.g. those from Terengganu. Johor, but only from blackwaters; it is not known
in the nearby peripheral habitats. Species such as
Sphaerichthys osphromenoides, have been found in
Classification offish habitat preferences Malaysian acid water habitats that are not strictly
blackwaters. They seem to grow to much larger
The division between blackwaters and usual for- sizes, and have the largest populations in black-
est acid waters (fide Johnson, 1967a) is not al- waters. This is true not only in the North Se-
ways clear, and both often gradually blend into langor peat swamps but also in Johor. Also,
each other. There has been no proper study of the Sphaerichthys osphromenoides has been collected
limnological and physico-chemical aspects of this predominantly (if not entirely) from blackwater
demarcation. Johnson (1967a, b) for example, habitats in Borneo (see later; M. Kottelat, pers.
recognises 'dilute blackwaters' which are brown- comm.) Betta livida, and Parosphromenus harveyi
coloured and higher in pH. Brown-tinted acid are rare in the acid waters of the Rawang area
waters are still common in many parts of Malay- (where the swamp forests have been severely dis-
sia, even in plantations. Many acid water species turned and destroyed), and extremely abundant
seem quite capable of coping with blackwaters. In in blackwaters. We prefer to classify these three
the present study, three tentative categories of species as stenotopic acid blackwater taxa. Many
species are recognised - eurytopic, stenotopic authors also closely associate Luciocephalus pul-
acid water, and stenotopic acid blackwater. Eu- cher with highly acidic blackwater streams and
rytopic species are those which are often common swamps. While it is true that they are quite com-
and thrive in slightly acidic to neutral waters or mon in blackwaters, substantial populations and
even slightly alkaline, as well as acid waters, being large breeding individuals also occur frequently
able to thrive in open-country as well as forest- outside blackwaters in forested acid water habi-
country habitats (fide Johnson, 1967b, 1973). tats (fide Johnson, 1967a; pers. obs.).
Stenotopic acid water species are usually three- The present categorisation tentative though it
country species, and occur only in acidic waters is, permits approximate comparisons and useful
(pH usually less than 6), although many are prob- discussions to be made of Malaysian blackwater
ably also able to thrive in blackwaters. Stenotopic fishes, especially with regards to the stenotopic
acid blackwater species are those that appear to acid blackwater fauna.
be confined to the high acidity, dark-coloured
blackwaters, usually in or adjacent to forest- Blackwater fishes in Peninsular Malaysia
country habitats.
We emphasise that this categorisation is ten- Other than the papers by Johnson (1967b, 1968)
tative. It is possible that the taxa classified here and Davies & Abdullah (1989) who dealt with
as stenotopic blackwater species might perhaps blackwater fishes from Johor and Selangor re-
be stenotopic to a special food item in blackwa- spectively, there have been no other papers which
ters, a particular substrate present etc., and not emphasise the blackwater fauna. Of the 47 spe-
207
cies recorded at present from the North Selangor noted that 11 species could be classified under his
swamp forest, Moshin & Ambak (1983) listed 38 'dilute blackwaters habitat' category. Of the 95
species although many of the species were not species reported from Tasik Bera by Mizuno &
collected by them. They collected only one of the Furtado (1982), only two, Sphaerichthys osph-
stenotopic blackwater species (Sphaerichthys os- romenoides and Ompok leiacanthus, can be re-
phromenoides); most of the other species were garded as stenotopic acid blackwater species.
classified as 'rare or extinct'. Of interest is the There were true blackwater areas in the area but
Puchong site in Selangor sampled by Moshin & these were not sampled or fully explored, and
Ambak (1983) where they obtained Sphaerichthys most of the sampling had been done in tree-
osphromenoides and Parosphromenus deissneri country acid water habitats with pale tea-coloured
(Bleeker, 1859). This area is a disturbed swamp waters (R. P. Lim, pers. comm.).
forest, a remnant of larger swamps that were The extensive freshwater peat swamps around
present in the area (S. Lim, pers. comm.). Kuantan, Pahang, have been poorly explored. The
The information on the peat swamp fish diver- recent discoveries of acid blackwater species from
sity of Terengganu is based primarily on the col- peat swamps in Pahang (Betta tussyae, Betta
lections of Tweedie (1952) from the Merchang waseri, Parosphromenus nagyi) (Schaller, 1985a, b;
and Kuala Brang areas. He noted that the habi- Krummenacher, 1986) reflect this neglect. In a
tat had acidic blackwaters and he described the recent survey of the swamps around Kuantan, we
blackwater Parosphromenus paludicola, and re- have also obtained other stenotopic acid black-
corded Rasbora kalochroma. Seven acid water water species not previously known from the
species were also listed Luciocephalus pulcher, swamps there: Channa bankanensis, Sphaerichthys
Puntius hexazona, Rasbora cephalotaenia, R. osphromenoides and Macrognathus circumcinctus.
pauciperj'orata, R. gracilis, R. dorsiocellata, Nandus Surveys of the blackwater fish fauna in south-
nebulosus and Kryptopterus macrocephalus. Wee western Johor have also revealed several new
B. T. (unpublished data) regarded Parosph- species and records (see Schaller, 1986; Lee &
romenus paludicola (incorrectly as P. deissneri) as Ng, in press).
an acid water species in her survey of the Tereng- Studies on the fish fauna of Taman Negara and
ganu waters and swamps. Aquatic life of large Endau-Rompin, have included little mention of
peat swamp forests in central Terengganu is, fishes from blackwaters (see Zakaria-Ismail,
however, still poorly explored. 1984, 1987; Lim et al., 1990a, b). In his studies of
The important study by Mizuno & Furtado the Malaysian freshwater fishes, Zakaria-Ismail
(1982) on Tasik Bera in Pahang was based mostly (unpublished data) observed that some of the few
around the more accessible area around Fort Is- endemic Malaysian species (Parosphromenus
kandar. Lim et al. (1982: 10) noted that about nagyi, Parosphromenus paludicola) were confined
two-thirds of Tasik Bera (ca 67%) was swamp to the blackwater habitat, and that the absence of
forest with peat substrate; and three habitat types species like Sphaerichthys osphromenoides, Lucio-
could be distinguished in the study site: Utricu- cephalus pulcher and Parosphromenus deissneri in
laria stands, Lepironia stands, and open water Thailand and Indo-China is correlated with ab-
areas. Although blackwaters were reported as sence of blackwater swamps there. Parosph-
present, the extent of the survey on this habitat romenus paludicola has since been found in south-
was not specified. The presence of large numbers ern Thailand (see Kottelat, 1989).
of submerged macrophytes in the study area (see
Mizumo & Furtado, 1982) suggests that the study The blackwater ichthyofauna of Sundaic Southeast
area had mainly dilute blackwater. Mizuno & Asia
Furtado (1982) did not indicate which species
were collected from blackwaters. They compared The Sarawak ichthyofauna as a whole is rather
their species with those of Johnson (1967b) and poorly sampled, although the blackwater fish
208
fauna there is probably diverse. Possible black- Many anabantoids have also been reported from
water species include Ompok leiacanthus (see acid blackwaters in Borneo: Sphaerichthys sela-
Roberts, 1989: 151), Betta livida and Parosph- tanensis Vierke, 1979, Betta anabatoides, Bleeker,
romenus allani Brown, 1987. Rasbora kalochroma 1850, Bettafoerschi Vierke, 1979, Parosphromenus
and an unidentified Sphaerichthys have also been parvulus Vierke, 1979, Parosphromenus filamento-
recorded (Brown & Brown, 1987). The freshwa- sus Vierke, 1981, and Parosphromenus ornaticauda
ter fish fauna of Sabah is well studied, but al- (see Kottelat, 1991; Linke, 1990).
though there are blackwater areas there (P. K. The blackwater swamps of Sumatra are exten-
Chin, pers. comm.), no specific reference have sive but poorly studied. Many species have been
been made to such habitats (see Inger & Chin, described or reported which are here recognised
1962, 1990). In fact, the colour of the water and as acid blackwater species, although this is rarely
pH were not mentioned in their habitat descrip- mentioned, e.g. Channa bankanensis (Ng & Lim,
tions; and only nine species were obtained that 1990), Mystus bimaculatus (see Zakaria-Ismail,
are here recognised as stenotopic acid water taxa. 1990), Rasbora kalochroma (Brittan, 1954b), and
Interestingly, no acid water anabantoids (Belon- Betta coccina Vierke, 1979 (see Linke, 1990).
tiidae), so prominent in Malaysian blackwaters, There have been almost no studies of black-
were collected. Proper surveys of the Sabahan water areas in southern Thailand. Some of the
blackwater habitats have yet to be made (P. K. stenotopic acid water and blackwater species
Chin, pers. comm.). could conceivably be there.
The important study by Roberts (1989) did not
concentrate on blackwater habitats. He did not
specifically mention blackwaters; although he A case study: the North Selangor peat swamp
listed many areas as having brown or brown- forest
tinted water. Only in two of his collecting sites
(1976-4 and 1976-5, Roberts, 1989: 13) were the Our study sites in the North Selangor peat swamp
waters acidic enough (pH 4.5-5), to possibly be forest (Fig. 1) were blackwater streams and
acidic blackwaters. In one site (1976-5) he noted ditches along the trunk road between Kampung
the waters as appearing black when viewed from Sungai Besar and Tanjung Malim. The vegeta-
above. Other than these two sites, the other acid tion is rather dense with Macaranga and Panda-
water sites had pH values from 5.5 to 6.5. In the nus spp. being dominant. Water depth ranged
two pH sites however, Roberts collected all his from about 30 to 160 cm, in tandem with the
specimens of Sphaerichthys osphromenoides, Pa- fluctuating rainfall. The pH of the water is very
rosphromenus parvulus (= P. ornaticauda), his low, ranging from 3.4 to 3.8. This pH level is
single specimen of Neohomaloptera johorensis, significantly lower than that reported by Johnson
most of his specimens of Puntius rhomboocellatus (1967a, 1968) for most blackwaters, but appears
Koumans, 1950, and all his specimens of Rasbora to be usual for the North Selangor swamp forest.
axelrodi Brittan, 1976 ( = R. kalbarensis Kottelat, Colour of the water varied from a dark tea- to
1991). Roberts (in litt.) commented that he had coffee-coloured. The substrate of the streams are
tried to reach good blackwater habitats in the usually composed of a layer of peat and wood
lower Kapuas but those which were accessible debris of varying thickness.
were mainly spoiled by canals. In a follow-up The North Selangor peat swamp forest was
study by Kottelat in the same area, which in- previously more extensive than it is now, extend-
cluded the sampling of blackwaters, Ompok lei- ing into parts of the Rawang area in the south.
acanthus, Betta rutilans Witte & Kottelat, 1991, Development has sharply reduced the total area
Parosphromenus ornaticauda Kottelat, 1991, and covered. For further details on the swamp forest,
Nagaichthys filipes Kottelat & Lim, 1991, were see Chan (1989), Low & Balamurugan (1989) and
found (Kottelat, 1991; M. Kottelat, pers. comm.). Prentice (1990).
209
A note on the material collected Table 1. List of fishes collected from blackwater streams in
the North Selangor Peat Swamp Fore
The present study by the National University of E - eurytopic (acid to neutral or slightly alkaline)
Singapore of the North Selangor peat swamp sur- S - stenotopic to acid blackwaters
vey covered a time-frame of 12 months (May A - stenotopic to acid waters
1991-June 1992), with eight sampling periods. Family CYPRINIDAE
This time-frame included periods of high water 1. Osteochilus spilurus (Bleeker, 1851) A
(May) as well as lowest water (August). The 2. Puntius johorensis Duncker, 1904 A
specimens obtained are still under study, and sev- 3. Puntius hexazona Weber & de Beaufort, 1922 A
4. Rasbora cephalotaenia (Bleeker, 1852) A
eral reports describing the new species of fish,
5. rasbora dorsiocellata Duncker, 1904 A
other aquatic vertebrates and some invertebrates, 6. rasbora einthovenii (Bleeker, 1851) A
discussing the diversity and ecology of the species 7. Rasbora gracilis Kottelat, 1991 A
are in preparation. Reference specimens have 8. Rasbora kalochroma (Bleeker, 1851) S
been deposited in the Institute of Advanced Stud- 9. Rasbora maculata Duncker, 1904 A
10. Rasbora paucipeiforata Weber & de Beaufort, A
ies (University of Malaya); Department of Zool-
1916
ogy, University of Malaya; and the Zoological
Reference Collection of the National University Family BALITORIDAE
11. Neochomalopterajohorensis (Herre, 1944) S
of Singapore. Readings of pH were made with
electronic pH meters, and the glass electrodes Family COBITIDAE
12. Lepidocephalichthys tomaculum Kottelat & Lim, A
were standardised with buffers before each set of 1992
readings. Approximate corroborative readings 13. Lepidocephalichthys pristes Roberts, 1989 A
were obtained with universal pH paper.
Family BAGRIDAE
14. Leiocassis micropogon (Bleeker, 1852) A
15. Mystus bimaculatus Volz, 1904 S
Some taxonomic comments 16. Mystus nemurus (Valenciennes, 1839) E
Family PARAKYSIDAE
Forty-seven freshwater fish species, representing 17. Parakysis verrucosa Herre, 1940 A
18 families, were obtained from blackwater Family SILURIDAE
streams in the present series of investigation in 18. Kryptopterus macrocephalus (Bleeker, 1858) A
the North Selangor Peat Swamp Forest (Table 1). 19. Ompok leiacanthus (Bleeker, 1853) S (A?)
There had been an unfortunate confusion in the 20. Silurichthys hasseltii Bleeker, 1858 A
identities of the cyprinids known as the Six- Family CLARIIDAE
banded and Striped Barbs, presently known as 21. Clarias meladerma Bleeker, 1847 E
Puntius johorensis Dunker, 1904, and P. eugram- 22. Ciarias cf. nieuhofii Valenciennes, 1840 A
23. Clarias teijsmanni Bleeker, 1857 A
mus Silas, 1956, respectively. Kottelat (1992)
24. Encheloclarias curtisoma, Ng & Lim, 1993 S
showed that the type of P. johorensis is actually
identical with the species now known as P. eu- Family HEMIRAMPHIDAE
25. Hemirhamphodon pogonognathus (Bleeker, 1853) A
grammus, and as P. johorensis is an earlier name,
it has priority over P. eugrammus. This is unfor- Family SYNBRANCHIDAE
26. Monopterus albus (Zuiew, 1793) E
tunate as Puntius eugrammus had been a replace-
ment name for P.fasciatus (which the Malayan Family CHAUDHURIIDAE
fishes had been called for a long period), a name 27. New genus, new species S
already preoccupied for an Indian species (see Family MASTACEMBELIDAE
Roberts, 1989). That means the name of the Six- 28. Macrognthus circumcinctus (Hora, 1924) S (A?)
banded Barb, presently incorrectly known as Family PRISTOLEPIDAE
'P.johorensis', should be called P. hexazona 29. Pristolepis grootii (Bleeker, 1876) A
Weber & de Beaufort, 1922, instead. Puntius
210
Table 1. (Continued) mens from Sambas, Banka and Johor reported

upon by Bleeker (1858, 1862) and Herre & Myers
Family NANDIDAE
30. Nandus nebulosus (Gray, 1835) A (respectively) are not con specific (Ng & Lim,
1993). Lepidocephalichthys pristes Roberts, 1989,
Family ANABANTIDAE
31. Anabas testudineus (Bloch, 1792) E
known previously only from western Kalimantan
(Roberts, 1989), Kalimantan Tengah, Kaliman-
Family BELONTIIDAE
tan Timur and Sarawak (M. Kottelat, pers.
32. Belontia hasselti (Cuvier, 1831) A (S?)
33. Betta bellica Sauvage, 1884 A
comm.), is now reported from Peninsular Malay-
34. Betta livida Ng & Kottelat, 1992 S sia. Betta, new species, belongs to a species-group
35. Betta, new species S known previously from Pahang and Sumatra
36. Parosphromenus harveyi Brown, 1987 S (Krummenacher, 1986; Kottelat& Whitten, 1993;
37. Sphaerichthys osphromenoides Canestrini, 1860 S
Ng & Kottelat, in press). The present specimens
38. Trichogaster leerii (Bleeker, 1852) A (S?)
39. Trichogaster pectoralis (Regan, 1910) E
differ from B. waseri s. str. in head shape and
40. Trichogaster trichopterus (Pallas, 1770) E markings. The blackwater balitorid loach, Neo-
41. TRichopsis vittata (Cuvier, 1831) E homaloptera johorensis and akysid catfish Paraky-
Family HELOSTOMATIDAE sis verrucosa has not been previously recorded as
42. Helostoma temminckii Cuvier, 1831 A far north (see Alfred, 1969; Roberts, 1989).
Family LUCIOCEPHALIDAE Rasbora kalochroma, previously known from only
43. Luciocephalus pulcher (Gray, 1830) A a few specimens from Johor and Terengganu
Family CHANNIDAE
(Brittan, 1954a; Tweedie, 1952; Zakaria-Ismail,
44. Channa bankanensis (Bleeker, 1852) S 1987), is now reported from northern Selangor.
45. Channa gachua Hamilton, 1822 A The range of Macrognathus circumcinctus, previ-
46. Channa lucius (Cuvier, 1831) A ously known only from Terengganu and Kelantan
47. Channa melasoma (Bleeker, 1851) A (Sufi, 1956; Cramphorn, 1983; Zakaria-Ismail,
unpublished data), is also extended to the west
coast of Peninsular Malaysia.
The identity of the poorly described Parosph-
hexazona had been previously regarded as a junior romenus harveyi (fide Brown, 1987) has caused
synonym of 'P. johorensis'. some taxonomic problems in that the original de-
Four species have been identified as new. They scription was far too brief and imprecise, with no
are one new cobitid loach of the genus Lepi- type designations, indication of number of speci-
docephalichthys Bleeker, 1863 (L. tomaculum) mens examined or site of specimen deposition.
(Kottelat & Lim, 1992), an undescribed genus Although P. harveyi is similar to P. nagyi, both are
and species of chaudhuriid eel (Kottelat & Lim, certainly valid species, despite the poor original
in prep.), a clariid catfish of the genus Enchelo- descriptions of both taxa. We have obtained
clarias Herre & Myers, 1937 (see Ng & Lim, topotypic material of P. harveyi from near Batu
1993), and a new species of Betta (B. livida) (Ng Arang, near Rawang, Selangor, and it agrees with
& Kottelat, 1992). The family Chaudhuriidae is the excellent series we now have from northern
recorded for the first time from Selangor, and the Selangor. Topotypic specimens of P. nagyi have
family, until 1985, was not known from Sundaic also been obtained and compared directly with
Southeast Asia (see Kottelat, 1985; Kottelat & P. harveyi. Parosphromenus harveyi, differs from
Lim, in Kottelat, 1991). The genus Encheloclarias P. nagyi distinctly in colour patterns and length of
in Peninsular Malaysia, was previously known the pelvic fin (shorter in P. nagyi) (see also
from a single specimen (identified as E. tapeinop- Schaller, 1985a; Linke, 1990). Redescriptions of
terus (Bleeker, 1852) collected from southeastern P. nagyi and P. harveyi are being prepared.
Johor. The genus Encheloclarias, until the present
survey, was believed to be monotypic. The speci-
211
Discussion uted mainly to two factors - collecting methods

and time of collection.
Diversity of Malaysian blackwater ichthyofauna Our collecting gear, rectangular hand-held push
nets measuring 60 by 45 cm with fine mesh, is
Of the 47 species recorded from the North Se- especially effective in the blackwaters as it is
langor blackwater peat swamp, nine are consid- highly portable, can be operated by one person
ered to be eurytopic, 25 stenotopic acid water and and will work even in shallow or narrow streams.
13 stenotopic acid blackwater taxa. The species Two-man teams however, are usually employed,
recognised as stenotopic acid blackwater taxa are with one holding the net and the other disturbing
Neohomaloptera johorensis, Rasbora kalochroma, the submerged vegetation, mud and detritus to
Channa bankanensis, Sphaerichthys osph- drive the animals into the net. The same spot may
romenoides, Betta aff. waseri, Betta livida, Parosph- be sampled several times, even after a short in-
romenus harveyi, Encheloclarias, new species, terval. The disturbed substrate tends to uncover
Ompok leiacanthus, Mystus bimaculatus, new small organisms, thus attracting predatory spe-
genus and species of chaudhuriid and Macrog- cies (e.g. Silurichthys, Clarias, Mystus, Macrog-
nathus circumcinctus. nathus and Channa). Choice of mesh size is criti-
Our list contrasts strongly with Johnson's cal. We used mesh size of about 5 mm, and even
(1968) report that Malaysian blackwaters seem to then, this was barely small enough to capture
be especially depauperate in fish species. Johnson species such as the new genus and species of
(1967b, 1968) commented that he had collected chaudhuriid eel.
only 26 species (only a partial list of species pro- The cast net proved difficult to use for two
vided) from blackwaters in Malaysia, of which reasons. First, the mesh size is too large to trap
only 15 are frequently found. Of these, only small pelagic species such as Rasbora maculata or
Sphaerichthys osphromenoides is a stenotopic acid R. gracilis. Second, the dark water tends to con-
blackwater species. Of the 15 'blackwater ceal the presence of submerged branches. Nets
frequents', he regarded three as typical of more that become entangled with these objects are often
polluted blackwaters: Trichogaster trichopterus, very difficult to extricate. The same is true for grill
Channa striata and Dermogenys pusillus. Three nets. Electrofishing was not attempted as the
were associated with gelam blackwaters (Anabas stunned fish that sink to the bottom are difficult
testudineus, Betta imbellis (as B. splendens) and to recover due to the dark colour of the water.
Rasbora maculata), eight from typical blackwaters Fishing with hook and line is effective, but tends
(Puntius hexazona (as P. pentazona), Puntius jo- to sample only the larger predatory species e.g.
horensis (as P. lineatus), Rasbora pauciperjorata, Channa or Clarias. Local fishermen use wicker
Rasbora cephalotaenia, Hemirhamphodon pogo- traps baited with oil palm nuts. Large species
noganthus, Sphaerichthys osphromenoides, Betta cf. such as Belontia, Pristolepis and Clarias were ob-
pugnax (as B. picta) and Channa lucius (as an served in such traps. Use of these traps will prob-
Ophicephalus). Only one was common in most ably be very effective for larger species, especially
kinds of blackwater, Rasbora einthovenii. if the traps are properly baited with strong-
Davies & Abdullah (1989) recorded 16 species smelling food. The collecting methods used in our
(in six families) from the blackwaters of the North surveys differ from those used by earlier workers
Selangor swamp forest, using mainly cast nets, (see Johnson, 1967b; Davies & Abdullah, 1989)
gill nets and baited lines. Not surprisingly, only in that they are more 'active' than 'passive'.
larger species were collected. Of their 16 species, As most of the water input in the swamp for-
only three were stenotopic acid blackwater taxa. est is by precipitation, the amount of rainfall thus
In our surveys fishes as small and hair-like as the largely determines the water level in the streams.
new genus and species of chaudhuriid eel were The level of water in the streams in August 1991
collected. The differences observed can be attrib- was found to be about half that measured in June
212
the same year. Some fish species that were abun- actual swamp forests (e.g. Betta bellica, Betta, new
dant in June were caught in smaller numbers in species waseri, Betta livida, and Parosphromenus
August, e.g. Clarias spp., Rasbora kalochroma, harveyi), suggesting that type localities of some of
Ompok leiacanthus and Betta livida. On the other these species are not the primary habitat. Like
hand, some species, such as Lepidocephalichthys Vierke (1987, 1988), we obtained B. bellica from
pristes, uncommon in June, were very abundant in dark-coloured acid ditch waters in disturbed for-
August and September. The same appears to be ests near Ayer Hitam, but in North Selangor the
true of Parosphromenus harveyi, with many of the species is much more common and the specimens
August/September males in full breeding colou- obtained are generally larger in size, suggesting
ration, suggesting that there is probably a higher that its preferred habitat is blackwaters.
incidence of breeding in this and probably other A significant proportion of the blackwater spe-
species during low water. Lepidocephalichthys cies are anabantoids. Thirteen species, of which
pristes not only becomes abundant in August, it four are eurytopic, five are acid water stenotopic
attains larger sizes. Our studies thus show that a and four are blackwater stenotopic. The belonti-
particular species recorded as 'rare' at a particu- ids are the best represented, with 10 species, i.e.
lar time, may be abundant at other times. 23 % of the total blackwater ichthyofauna.
The high species diversity in the blackwaters of Johnson (1967b) had noticed the same trends in
the North Selangor swamps should not be sur- his studies; of the 15 blackwater species he con-
prising. Even in the Amazon, a much more in- sidered important, nine (60%) were air breathers
tensely studied system, blackwaters (e.g. the huge or lived near the water surface. Our results are
blackwater river Rio Negro) have species-diver- similar. For the 47 fishes obtained in the present
sities comparable to that in clearer waters (Goul- study, 31 species are air breathers (22) or often
ding, 1985). Johnson (1967b) noted that the swim near the surface (9). This represents some
blackwater fish fauna he obtained represented 66 % of the total fauna. The ability to breathe
only 10% of the total Peninsular Malaysian fauna. atmospheric air is clearly an asset in the extreme
About 260 species of primary freshwater fishes blackwater habitat.
are known from Peninsular Malaysia at present
(fide Moshin & Ambak, 1983; Kottelat, 1989;
Zakaria-Ismail and authors, unpublished data), Fish biomass in Malaysian blackwaters
and the 47 species we have obtained represents
18% of the total known ichthyofauna. This is Johnson (1968) suggested that blackwaters have
higher than believed previously, and considering a very low productivity, not exceeding five pounds
the small area we have sampled, this high per- per acre (equivalent to 5.6 kg per hectare). This he
centage is even more significant. attributed partly to the absence of primary pro-
Many known Malaysian species previously reduction (e.g. by aquatic macrophytes and phy-
garded as rare or very rare, such as Rasbora toplankton) because of the forest cover and dark-
kalochroma, Neohomaloptera johorensis, Ompok coloured acidic waters. Johnson did not state how
leiacanthus, Sphaerichthys osphromenoides and this figure was obtained but he was probably re-
Macrognatus circumcinctus were collected in large ferring to standing stock or biomass, not produc-
numbers in the North Selangor peat swamp for- tivity which has a temporal parameter. Whitten
est. It is obvious that the apparent rarity is due to et al. (1987) in their review of Sumatran black-
imperfect sampling of their preferred habitat, i.e. water aquatic life, agreed with most of Johnson's
blackwaters. The same is true for known species findings. The allochthonous nutrient input into
like Channa bankanensis, Mystus bimaculatus, blackwaters, though recognised by Johnson, is
Clarias aff. nieuhofii. Species previously obtained probably much more important. Junk & Furch
by aquarists in small numbers from drainage (1985) commented that the great number of fish
ditches are also present in large numbers in the recorded from Amazonian blackwaters was pre-
213
dominantly due to allochthonous inputs - insects, Table 2 List of North Selangor blackwater peat swamp fishes
fruits, pollen etc. collected for consumptJon by local people and of aquanum
Importance
An intensive and thorough collection was made
at one North Selangor site which covered about Food species AquarIUm/ornamental species
50 square metres. The fish field, comprising
Mystus nemurus Rasbora cephalotaema
mamly of the snakehead, Channa bankanensls, MyslUS blmaculatus Rasbora dorSlOcellata
and the spiny eel, Macrognathus circumcinctus, Kryptopterus macrocephalus Rasbora emthovenu
both edible species, totalled about 0.2 kg. This Cianas telJsmanm Rasbora graCilIS
Cianas cf meuhofo Rasbora kahchroma
quantity, equivalent to 40 kg per hectare, is a con- Cianas meladerma Rasbora maculata
servative estimate as smaller species (e.g. Mystus Pnstoiepis grootu Rasbora pauclpeiforata
bimaculatus, Betta, new species etc.) were not Ananbas testudmeus Punttus johorensls
Belontta hasseltl PuntlUS hxazona
considered. Nonetheless, this figure is still con- Tnchogaster leeru Osteochtlus spllurus
siderably higher than that reported by Johnson Trtchogaster pectoralIS Hemtrhamphodon pogonognathus
(1968). The fish biomass of blackwaters is Tnchogaster trlchopterus Mystus b,maculalus
Helostoma temmmckll LelOcasslS mlcropogon
probably less than large clear water drainages. Macrognathus ctrcumcmctus Kryptopterus macrocephalus
Bishop (1973) reported a value of some 180 kg Monopterus albus Stlunchthys hasseltll
per hectare for the Sungei Gombak river system Channa bankanensls M acrognathus ctrcumcmctus
Channa lUCIUS Belonlta hasseltl
in Selangor. Channa melasoma Betta Imda
Betta belhca
Betta, new species
Parosphromenus harveYl
Economic significance of the blackwater Sphaertchthys osphromenoldes
ichthyofauna Tnchogaster leertl
Tnchogaster /Tlchopterus
Helosloma temmmcku
Blackwater peat swamps have been recognised as Luclocephalus pulcher
important catchment areas. The thick peat layers Channa gachua
tend to absorb and store large quantities of water
in the rains, and this water can be tapped for
agricultural use. In North Selangor, the rice fields
are irrigated by water channeled from the peat
swamps (Low & Balamurugan, 1989) as well as The potential for the aquarium trade is very
from the Sungai Bernam via the Karang River. promising. Among the 47 species collected in the
The economic importance of the ichthyofauna present survey, 27 are already or having very good
in Peninsular Malaysian blackwater peat swamps potential to be exploited for the aquarium trade
should not be overlooked or underestimated. Al- (see Table 2). Well known and popular blackwa-
though blackwater swamps are threatened habi- ter fishes already established in the trade include
tats, there is no reason, with foresight and proper the 'Pearl Gouramy' (Trichogaster leerii), 'Choco-
management, why the rich fish resources there late Gouramy' (Sphaerichthys osphromenoides),
cannot be exploited for consumption and the 'Clown Rasbora' (Rasbora kalochroma), 'Six-
aquarium trade for the benefit of the local popu- Banded Barb' (Puntius hexazona), and the 'Two-
lace. spot Catfish' (Mystus bimaculatus). With the gen-
Harvesting of the larger species for consump- eral popularity of anabantoids, there is also a
tion, albeit in a small scale by the local people demand for colourful or exotic species for the
have been observed. The 18 species caught for trade - and species like 'Claret Fighting Fish'
this purpose are listed in Table 2. Most of the (Betta livida), 'Bellicose Betta' (Betta bellica),
species are sold for very low prices and have only 'Giant Fighting Fish' (Betta, new species) and
local importance. None of the species is trans- 'Harvey's Licorice Gouramy' (Parosphromenus
ported far for sale. harveyi) fit the requirements very well.
214
Conservation of the Malaysian blackwater We observed that virtually all the areas sampled
ichthyofauna by us have been logged before. In fact one par-
ticularly rich stream flows through an oil palm
The current literature review and studies, using estate from the swamp forest. The high fish di-
the North Selangor blackwater peat swamp for- versity in these areas suggests that selective and
est as a case study, show that there is great po- controlled harvesting of timber apparently does
tential for research work to be carried out in no real immediate damage to the peat swamp
blackwater peat swamps. Contrary to published forest ecosystem. In fact, the Asian Wetland
literature, blackwater swamps are not species- Bureau has drawn up several proposals in favour
poor and low in biomass. The number of fish of controlled logging in the North Selangor peat
species recorded at present from the North Se- swamp forest (see Chan, 1989; Prentice, 1990).
langor peat swamp forest far exceeds those ob- Similarly, Davies & Abdullah (1989) recommend
tained in previous collections. It shows that the that landuse in the peat swamp should be based
fauna, at least where freshwater fish is concerned on the retainment of forest cover and avoiding
is not poor, and strongly suggests, especially in total land clearance. Within this proposal, proper
the discovery of four new species, that there is management of the habitats and controlled har-
much to learn and discover in this neglected habi- vesting for the aquarium trade will enable the local
tat, which is fast losing ground to human exploi- population to derive long term, direct financial
tation and encroachment. As has already been returns from conserving the main swamp forest
demonstrated in several studies (see Prentice, area.
1990), the conservation and proper management The value of the aquarium trade is not often
of peat swamps is important not only in ensuring obvious as complete records and data are lack-
the survival of a rich and poorly studied, unique ing. In Indonesia for example, many unregistered
ecosystem, but also as a natural resource from foreign collectors organise large scale 'harvests',
which to tap water, timber and fishes for human with the live fishes exported directly back to their
use. countries without passing through local authori-
The value of wild aquarium fish has not been ties. Undoubtedly this also occurs in Malaysia, to
widely discussed in scientific communities other an unknown degree. This has led local authorities
than with concern for endangered species such as to underestimate the value of this resource. To get
Ikan Kelesa or Golden Dragon Fish (Scleropages just a rough idea of the value of the resource at
formosus). The freshwater aquarium fish resource hand, we calculated the profits of selling, for ex-
is a very lucrative trade. In Singapore, this trade ample, 1000 individuals of each of the 25 species
exceeded US$50 million in 1990, and although we have recognised here as worthy of the
much of this is in cultured species, a significant aquarium trade. The individual species costs are
proportion consists of wild-caught fish (Ng, based on what we know of retail prices in Sin-
1991). Johnson et al. (1969) noted some of the gapore. In Europe and America, prices are almost
species they felt were valuable and discussed the certainly much higher. The retail profit amounts
potential of harvesting Malaysian wild fishes for to at least US$12000. While such an exercise is
the aquaria. The aquarium trade however, has purely hypothetical and approximate, it gives a
grown greatly since. Concern about overexploit- rough idea of what income to expect if the re-
ing the fish resources are of course valid (see sources are properly managed. Considering the
Alfred, 1968; Tweedle, 1961; Johnson, 1961; size of the forest, extent of the swamp forest
Johnson et al., 1969). This has led to proposals drainage and biomass of the waters, very large
on banning collections and exports (or imports) numbers of some of the smaller species might be
in some countries keen on conservation, but un- harvested without serious impact on their popu-
fortunately, this idea is too simplistic and does lations, as long as it is properly managed. Some
not really help conservation ultimately. aquarium species such as Sphaerichthys osphro-
215
menoides or Macrognathus circumcinctus, which ours in North Selangor peat swamp forest are
the published literature regards as rare or endan- now no longer possible in Johor, where the most
gered in Malaysia, are in fact, common in acidic extensive peat swamps and blackwater habitats
blackwaters, the species being common enough in in Peninsular Malaysia once occurred. It would
these habitats to sustain controlled exploitation. be pointless to speculate what undiscovered or
It is our belief that retention of the intact forest unrecorded species once occurred in these black-
is more crucial to the survival of these species in waters. A small hint of its vast wasted potential
the wild than an ineffectively conceived but per- can be perhaps gleaned from the discovery of the
haps well meaning total ban. A problem associ- blackwater snakehead Channa bankanensis in the
ated with the current fish trade that must be men- western remnants of the J ohor peat swamp for-
tioned is that many of the specimens collected do ests. It is however, not impossible that some of
not reach their eventual destinations (i.e. aquar- these species (like C. bankanensis) still survive (al-
ists) and even when they do, most do not live very beit rather tenuously) in small and as yet, intact
long. The compromises involved in harvesting the blackwater areas, and await discovery. System-
wild fishes for economic gain and trying to reduce atic surveys should be made in these areas before
the tragic waste of so many lives are many. development completes its coup-de-grace on
Johnson (1968) had commented that the chem- southern Malaysian blackwater habitats.
istry of blackwaters tends to limit productivity,
and indicated that ' ... high fields of fish can be
obtained from ponds built on gelam lands [with Acknowledgements
blackwater] if the water is limed and treated with
phosphate' (p. 309). He realised the conflicting Mr Duncan Parish of the Asian Wetlands Bureau
problems of producing traditional food fishes first broached the idea of a detailed fish survey in
against that of valuable aquarium species, al- the North Selangor peat swamp forest, and
though he felt that blackwaters are poor to mod- throughout this exercise, he has been most help-
erately good for only a few species of aquarium ful. Mr Jonathan Davies and Mr Abdullah Rahim
species. Our study suggests otherwise. kindly shared their observations and experiences
The association between the loss of specific of their first survey with us, and provided many
blackwater habitats and the fate of a species can useful tips. Dr Maurice Kottelat (Munich Mu-
be perhaps best exemplified by Betta persephone, seum) and Dr Tyson Roberts (California Acad-
a beautiful belontiid recently described by emy of Science) were, as always, generously
Schaller (1986). It is known for certain only from forthcoming with their help and advice in identi-
one small stream - its type locality near Ayer fying many of the taxa. Dr Susan Lim (Univer-
Hitam in Johor. This locality has been developed, sity of Malaya) was infectious with her enthusi-
a new highway and other projects have destroyed asm and urging us to make known our findings.
most of the forest in the area. The species can be Discussions with Dr Mohd. Zakaria-Ismail (Uni-
found only in the remnants of the peat swamps in versity of Malaya), Dr Guy Teugels (Tervuren
the Ayer Hitam area. It appears however, that the Museum), Dr Jorg Vierke (Husum, Germany),
species also occurs in several areas south of Ayer Mr Dietrich Schaller (Munich, Germany), Pro-
Hitam, and some specimens from this area have fessor Yong Hoi Sen (University of Malaya), Dr
been imported into Germany for the aquarium Richard P. Lim (Australia) and Datuk Chin Pui
trade (pers. observ.; M. Kottelat, pers. comm.). Kong (Sabah Fisheries) were helpful on taxo-
It can only be hoped that the habitats where nomic and ecological aspects during the collation
B. persephone still occurs, which are almost cer- of this report. We are grateful to Dr Mohd.
tainly vulnerable, can be protected in the near Zakaria-Ismail for so generously permitting us
future. the use of his unpublished thesis. Dr Maurice
It is unfortunate that detailed surveys such as Kottelat, Dr Tyson Roberts, Dr Mohd. Zakaria-
216
Ismail and an anonymous reviewer kindly re- Brown, B., 1987. Special announcement - two new anaban-
toid species. Aquarist & Pondkeeper, June 1987: 34.
viewed the manuscript. Field assistance by the
Brown, A. & B. Brown, 1987. A survey of freshwater fishes
Department of Zoology (National University of of the family Belontiidae in Sarawak. Sarawak Mus. J. 37:
Singapore) Honours class of 1991, Mr Tommy 155-170.
Tan, Ms Diana Chia, Mr Jeffrey Lee, Mr Abdul Cramphorn, J., 1983. Sungai Terengganu fish survey, 1980.
Latiff bin Zainal, Mr N. Sivasothi and Mr Lau Malay. Nat. 3: 16-20.
Chan, H. T., 1989. A Forestry Action Plan for the North
Ching Ong is much appreciated. This study has
Selangor Peat Swamp Forest. Asian Wetland Bureau Re-
been supported by a research grant 910410 to the port No. 46c, IPT Asian Wetland Bureau/WWF Malaysia,
first author from the National University of Sin- Kuala Lumpur, 8 pp.
gapore. Davies, J. & A. R. Abdullah, 1989. Freshwater Fish Survey
of the North Selangor Peat Swamp Forest. Asian Wetland
Bureau Publication No. 46, IPT Asian Wetland Bureau/
WWF Malaysia, Kuala Lumpur, 8 pp.
Note added in proof Goulding, M., 1985. Forest fishes of the Amazon. In G. T.
France & T. E. Lovejoy (eds), Key Environments: Ama-
A new genus and new species of minute cyprinid zonia. Pergamon Press, Oxford: 267-276.
was recently discovered from the North Selangor Inger, R. F. & P. K. Chin, 1962. The fresh-water fishes of
peat swamps and is currently being described by North Borneo. Fieldiana, Zool. 45: 1-268.
Inger R. F. & P. K. Chin, 1990. The fresh-water fishes of
K.-E. Witte. This is a stenotopic blackwater spe- North Borneo. Fieldiana: Zoology, Volume 45 (1962). Re-
cies. The number of fishes now known from the printed by Sabah Zoological Society, Sabah, Malaysia, with
peat swamp totals 48. supplementary chapter by P. K. Chin, 1990. 268 pp.
Johnson, D. S., 1961. Freshwater Life in Malaya - its con-
servation. Malay. Nat. J. special issue 1961: 232-236.
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Red tide phenomena in Brunei Darussalam - some implications for

fisheries
Selvanathan Subramaniam, Siti Amin Mahali & Sabri Hj Mohd Taha

Department of Fisheries, Ministry of Industry & Primary Resources, P.O. Box 2161, Bandar Seri
Begawan 1921, Brunei Darussalam
Abstract
Since the first recorded occurrence in 1976, the coastal waters of Brunei Darussalam have been spo-
radically subjected to the effects of red tide. The 1976 incident overwhelmed the nation which had no
previous experience of red tides. Subsequently, a routine was established to monitor the red tide phe-
nomenon. This included plankton monitoring and shellfish toxicity testing, measures which helped to
prevent or mitigate adverse human impacts and economic losses to the fishing industry.
The second red tide occurrence was in 1980. However, this time there was greater awareness and with
the experience gained from the 1976 incident, the situation was handled effectively.
A subsequent incident in 1988 was discovered in a slightly different manner. High densities of
Pyrodinium bahamense var. compressum, the causative organism, were found in the stomach contents of
a pelagic fish, Sardinella spp. Cats which had fed on contaminated Sardinella suffered sickness and
mortality. This led to a ban on the import and sale of Sardinella spp. and the closure of a local shell-
fish farm for almost a year.
Red tide occurrences have had some impact on the fisheries of Brunei Darussalam, mainly due to the
need to impose bans on the import, sale and consumption of certain species of fish and shellfish.
It is suggested that the effects of red tides on finfish capture fisheries, especially planktivorous fish can
be minimised by implementing simple precautions. Shellfish under culture would invariably face losses
in the event of a prolonged red tide occurrence and it is proposed that an insurance scheme be imple-
mented to cover such losses.
A Red Tide Action Plan is currently in force in Brunei Darussalam. It is a contingency plan for red
tide vigilance, monitoring and management, and will serve to reduce the negative impacts of red tides.
Introduction only 0.14%. Despite this apparent low signifi-

cance, fish and fisheries remain close to the hearts
Brunei Darussalam is situated on the north west of Bruneians. A total of 585 fishermen are en-
of the island of Borneo, between the latitudes 4 ° gaged in fishing on a full-time basis. In addition
and 5°05' N and longitudes 114°04' E and to this, there are 1218 part-time fishermen who
115° 22' E. are employed in the public or private sector but
The mainstay of its economy is an extensive oil conduct fishing operations during their holidays
and gas industry, which in 1988 contributed about or after work hours. The higher numbers involved
60% of the National Gross Domestic Product. In in part-time fishing as opposed to full time fish-
contrast to this, the fisheries sector contributed ing is testimony to the fact that despite the avail-
220
ability of comfortable, land-based jobs, fishing Red tide occurrences in Brunei Darussalam
retains a place of importance for many Brune-
ians. The red tide of 1976
Fish is also an important component in the diet
of Bruneians. It is a highly regarded source of The general areas of red tide blooms and sam-
protein. The per capita consumption of 40 kg/ pling stations are shown in Figs l(a) and (b).
annum is comparable with that of most South The 1976 red tide was discovered almost by
East Asian countries. chance (Beales, 1976). Fisheries Department bi-
Red tides are a relatively new phenomenon in ologists were out at sea on 11 March, 1976 when
Brunei Darussalam, the first recorded incident they spotted on extensive reddish brown disco-
being as recent as 1976 (Beales, 1976). In the louration of the sea surface about 4 nautical miles
neighbouring state of Sabah, red tides were also north-northeast of Muara port. Samples were
unheard of prior to 1976 (Wong & Ting, 1984). It taken and initially identified as a species of
is possible that earlier incidents may have oc- Gonyaulax. This was later verified as being
curred without being noticed but this is unlikely, Pyrodinium bahamense var. compressum.
given the deleterious effects that red tides would Five patients were treated on the same day at
have caused. Seliger (1989) suggests mechanisms the Government Hospital after having had a meal
to explain the development of red tides in Brunei of Chub Mackerels (Rastrelliger sp) and Scads
Darussalam. (Selar sp). They complained of numbness, giddi-
This paper attempts to review the red tide phe- ness, fatigue, a bitter taste in the throat, weakness
nomena and its effects on fisheries in Brunei in the limbs and tightening oflips, tongue, fingers
Darussalam. and toes (Beales, 1976). It was also reported from
I~'I ::: ~::: ::::

0 2 :3 4
I I I I
10 Nau tical miles SABAH
(MALAYSIA)
t
N
Bandar
Ser' ,
Begawan I "-
Tutong ,- , -' ,
/ ' I
BRUNE I /SARAWAK (' Temburonll
OARUSSALAM,l MALAYSIA) I BRUNEI I
, ' ARUSSALAM
Fig. I. (a) Known locations of 1976 and 1980 Pyrodinium red tides in Brunei Darussalam's coastal waters and adjacent areas
(Beales, 1976; laafar & Subramaniam, 1984; laafar et al., 1989).
221
• Sampling for Pyrodlnlum N

o
{
Pelong
Sampling for PSP tox Ins Flock 0 •
• • •
. \:SO Besor
(J P. Bedukang
-.
Sg. Teritip •
Inner Boy
BRUNEI
Bandar
Seri Begawan tJP. Baru Baru
o I 2 :3
/ . <7
P. Kirang Nautical miles
Fig. J. (b) Locations of sampling stations from which high densities of Pyrodinium bahamense var compressum were recorded in
vertical plankton haul samples (Jaafar et af., 1989).
Sabah that four children had died after partaking tide phenomenon and the fish kills. Indeed at
of a meal of bivalve molluscs and that some others Pelong Rock (a coral reef area) where the great-
were taken ill (Roy, 1977). est number of dense blooms occurred there was
These related incidents led the officials to sus- no evidence of fish mortality during or after the
pect a possible occurrence of red tides. Various incident (Beales, 1976).
actions were immediately taken to prevent or
minimise ill effects to the public. These included
a public announcement on the dangers of red The red tide of 1980
tide; the destruction of food organisms suspected
to be contaminated; the monitoring of the plank- After the 1976 red tide incident, various institu-
ton bloom in the affected coastal areas and the tions in Brunei Darussalam, such as the Navy,
bioassay of shellfish and related food organisms. Air Force, Commercial Airlines and the public at
This episode lasted till May, 1976. The red tide large were advised to maintain a vigil for red tide
at its height was reported to stretch for about blooms and to inform the Fisheries Department
200 km, from Kudat in Sabah to the western tip accordingly. In addition, the Fisheries Depart-
of Brunei Darussalam. ment initiated a programme of regular plankton
The 1976 incident was the only one in which monitoring and shellfish bioassays.
fish kills were reported but this was on a rather With public co-operation the sighting of red
limited scale. It was restricted to the deaths of a tide blooms in Brunei Bay on 28 April, 1980 was
few hundred specimens of Lethrinus spp. while reported to the Fisheries Department. The Fish-
Stolephorus spp. were reportedly found washed eries Department officials confirmed that it was a
ashore in the vicinity of Muara Port during the red tide caused by Pyrodinium bahamense var.
initial stages of the outbreak. However, there was compressum. and appropriate warnings were im-
no conclusive evidence of a link between the red mediately issued to the public. This early warning
222
probably prevented the occurrence of any para- Sabah. Following the incident an immediate ban
lytic shellfish poisoning (PSP) incidents in Brunei on import and sale of planktivorous fish was en-
Darussalam. forced.
Whereas in the 1976 incident, the blooms were In mid-January 1988 high densities of Pyrod-
found in surface waters, during the 1980 incident inium cells were detected in plankton samples
dense blooms were found at a depth of about taken off the Brunei coast, subsiding towards the
10 m. The progressive sequence ofthis occurrence end of March, recurring at the end of April and
is described in Jaafar & Subramaniam (1984). lasting up to August 1988 (Jaafar etal., 1989).
After the 1980 incident of red tide, there were Despite the high Pyrodinium numbers, there were
no further incidents of red tide blooms. However, no visible blooms, unlike the earlier incidents.
Brunei Darussalam remained vigilant and was Toxin levels in shellfish (Perna viridis) and a fish
placed on alert for possible red tide effects fol- (Selar kala) on 23 January 1988 were found to be
lowing red tide phenomena in the neighbouring 77 Jlgj100 g and 51 Jlg/100 g of flesh respectively.
state of Sabah in July 1983, December 1985 and The levels in the shellfish increased to 5354 Jlg/
December 1986 (Jaafar et al., 1989). 100 g of flesh in wild stock and 1054 Jlg/I00 g of
flesh in the cultured specimens by 3 February
1988. It has been established that levels above
1987j 1988 Occurrence 80 Jlg/100 g flesh can be considered unsafe for
human consumption.
J aafar (loc. cit) describe in detail the series of In view of this, the earlier ban, which was lim-
events associated with this occurrence. This in- ited to the import and sale of planktivorous fish
cident was detected in a rather different manner was extended to cover molluscan shellfish as well.
from that of the two earlier incidents in 1976 and
1980.
In December 1987, the Government Veterinary Effects on fisheries
Clinic in Bandar Seri Begawan treated 40 cats for
suspected poisoning. Investigations revealed that In the 1976 red tide incident 55578 kg of fish
all these cats had earlier been fed with a clupeoid valued at B$198 000 were confiscated and de-
fish, Sardinella spp. Seven of the cats perished stroyed, on suspicion that they may have been
and many of the others were reported to be in contaminated (Jaafar et al., 1989). These actions
serious condition. As all these cats had consumed were necessary considering the fact that very little
similar material, it was naturally assumed that the information was available on the red tide phe-
fish was the probable cause of the poisoning, es- nomenon, and safeguarding public health was of
pecially so, as Sardinella spp. is known to be a utmost importance. The fact that there were no
plankton feeder. Examination of the stomach human fatalities at that time vindicates the rather
contents of the fish revealed large numbers of severe measures taken.
Pyrodinium bahamense var. compressum. Mortal- The net effect on the commercial fisheries was,
ity following the consumption of fish contami- however, quite significant. The bans imposed on
nated by Pyrodinium is rather unusual, as it is the sale of fish caused the fishermen to suspend
generally unknown for dinoflagellate toxins to operations for a period of time, compelling the
bioaccumulate in the flesh offish. In this instance, provision of welfare assistance to those badly
the poisoning was explained by the fact that the affected (Beales, 1976). Red tide also placed new
fish had been consumed whole, with guts, gill, etc. limitations on the culture of marine molluscs
Ting & Wong (1989) reported a similar poisoning (Jaafar & Subramaniam, 1984).
via Sardinella and Decapterus spp. in Sabah. In the 1980 outbreak of red tide a more sub-
The source of the fish in the Bandar Seri dued approach was taken, possible then because
Begawan cat mortality incident was traced to of the experience gained from the previous inci-
223
dent. A ban was imposed on the collection and tide and PSP to be extended also to countries
sale of molluscs, but other seafood such as fish from which imports offish and shellfish are made.
continued to be freely sold (J aafar, loco cit). Generally, the food organisms that have been
Though there were economic losses to the fish- implicated with red tide and PSP in Brunei
eries sector, these were not recorded. The losses, Darussalam are molluscan shellfish and some
however, would have been lower than those in- planktivorous fish. The production and consump-
curred in 1976 when the ban was imposed on tion patterns offish and shellfish in Brunei Darus-
both fish and molluscs. salam are shown in Table 1. The main planktivo-
The 1988 incident brought more serious effects rous fish considered (viz Rastrelliger spp.,
on commercial fisheries than the 1980 incident. Decapterus spp., Selar spp. and Sardinella spp.)
This is due to a number of factors: comprise a little under 25 % of the total fish con-
sumed, of which more than 50% is imported.
- the existence then of a commercial mussel farm
Molluscan shellfish comprise less than 5 % of the
in Brunei Darussalam;
total fish consumption, and the bulk of this is
- the 'implication' of a planktivorous fish in
imported from the neighbouring states of Sabah
paralytic shellfish poisoning (PSP).
and Sarawak.
The ban on harvesting and sale of shellfish caused Local fishing gears and the number of fisher-
losses of about B$112 000.00 for the local mussel men exploiting the aforesaid planktivorous fish in
farm (Jaafar et ai., 1989). Difficulties were also 1990 are given in Table 2. It is evident that there
encountered by importers of molluscan shellfish are a substantial number of fishermen involved in
as well as by fishermen operating fishing gears fishing for pelagic fish species. Their livelihood
such as purse-seines which capture planktivorous would therefore be affected if bans were imposed
fish such as Sardinella spp. and Rastrelliger on the sale of these fish. However, as was men-
spp. The losses incurred by the fishermen were tioned earlier, this problem has been somewhat
quite considerable immediately following the overcome with advice given to the public on the
announcement that fish had also been 'res-
ponsible' for PSP.
Table 1. Production and consumption offish and shellfish in
Consumer confidence in the safety of fish and
Brunei Darussalam (m. ton).
fish products declined. However, the situation
improved shortly after, when advice was given to Year 1988 1989 1990
the public on the safe preparation of fish cooking
Total consumption (fish, 4743.3 5805.8 6051.8
- consumers were advised to completely remove shrimp, molluscs)
the gill, guts and internal organs of fish before Total consumption of 639.5 1148.6 1158.0
cooking. *p1anktivorous fish
% *planktivorous fish to 13.5 19.8 19.1
total consumption
Local production of 241.2 510.7 495.6
Implications for fisheries *planktivorous fish (%) (37.7%) (44.5%) (42.8%)
Total mollusc consumption 117.3 231.3 278.9
% mollusc to total 2.5 4.0 4.6
It has already been established that fish is an consumption
important component in the diet of Bruneians. Local production of 10.6 14.4 8.3
mollusc (%) (9.0%) (62.3%) (3.6%)
Owing to the early stage of development of com-
mercial fisheries in Brunei Darussalam, the local * Only the following species known to be implicated in red
production is insufficient to meet local demand. tides and PSP are considered:
Rastrelliger brachysoma.
Consequently more than 50% of the fish con-
Rastrelliger kanagurta.
sumed locally is met by imports from neighbour- Decapterus spp.
ing areas. Sardinella spp.
Such a situation necessitates vigilance for red Selar spp.
224
Table 2. Number of fishermen exploiting planktivorous fish in to be affected during a red tide/PSP occurrence.
Brunei Darussalam - 1990.
In this context, it must be stated that evidence
Fish group Fishing gear Fishermen (no.) to date indicates that the toxin does not accumu-
late in the fish flesh (White, 1980). It has also been
F/time P/time reported from USA that the toxin produced by
red tide alga was confined to the liver and other
Rastrelliger spp. Purse-Seine 45*
viscera of the fish. Thus, removal of guts, gills and
Ring net 15 5
Gill net (large mesh) 65 66 all internal organs as advised would render the
Hand Line 83 132 fish flesh edible. It is therefore foreseen that with
Decapterus spp. Purse-Seine 45*
the implementation of this simple precaution,
Ring net 15 5 there will be little problem in harvesting or mar-
Gill net (large mesh) 65 66 keting this group of fishes during red tide occur-
Hand Line 83 132 rences. Indeed, during the 1988 incident there
Selar spp. Purse-Seine 45* were substantial landings of planktivorous fish
Ring net 15 5 following the public announcement as to the safe
Trammel net 252 846 preparation of fish. Thus it seems unlikely that
Gill net (large mesh) 65 66
these fishermen would be unduly affected by new
Hand Line 83 132
Cast net 28 159 red tide/PSP incidents.
With respect to shellfish harvesters in Brunei
Sardinella spp. Purse-Seine 45*
Ring net 15 5
Darussalam, they are small in number. All of them
Large Corral 5 3 operate on an artisanal basis merely to supple-
ment their income. This is fortunate, for as they
* Averaged from 15 fishermen/vessel.
are not dependent on shellfish for their sole
wherewithal, they are able to simply suspend
safe method of preparation of pelagic fish prior to operations during the course of a red tide/psp
cooking. situation. There is no alternative to this for unlike
Molluscan shellfish is mainly collected from the finfish, shellfish are known to accumulate the toxin
wild. Production levels are very low (Table 1). in the flesh.
Production from a commercial mussel culture Another category of persons likely to be
farm is also low at present. affected are shellfish culturists. J aafar & Subra-
Those in the fisheries sector who would there- maniam (1984) reported that early investigations
fore be affected by a red tide or PSP are fish into the culture of Perna viridis in Brunei Darus-
(including shellfish) dealers, fishermen and shell- salam were encouraging. A commercial shellfish
fish culturists. farm was subsequently established in 1985. Whilst
The concern of fish dealers is the general sus- Brunei Darussalam does have many factors con-
picion with which seafood is held during a red ducive to shellfish culture, red tides pose a prob-
tide situation. During the 1976 incident, a high lem. Jaafar et al. (1989) reported that the com-
level of wariness was evident. Subsequent occur- mercial shellfish farm suffered a loss of about
rences showed considerable improvement in this B$112000.00 as a result of the ban on harvesting
regard, thanks to the educative efforts taken by molluscan shellfish during the 1988 red tide inci-
the Fisheries Department to enlighten the public. dent. It is submitted that the situation is not totally
Nevertheless, during each red tide situation, the dismal because with an appropriate insurance
Fisheries Department does receive calls from the coverage as a contingency, the problem of finan-
public enquiring as to whether a particular sea- cial losses due to red tides/psp can be mitigated
food is edible or otherwise. or overcome. With an insurance umbrella in place
Fishermen, especially those exploiting the there is little reason not to proceed with shellfish
planktivorous fish mentioned above are also likely culture in Brunei Darussalam.
225
Conclusion Centre, Singapore and International Development and

Research Centre, Ottawa, Canada: 17-24.
Jaafar, H. M., M. W. R. N. De Silva & P. H. Y. Sharifuddin,
It is evident that red tide phenomena has caused 1989. Pyrodinium red tide occurrences in Brunei Darus-
some degree of negative impact on fisheries in salam. In G. M. Hallegraeff & J. L. Maclean (eds), Biol-
Brunei Darussalam. This is mainly in relation to ogy, Epidemiology and Management of Pyrodinium Red
contamination of shellfish and planktivorous fish Tides. ICLARM Conference Proceedings 21. Fisheries De-
by the causative agent in Brunei Darussalam, partment, Ministry of Industry and Primary Resources,
Brunei Darussalam and International Centre for Living
Pyrodinium bahamense var. compressum. Never- Aquatic Resources Management, Manila, Philippines:
theless, the adverse impacts on fisheries may be 9-17.
overcome or minimised through the measures Jaafar, H. M., S. Subramaniam, M. W. R. N. De Silva &
described above. S. A. Mahali, 1991. Procedures of red tide action plan for
Brunei Darussalam (Unpublished Draft). Fisheries Depart-
Moreover, the Fisheries Department of Brunei
ment, Ministry of Industry and Primary Resources, Brunei
Darussalam has developed a highly reliable red Darussalam.
tide monitoring and response programme, which Roy, R. N., 1977. Red tide and outbreak of paralytic shellfish
is being upgraded into a Red Tide Action Plan poisoning in Sabah. Med. J. Malaysia, 31: 247-251.
(De Silva et al., 1989; Jaafar et al., 1991). Seliger, H. H., 1989. Mechanisms for red tides of Pyrodinium
With such action programmes, it is expected var. compressum in Papua New Guinea, Sabah and Brunei
Darussalam. In G. M. Hallegraeff & J. L. Maclean (eds),
that fisheries development, both in the capture Biology, Epidemiology and Management of Pyrodinium Red
and culture sectors, will continue to grow in Tides. ICLARM Conference Proceedings 21. Fisheries De-
Brunei Darussalam. partment, Ministry of Industry and Primary Resources,
Brunei Darussalam and International Centre for Living
Aquatic Resources Management, Manila, Philippines: 53-
71.
References Ting, T. M. & J. T. S. Wong, 1989. Summary of red tide and
paralytic shellfish poisonings in Sabah, Malaysia. In G. M.
Beales, R. W., 1976. A red tide in Brunei's coastal waters. Hallegraeff & J. L. Maclean (eds), Biology, Epidemiology
Brunei Mus. J., 3: 167-182. and Management of Pyrodinium Red Tides. ICLARM Con-
De Silva, M. W. R. N., A. H. M. Salleh, S. A. Mahali & ference Proceedings 21. Fisheries Department, Ministry of
S. Subramaniam, 1989. Management of Pyrodinium red Industry and Primary Resources, Brunei Darussalam and
tides in Brunei Darussalam. In G. M. Hallegraeff & J. L. International Centre for Living Aquatic Resources Man-
Maclean (eds), Biology, Epidemiology and Management of agement, Manila, Philippines: 19-26.
Pyrodinium Red Tides. ICLARM Conference Proceedings White, A. W., 1980. Recurrence of kills of Atlantic herring
21, Fisheries Department, Ministry of Industry and Pri- (Clupea harengus harengus caused by dinoflagellate toxins
mary Resources, Brunei Darussalam, and International transferred through herbivorous zooplankton. Can. H.
Center for Living Aquatic Resources Management, Manila, Fish. aquat. Soc. 87: 2262-2265.
Philippines.: 125-134. Wong, J. T. S. & T. M. Ting, 1984. Red tide and paralytic
Jaafar, H. M. & S. Subramaniam, 1984. Occurrences of red shellfish poisoning in Sabah, Malaysia. In A. W. White,
tide in Brunei Darussalam and methods of monitoring and M. Anraku& Kok-Kuang Hooi (eds), Toxic Red Tides and
surveillance. In A. W. White, M. Anraku & Kok-Kuang Shellfish Toxicity in Southeast Asian Fisheries Develop-
Hooi (eds), Toxic Red Tides and Shellfish Toxicity in ment Centre, Singapore and International Development
Southeast Asia. Southeast Asian Fisheries Development and Research Centre, Ottawa, Canada: 35-42.
Hydrobiologia 285: 227-235, 1994.
Water quality of Inanam River estuary and the Ko-Nelayan tiger prawn
aquaculture ponds in Sabah, Malaysia
Mazlin B. Mokhtar, Almah Bt. Awaluddin & Low Yew Guan

Faculty of Science and Natural Resources, Universiti Kebangsaan Malaysia, Sabah Campus, Locked Bag
62,88996 Kota Kinabalu, Malaysia
Abstract
Concentrations of selected metals (Cd, Co, Cr, Fe, Mn and Pb), oxidizable organic carbons, sediment
acid potentials, dissolved oxygen (DO), dissolved solids, suspended solids, pH, conductivity, salinity and
temperature in the Inanam River Estuary and the KO-Nelayan tiger prawn aquaculture ponds were
monitored during the period March to August 1989. Dissolved Co and Pb were found to be higher than
the recommended values of 0.05 mgl- I (Krenkel & Novotny, 1980; Nemerow, 1985), whereas the other
metals were comparable to the recommended safe levels. DO concentrations of the river and pond water
were in the range 2.6-4.7 mgl- I and 3.0-5.3 mgl- I respectively, both with an average which was lower
than the optimum value for the growth of prawns which is 5 mgl- I. Ferrous sulfide concentrations were
in the range 0.14-1.20%. Suspended solids were higher than the maximum (40 mgl- I ) recommended
value by WHO (1978). Other physical parameters were within the recommended range for optimal
growth of tiger prawns.
Introduction shops and poultry farms along its banks, and

these are potential sources of pollution. Domes-
The Ko-Nelayan aquaculture project for the cul- tic waste discharge from the village at the estuary
ture of tiger prawns has been in existence for a is also an important source of pollution.
number of years in an area situated close to the The purpose of this investigation was:
Inanam River Estuary (Fig. 1). In 1989 the farm
was not able to produce a satisfactory yield of (i) to study the water quality of the Inanam River
tiger prawns. A culture period of about four and its estuary;
months could only produce prawns with an av- (ii) to obtain information on the water quality
erage weight of only about 25 g which is smaller and sediment of the Ko-Nelayan Aqua-
than the average weight of about 40 g previously culture ponds in Inanam;
produced. Since disease was not the obvious (iii) to study the relationship between the water
cause of this problem the effect of metals present quality of the aquaculture ponds and the
in water and sediment of the pond and the Inanam Inanam River (water source for the aqua-
River was investigated. culture ponds).
The relationship between the water and sedi-
ment quality of these ponds and the Inanam River
was also investigated. Metals are given special Background of study area
focus due to the fact that the polluted Likas River
flows into the Inanam estuary. The Likas River The aquaculture ponds lie on the banks of the
has a number of light industries, motor work- Inanam River which is about 2 kilometers from
228
SCALE
o 0.5 KM
-===
, ,
,,
,
I
I
I
I
I
I
I
I
I
I
I
I
inanam ·River 11 /
I
,
,-- --
_ ••DaKlIOJoD , , .-
.,..,.......-
------
Key
• Sampling location
Road
Fig. 1. Location of sampling stations along the Inanam River, Sabah, Malaysia.
the mouth of the Inanam estuary (Fig. 1). The culture ponds is about 76 ha. The culture of these
Likas River comprises of two tributaries that is prawns comprises three stages using three ponds,
the Inanam River II which is short and the Likas viz. the nursery, transitional and grow-out ponds.
River which is long. Pig farms, poultry farms and Beginning with the nursery pond, these prawns
mechanical workshops are found along the are later transferred into the transition pond
Inanam River II. Domestic effluents also flow into followed by the grow-out pond. The water source
this river. The sources of pollution along the Likas of these ponds is the Inanam River, and about
River are industrial effluents such as food and 20-30% of the pond water is renewed weekly to
beverage factories, and also domestic effluents ensure its quality. Renewal is done during high
from the population living along this river. In the tide and the river water is first filtered but not
vicinity of the Ko-Nelayan aquaculture ponds, a chemically treated. Temperature, salinity and pH
bridge building project and a housing park project are routinely monitored at Ko-Nelayan. Dis-
were in progress at the time of this study. solved oxygen (DO) content in the ponds are con-
The area of the Ko-Nelayan tiger prawn aqua- trolled using mechanical wheel-paddlers.
229
Water and sediment samples were taken weekly pond AT3 and finally into the grow-out pond
during March 1989 to August 1989 from nine AG3. Group B started in AN7, then shifted to
aquaculture ponds in Ko-Nelayan and from three AT4 and finally to AG4. Prawns of group C were
different locations along the Inanam River cultured in BN1, then move to BTl and finally to
(Fig. 2). ANI is the water intake point for the BGl.
ponds from the Inanam River. Temperature, salinity, conductivity, pH, sus-
pended solids, dissolved solids, DO, total
alkalinity, acid potential of sediments, oxidizable
Materials and methods organic carbon and concentrations of metals such
as cadmium (Cd), cobalt (Co), chromium (Cr),
Water and sediment samples from the aquacul- iron (Fe) and lead (Pb) were studied.
ture ponds were taken weekly during this study. Surface water samples and bottom sediments
Three groups of P. monodon prawns were chosen were taken at random from three different points
for this study, i.e. group A, B, and C. Prawns in each pond. These samples were mixed to form
from group A were cultured in the nursery pond a single composite sample (Banerjee, 1967). Sur-
AN3, and later transferred to the intermediate face water and sediment samples from Inanam
SCALE
lcm:20m
8G1
Fig. 2. Location of Ko-Nelayan Aquaculture ponds in Likas, Sabah.

230
River at location 11 and 12 were taken from the nation of acid potential (Simpson et al., 1983;
middle of the river whereas samples from ANI Rosly & Ti, 1986) and metal concentrations. The
was taken near the banks. Sampling time was metals were divided into different fractions in the
between 0900 and 1200 hours. sediment, this was done according to the sequen-
Samples for the analysis of dissolved metals tial extraction method proposed by Tessier et al.
were obtained by filtering the water samples (1979). These are the exchangeable (Fl), carbon-
through a 0.45 f..tm millipore membrane. Dis- ate (F2), ferromanganese (F3), organic (F4) and
solved solids, suspended solids and total alkalin- residual (F5) fractions.
ity, were determined, on the same day. Sediment
samples were taken using the Ekman grab, and
frozen at less than 4 °C before analysis. Results and discussion
Physical parameters such as conductivity, tem-
perature, and salinity were measured in situ using Inanam River
a YSI 33 meter whereas pH was measured using
a Corning 115 pH meter. DO was measured with The environmental parameters such as tempera-
YSI 52 meter. Determination of suspended solids, ture, salinity and conductivity obtained from this
dissolved solids and total alkalinity was based on study were comparable to the previous studies
standard methods (APHA, 1985). done (Table 1). The values for the suspended
Sediment samples were dried to constant solids obtained from this study (93-99 mgl- 1)
weight at 105 degrees Celsius. Dried samples were were higher than those recorded by Murtedza &
later ground using mortar and pestle then sieved U sup (1987) (18-62 mgl- 1). This is caused by
using a 1 mm mesh sieve for the determination of the housing project currently under construction
oxidizable organic carbon (Walldey & Black, nearby. The maximum level recommended by
1934), and a 63 f..tm mesh sieve for the determi- WHO for suspended solids is 40 mgl- 1 (Sarmani,
Table 1. Physical and chemical parameters of Inanam River surface water obtained in this study and previous studies.
Parameter 1984" 1986 b 1987 c This study

1989
pH 7.2-7.5 6.5-7.5 6.64-7.97 7.32-8.03

Temp.oC 31.0 28.9-29.9 30.1-31.2 29.9-30.6
Salinity (%0) 20.0-28.0 26.5-28.7 23.6-30.9 21.6-24.8
Conductivity (mS m - 1) 41.0 45.0-46.6 38.0-47.4 39.82-44.8
DO (mg 1-1) 2.3-4.9 3.7-6.8 1.7-5.7 2.57-4.66
Suspended solids (mg 1- 1) 825-1408 490-710 18-62 93-99
Dissolved solids (mg 1- 1) 28.2-31.1 24.2-37.1 n.a. 27.5-28.7
Dissolved metals (mg 1- 1)
Mn 0.01-0.17 0.02-0.08 n.a. 0.08-0.11
Co n.a. 0.02-0.04 n.a. 0.17-0.19
Cr t.k.-1.15 n.a. n.a. 0.04-0.05
Cd n.a. O.oI-O.03 0.10-0.15 0.05
Pb 0.44-0.45 0.01-0.04 0.17-0.70 0.38-0.44
Fe 0.22-0.43 0.40-1.17 n.a. 0.13-0.21
a Foong (1984).
b Othman (1986).
c Murtedza & U sup (1987)
t.k. Not detected.
n.a. Not analysed.
231
1985}. Concentrations of dissolved solids were in Ko-Nelayan aquaculture ponds

the range of 27.69-28.72 gl- 1 which is similar to
previous studies (Table I). Minimum alkalinity in Physical & chemical parameters
the estuary is about 140-147 mgl- 1 Ca C0 3 and pH of the pond water was between 7.S-8.S and
this result is comparable to the normal value for this is within the range recommended for aqua-
sea water (Burton, 1976; Phillip, 1972). culture ponds (Law, 1988) The temperatures of
Overall the concentrations of dissolved metals surface and bottom water samples were within
were less than 1 mgl- I. The concentration range the range 29-31 °C. Salinity of water at the sur-
of Cd, Co, Cr, Fe, Mn and Pb for the three dif- face and the bottom of the ponds was in the range
ferent locations along the Inanam River is shown 22-24%0'
in Table 1. Pb was found to be high in all loca- DO of all ponds exceeded 2 mgl- 1 (3.0-
tions. The recommended safe levels of dissolved S.3 mgl- I ), the minimum value recommended by
Pb should be less than O.OS mgl- 1 (Krenkel & Law (1988). It seldom exceeded S mgl- 1, the op-
Novotny, 1980). Pb normally exists in undis- timum value for the growth of prawns (Chiang,
solved forms and its concentration in aquatic 1982). Alkalinity of the water lies within the range
systems is normally low except when pollution of 96-186 mgl- 1 CaC0 3 that is suitable for the
is indicated (Waite, 1984). It is apparent that growth of tiger prawns (Law, 1988). The concen-
the Inanam River is contaminated with Pb tration of suspended solids in all the ponds were
(Murtedza & Usup, 1987). All three locations in the range 7S-116 mgl- 1, but according to the
show increasing trends of Pb concentration with classification of Liong (1984) it is slightly pol-
time which is significant (p<0.01). luted.
The concentration of Co in the Inanam River
throughout this study remained in the range 0.17- Metals
0.19 mgl- 1 which is higher than the normal con- Comparisons between the concentrations of dis-
centration of dissolved Co in natural water (0.01- solved metals in the aquaculture ponds and the
0.03 mgl- I) (Sarmani, 1985). The increasing concentrations in the Inanam River indicate that
trend of Co with time is only significant (p<O.OI) the water quality of the ponds is influenced by the
for location ANI. The concentrations of Cd and water quality ofInanam River. The concentration
Cr were within the range of 0.04-0.0S mgl- 1 and of dissolved Pb was found to be the highest
this is equivalent to the normal value of these amongst the metals studied. The concentration of
metals in natural water (Sarmani, 1985). This dissolved Pb increased steadily from 0.3S-
range is lower than the values obtained by previ- 0.42 mgl- 1 in the nursery ponds to 0.44-
ous studies (Foong, 1984; Murtedza & Usup, 0.46 mgl- 1 in the juvenile ponds, and then in-
1987). The concentration of Mn (0.08- creased to 0.61-0.64 mgl- 1 in the grow-out
0.11 mgl- I) found is consistent with previous ponds. Similar increasing trends were also ob-
findings (Foong, 1984; Othman, 1986). served in the Inanam River during the same du-
The concentration range of Fe was 0.14- ration. The water quality of the ponds changed
0.21 mgl- 1 which is similar to values obtained in with changes in the water quality of the river. On
previous studies. Concentrations of undissolved other hand the concentration of dissolved Fe
Fe were in the range of 0.22-0.31 mgl- 1 and showed the opposite trend. Fe concentration de-
28-40 % of this was found to be adsorbed to creased from the range of 0.2S-0.32 mgl- 1 in the
suspended solids. Typical distribution of metal in nursery ponds to 0.IS-0.19 mgl- 1 in the juvenile
sediment is shown in Figs 3 & 4. The percentage ponds and further decreased to 0.11-0.13 mgl- 1
of metals in the residual fraction is high for all in the grow out ponds.
metals except for Pb. Pb was found mainly in the The water in the ponds was also contaminated
carbonate fraction. with Co. The concentrations of dissolved Co in
the ponds lie in the range 0.16-0.24 mgl- I which
232
Pond AN7
Pond AT4
..
..'"'"
;:
<.>
i
a.
Pond AG4
..;:''""
8
i
a.
~ Fl IS:SI f2 !Z1:3 FS ~ F4 ~ F6
Fig. 3. Percentage fractionation of metals in sediment samples of the ponds AN7, AT4 and AG4.
Station ANI 233
Station 12
"01
"
;:
8
;
"'-
Station 11
'"01
"~
u
;
Q.
~ Fl ISSI f2 ~ Fa ~ f4
Fig. 4. Percentage fractionation of metals in sediments of the Inanam River, Sabah, Malaysia.
234
exceeds the maximum concentration recom- Table 2. Potential acidity, contents of FeS and oxidizable
mended for the purposes of irrigation and agri- organic carbon in sediments of Ko-Nelayan aquaculture
ponds.
culture (Nemerow, 1985). The concentrations of
Mn, Cr and Cd were low with values below Pond Potential acidity FeS Organic carbon
0.1 mgl- 1 • (milli equiv./IOO g) (%) (%)
A typical distribution of heavy metals in the
AN3 28.6 0.87 0.11
different fractions of the sediment sample is
AN7 19.4 0.59 0.7
shown in Fig. 4. Similar to the river sediments BNl 39.6 1.20 0.8
(Fig. 3), the concentration ofPb was mainly found AT3 12.3 0.37 4.0
in the carbonate and ferromanganese oxides AT4 29.9 0.91 0.2
fraction. BTl 11.8 0.36 0.9
Am 4.7 0.14 1.3
In the sediment samples of all ponds, more
AG4 29.0 0.88 0.3
than 70 % of the iron were found in the residual BGI 8.9 0.20 1.2
fraction. The non-residual fraction of iron was
mostly found in the ferromanganese oxides
fraction.
Sediment-Mn was mainly found in the carbon-
ate fraction, whereas Co was mostly found in the
residual and the exchangeable fractions. Cr in the Conclusions
sediment was mainly found in the residual frac-
tion and its non-residual fraction being found This study shows that the water in the aquacul-
mainly in the organic fraction. Sediment-Cd was ture ponds and the estuary of the Inanam River
mostly concentrated in the residual and exchange- is contaminated by suspended solids, Co and Pb.
able fractions. These metals probably originated from light in-
The concentration ranges of these metals in dustries, and motor workshops and leaded fuel in
sediment samples of the ponds were lower or boat engines. All the metals in the sediments ex-
about equal to the mean baseline values reported cept Pb were mostly distributed in the residual
for coastal sediments (Aston & Chester, 1976) fraction. Pb in the sediments was mainly found in
and when compared to the sediment of the the carbonate and Fe-Mn oxide fraction. Changes
unpolluted Krka River Estuary in Yugoslavia in the water quality in the Inanam River influ-
(Prohic & Kniewald, 1987). enced the water quality in the ponds. DO content
was below the optimum level whereas tempera-
Sediment acid potential ture, salinity and conductivity were within the op-
Table 2 shows the mean values of potential acid- timum range for the cultivation of tiger prawns. In
ity, percentage content of ferrous sulfide, and the all the ponds the organic carbon content was low
percentage content of organic carbon in the sedi- and concentration of the pyrites was also low
ment samples of the aquaculture ponds. Ferrous (0.2-1.2%).
sulfide with a percentage content range of 0.14-
1.2% is lower than the 3.9% value reported by
Rosly & Ti (1986) for the aquaculture ponds in Acknowledgements
Gelang Patah, Johore. The low organic content of
the sediment is consistent with the low content of We would like to express our appreciation to the
heavy metals in the organic fraction. pH of the staff of KoNelayan, especially Mr Samson Shak,
bottom water samples, i.e. the water just above Mr Julius Lojungin and Ms Claire Lojiu. Our
the surface of the sediment, was acidic (pH < 6). thanks to Universiti Kebangsaan Malaysia for
This might be an indication for the oxidation of financial assistance (Research Grant UKM 29/
ferrous sulfides (iron pyrites) and organic carbon. 89).
235
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Assoc., A WWA, A WPCF. Washington. 493 pp.
Chemistry, National University of Malaysia (Sabah Cam-
Aston, S. R. & R. Chester, 1976. Estuarine sedimentary
processes. In J. D. Burton & P. S. Liss (eds), Estuarine pus), Kota Kinabalu, Malaysia.
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Banerjee, S. M., 1967. Water quality and soils condition of Barnes & J. Green (eds), The estuarine environment.
fish ponds in some states of India in relation to fish pro- Applied Science Publishers Ltd: 33-47.
Prohic, E. & G. Kniewald, 1987. Heavy metal distribution
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Burton, J. D., 1976. Basic properties and processes in estua-
of sequential extraction analysis. Mar. Chern. 22: 279-
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rine Chemistry. Academic Press, London: 1-31. 297.
Chiang, D. M., 1982. Feed, feeding culture technology and Rosly, B. H. & T. L. Ti, 1986. Observation on acid runoff and
iron in brackish water fishponds. Problems and implica-
management. Hanaqua Feed Corporation, Taiwan: 1-6.
tions. Fisheries Bulletin 43. Fisheries Dept., Ministry of
Chiang, P. & I. C. Liao, 1985. The practice of grass Prawn
(Penaeus monodon) culture in Taiwan from 1968 to 1984. J.
Agriculture, Kuala Lumpur, Malaysia.
World Maric. Soc. 16: 297-315. Sarmani, S., 1985. Kajian pemonitoran kualiti air Lembangan
Foong, C. S., 1984. Kajian Garis Dasar Kualiti Air Di Sungai Langat, Selangor. Sains Malaysiana 14: 245-255.
Lembangan Sungai Inanam. Hons. thesis, Dept. of Chem- Simpson, H. J., H. W. Ducklow, B. Deck & H. L. Cook,
1983. Brackish-water aquaculture in pyrite-bearing tropical
istry, National University of Malaysia (Sabah Campus),
soils. Aquaculture 34: 333-350.
Kota Kinabalu, Malaysia.
Krenkel, P. A. & V. Novotny, 1980. Water quality Manage- Tessier, A., P. G. C. Campbell & M. Biosson, 1979. Sequen-
ment. Academic Press, New York: 112-147. tial extraction procedure for speciation of particulate trace
metals. Analyt. Chern. 51: 844-851.
Law, A. T., 1988. Water quality requirements for Penaeus
Waite, T. D., 1984. Principles of Water Quality. Academic
monodon. In K. J. Ang& S. Tarlochan (eds), Marine Prawn
Farming in Malaysia. Proc. Seminar Marine Prawn Press, London.
Walkley, A. & I. A. Black, 1934. An examination of the
Farming in Malaysia. Malaysian Fisheries Society,
Serdang: 53-60. Degtjareff methods for determining soil organic matter, and
Liong, P. C., 1984. Water quality standards for aquaculture in a proposed modification of the chromic acid titration
Malaysia. Fisheries Bulletin 20. Dept. of Fisheries, Minis- method. Soil Sci 37: 29-37.
WHO, 1978. Environmental health criteria. Principles and
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Murtedza, M. & G. Usup, 1987. Water Quality and Hydro- methods for evaluating the toxicity of chemicals. Part I:
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graphy of Buat-Inanam-Likas Estuary. Report to Jurutera
Konsultant Sdn. Bhd. Kota Kinabalu, Malaysia, 60 pp.
Hydrobiologia 285: 237-247, 1994.
Prolonged inundation and ecological changes in an Avicennia mangrove:

implications for conservation and management
Satish C. Choyl & Webber E. Booth

Universiti Brunei Darussalam, Gadong 3186, BSB, Brunei Darussalam, Borneo
I Present addres: Australian School of Environmental Sciences, Griffith University, Nathan, Qld., Australia
Key words: prolonged flooding, A vicennia, ecological changes, mangrove management
Abstract
The mangrove around Sungei Pemburongunan, just west of Tanjong Batu and Istana Darul Aman in
the Brunei-Muara District, is unique in that it contains the only pure stand of Avicennia marina in Brunei
Darussalam. In mid-October, 1990 the mouth of Sungei Pemburongunan was closed due to a natural
build-up of a sandbar across it. The mangrove quickly became flooded and adverse effects on the flora
and fauna were observed during the following weeks. In mid-December a channel was dug across the
sand bar to reduce the flooding and normalise the ecology. Some of the dying A vicennia and Lumnitzera
have since recovered but others such as Acrostichum and Casuarina have not. Avicennia showed some
interesting responses to prolonged inundation. Many of the A vicennia that had flowered during the flood
produced seedlings which are now well rooted and 40-80 cm in height. The survival of large numbers
of these seedlings is attributed to the floor-related decrease in the number of grapsid and sesarmid crabs
which predate on them. The crab and mollusc populations have largely recovered. Unlike periodic
short-term flooding which does not seem to have any apparent adverse affect on the ecosystem, pro-
longed inundation can result in the loss of the A vicennia and its associated flora and fauna. Thus, fragile
mangrove ecosystems such as this need to be monitored and managed closely.
This work demonstrates that contrary to conventional wisdom, human intervention and management
can be beneficial to more fragile ecosystems, which could otherwise succumb to natural processes.
Furthermore, it highlights the importance of the dynamic nature of the environment which should be
considered in management and conservation programmes.
Introduction used Brooketon coalmine and covers an area of

just over 50 ha. The 1: 10 000 Brunei map for the
The Meragang wetland (Fig. 1) around Sungei area (Sheet 55/95 NE) does not indicate the pres-
Pemburongunan, just west of Tanjong Batu and ence of any vegetation in the 'swamp', about 50%
Istana Darul Aman in the Brunei-Muara district, of which is actually covered by A vicennia marina
is unique in that it contains the only pure stand and, to a lesser extent, by other plants such as
of Avicennia marina (Forsk.) Vierh. in Brunei Casuarina, Acacia, Lumnitzera, Nypa, Excoecaria
Darussalam. In other areas of the country, this and Pandanus. Only the southern and western
plant occurs as mixed stands with other man- fringes are devoid of any macrophytes. They are,
groves, such as Rhizophora, Sonneratia and Bru- however, covered by a dense layer of fine algae.
guiera. The wetland is just west of the now dis- There is evidence that these areas were, until
238
0°
100°
T
\
\
I
/
/
I
Fig 1 Locality map of Brunei Darussalam (BD), the Avicenma manna wetland at Meragang (M) and the samplmg statIons (1-
14) ----, wetland hmlts, .... , beach sand limits, - ' - , road.
recently, covered by Pandanus sp., root stumps gillnets, crab nets, crab traps and tidal weirs),
being still visible. mollusc harvesting and recreation.
Apart from its ecological importance, the Ecological baseline studies were carried out
Meragang wetland is of social and aesthetic sig- during the early part of 1990. In mid-October of
nificance. Human activities observed in the area the same year the mangroves became flooded due
include fishing (using a variety of gear such as to a natural build-up of a sandbar across the
239
mouth of Sungei Pemburongunan, the stream 3. Soon after the flood receded (January, 1991)
flowing through the area. The whole wetland 4. Nine months later (September, 1991)
quickly became flooded; water level in the A vi-
cennia marina grove and on the adjacent mudflats Numerous visits to the site were also made
reached a height of about 0.5 m above MHWS. between these periods. On these occasions and
The area remained under flood for eight weeks, during the flood, however, only qualitative obser-
during which time increasing mortality of the vations were recorded.
mangrove flora and fauna was observed. On
16 December, a channel was manually dug to re-
lease the floodwater. Results
These events provided an unique opportunity
for us to study the ecological responses of the The Meragang wetland has, as its dominant spe-
mangrove flora and fauna to prolonged inunda- cies, A vicennia marina. Other components of the
tion and their subsequent recovery. The episode flora include Nypa fruticans (Thunb.) Wurmb.,
also forced us to reassess our views on certain
approaches to mangrove conservation and man-
agement. Table 1. Plant denSity and the percentage mortality m the
A Vlcenma wetland, after eight weeks of mundatlon Hyphen-
ated station numbers mdlcate samplmg plots between the two
deSignated stations.
StatIon no SpecIes DenSIty Percentage
(no 100 m - 2) mortalIty
Standard ecological methods were employed to
survey the flora and fauna of the Meragang wet- 3 A vlcenma marina (Forsk ) 25 0
land. Using 1: 10 000 maps and aerial photo- Excoecarla agallocha L 1 0
Acrosllchum aureum L 100
graphs, field surveys were carried out to map the
4 A marina 17 0
vegetation types. Based on this information, 4-5 A marina 32 50
stratified random sampling was performed. Ten Cassuarma equIsltzjoiza L 100
A marina 33 73
by ten metre plots were used to estimate the
C equIsltzjoiza 7 100
density of the mangrove plants at the different E agallocha 3 0
sampling stations (Fig. lc). The same plots were Lummtzera racemosa Wdld 6 50
revisited at different sampling times. Macroepi- Pandanus sp 3 100
Grass 75
faunal (particularly crustaceans and molluscs) Rush 100
densities were estimated using five replicates of 6 A marina 15 0
either 0.016 m2, 0.25 m2 or 2 m2 quadrats within 7 A marina 29 86
7-8 A marina 74 31
and between the permanent, demarcated vegeta- 8 Nypa Jrutlcans (Thunb ) 3 0
tIon plots. The smaller quadrats were used for 9 A martna 25 100
10 A manna 17 59
high density, less mobile molluscs such as Cer-
E agallocha 2 50
ithidea while the largest quadrat was for more Grass 75
widely dispersed or mobile organisms such as 11-12 A manna 50 0
11-14 A manna 43 47
Telescopium and the crabs. The faunal quadrats
E agaUocha 6 0
samples were taken within the same general area Acrostzchum speclOsum 0
at different sampling times. Fish fauna in the DerriS trzJolzata Lour 3 67
stream were recorded from catches made by fish- C equIsltzjolza 2 0
Asplemum sp 0
ermen. Four sets of surveys were carried out: Pandanus sp 0
CymbIdIum sp 0
1. Prior to flooding (March-June, 1990) Legummosae 0
13-14 A manna 61 0
2. During the flood (November-December, 1990)
240
Excoecaria agallocha L., Lumnitzera racemosa (1.5-5 m) plants (Fig. 2). There was total mor-
Willd. and Derris trifoliata Lour. Casuarina equi- tality at stations 7 and 9. Most tree-sized (> 5 m
sitifolia L., Acrostichum aureum L. and A. specio- in height) Avicennia survived although 50% of the
sum L. occupy slightly raised sites within the wet- low density plants at stations 4-5 and 7-8 died.
land. Several species of rushes and grasses are The A vicennia seedlings shown in Fig. 2 actually
also present on higher, firmer ground. Perched on appeared soon after the flood and have grown to
old A vicennia stumps are the epiphytic orchid, a height of 40-80 em in nine months.
Cymbidium sp. and the ferns, Asplenium sp. and Some A vicennia displayed necrosis of the bark
Drynaria sp. A dense cover of micro- and meio- and surface wood tissues (to a depth of about
algae is present on the mud surfaces of the wet- 1 cm) in the region of the stem that was sub-
land. merged during flooding; yet they survived. Large
Most plant species suffered some mortality diameter (0.5-1.5 cm) adventitious roots have, in
during the prolonged inundation (Table 1). Avi-
cennia suffered substantial losses of shrub-sized Table 2. Mollusc abundance in the A vicennia wetland during
different periods. 1, prior to flooding; 2, just after flooding had
receded; 3, nine months later; + rare; + + occasional; + + +
Denslly
60.-~-----------------------------------, common; + + + + abundant.
50 . Relative abundance
40
2 3
30
Gastropoda
20
Assiminea brevicula (Pfeiffer) +++ + ++++
Cassidula mustelina Deshayes ++ + +
C. pilosa Gassus + + +
4- 1) 5 6 7-6 11-12 11-14 13- 14
Cerithidea cingulata (Gmelin) ++++ + ++++
Station Number C. obtusa Lamarck +++ + ++
_ Trees _ Shrubs I3ZJ Seedlings
Chicoreus capunicus (Lamarck) ++ + +
Ellobium aurisjudae L. ++ + +
E. aurismidae L. + + +
Liltoraria carini/era (Menke) ++ ++ +
L. scabra (L.) ++ ++ +++
Nassarius olivaceus Brug. ++ + +
Percent Moneilly Nerita planospira Anton ++ + +
N. balteata Reeve ++ + +
Telescopium telescopium (L.) +++ +++ +++
100
Terebralia sulcata (Born) +++ + +++
60 Thais tissoti (Petit) ++ + +
Theodoxus oualauiensis (Lesson) ++ + +++
60 Heminoea sp. ++ + +
40 Onchidium sp. ++ ++ +++
20 Bivalvia
Anadara granosa lredale + + +
0'--'----'--
3 4-0 I) 6 7- 6 10 11-12 11-14 13-14 Barbatoa sp( + + +
Ststi on Number
Enigmonia enigmatica (Ired ale) + + +
_ T,ee8 _ Shrubs £illI] Seedlings Gafrarium tumidum (ROding) ++ + +
Geloina coaxons +++ ++ ++
Fig. 2. Density and percentage mortality (as a result of the Glauconome sp. ++ + +
flood) of Avicennia marina. Trees, > 5 m; shrubs, 1.5-5 m; /sognomon amonoides (Reeves) ++ ++ ++
seedlings, < 1.5 m in height. Density of trees and shrubs, Saccostrea echinata (Q & G) +++ + ++
no. 100m - 2; density of seedlings, nO.m - 2. Plots between des-
Solen sp. ++ ++ ++
ignated stations are hyphenated.
241
many cases, begun to grow from just above the the mudflats have since been reduced markedly in
necrotic region. A large percentage of A vicennia numbers. Assiminea brevicula (Pfeiffer) and Theo-
pneumatophores died back either partially or doxus oualauensis (Lesson) completely disap-
completely as a result of the flooding. Side shoots peared during the flood but since then, have re-
have subsequently developed from the basal living appeared in some places in large numbers.
portion of some of the original pneumatophores. Assiminea shows preference for drier areas of the
Small numbers of new ones have also been mudflat (P < 0.0 1). Telescopium, Terebralia and
produced. An additional response of A vicennia to Theodoxus have reappeared in the wet zone
the flooding has been the stimulation of flowering (mudflat streams) but have yet to fully recolonise
and fruiting by plants of all ages and sizes; some the drier mudflat areas (cf. Figs. 3 & 4). Cerithidea
as small as 60 cm in height. cingulata (Gmelin) also suffered extensive mortal-
The relative abundance of m.olluscs prior to ity as a result of the flood. However, it has re-
flooding, soon after the flood receded and colonised the area in greater than pre-flood den-
9 months later is given in Table 2. Densities and sities, particularly in the wet zones (Fig. 5). The
percentage survival of the more common species length of the dead shells were 17 ± 2 mm while
are given in Table 3 and Figs. 3-5. Densities of those newly recruited in January were 2 ± 1 mm
Terebralia sulcata (Born) and Telescopium tele- long.
scopium (L.) increased significantly (P< 0.01) Telescopium and Terebralia from within the area
during the flood. However, those which colonised reproduced soon after the flood receded and ju-
Table 3. Mollusc density (x ± S.E., no. m - 2) and mortality (% dead, in brackets) in the Avicennia wetland. 1, preflooding; 2, a
few days after the flood receded; 3, nine months later. Dry, moist mudflat zone; wet, mudflat stream and pool zones. At each
station, densities of species between episodes 1 & 2 and 2 & 3 are significantly different (P<O.OI).
Stat. no. Species 2 3
6 (dry) Cerithidea cingulata 344 ± 50 (8) 390 ± 48 (10) 1920 ± 300 (60)
Terebralia sulcata 2±1 (0) 13 ± 2 (0) 1.25 ± 0.2 (75)
Telescopius telescopius 1± 1 (0) 5±1 (50) 0.5 ± 0.1 (100)
Assiminea brevicula 11 ±4 (0) 0 96±5 (0)
Theodoxus oalauensis 13 ±2 (0) 0 0
Littoraria spp. 5±3 (0) 2 (0) 12±2 (0)
Geloina coaxons 2±1 (100) 8±2 (100) 2±1 (100)
6 (wet) C. cingulata 731 ± 100 (5) 56 ± 10 (93) 3156 ± 95 (6)

T. sulcata 6±2 (5) 28 ± 7 (15) 192 ± 24 (5)
T. telescopium 3± I (5) 16±2 (0) 64± 15 (0)
A. brevicula 0 0 0
T. oalauensis 12 ± 4 (0) 0 96±7 (0)
9 C. cingulata 324 ± 54 (5) 46 ± 10 (98) 1500 ± 100 (40)

T. sulcata 4±2 (0) 15 ± 3 (10) 16 ± 3 (20)
T. telescopium 3± I (0) 7±2 (10) 10 ± 3 (10)
10-11 C. cingulata lOll ± 135 (5) 350 ± 55 (95) 1280 ± 150 (0)
T. sulcata 2±1 (0) 12±3 (5) 2±1 (30)
T. telescopium 6±2 (5) 9 ± 3 (5) 5±2 (6)
14 C. cinqulata 480 ± 87 (10) 410 ± 78 (100) 380 ± 65 (90)

T. sulcata 1± 1 (0) 3 ± 2 (10) 1 ± 1 (0)
T. telescopium 1± 1 (0) 3 ± 2 (0) 1 ± 1 (0)
242
Terebralla Telescoplum Asslmlrl9a ThaodOxus Asslmlnea Thaodoxus
_ Prallood _ JuSI 811er llood C!El g mlhs. laler _ Pre fl ood _ JUSI allar flood []ill 9 mlhs. later
Percent Survival
Peroent Survival
Asslmlnea Thaodoxus
_ Preflood _ J usl aller flood I'§D 9 mlhs. lale(

_ Prell ood BJuslaller flood [fZJg mlhs. later
Fig. 4. Density (no.m - 2) and percentage survival offour spe-
Fig. 3. Density (no.m - 2) and percentage survival offour species of molluscs in the wet mudflat areas of the mangrove.
cies of molluscs in the dry mudflat areas of the mangrove.
crustacea (Table 4). Most of the more common

veniles from that single cohort have grown to species, which had virtually disappeared during
23 ± 4 mm and 18 ± 4 mm, respectively in nine the flood have since reappeared (Fig. 6). During
months. Assiminea, Theodoxus and Cerithedia the flooding, mudlobsters Thalassina anomala
seemed to have been recruited from outside. In (Herbst) were commonly seen out of their flooded
nine months, Assiminea grew to 3 ± 2 mm and burrows during the daytime and, along with
began breeding in July. Theodoxus grew to sesarmid crabs, climbing trees. Newly moulted
5 ± 2 mm in the same period while Cerithedia has specimens were often observed being killed and
had two waves of recruitment, the first just after eaten by crabs such as Scylla serrata (ForskAl)
the flood receded and the second in May-June. and Thalamita spp. Areas where the mangroves
These have grown to 11 ± 2 mm and 6.7 ± 1.4 mm had died as a result of the flood and, previously
in nine and five months, respectively. Post-flood occupied by small sesarmid crabs (such as Chi-
Cerithedia and Theodoxus have yet to breed in the romantes, Parasesarma and Clistocoeloma), have
Meragang wetland. been colonised by the larger Metaplax (Fig. 6,
The other very abundant and conspicuous stn.6). Uca annu/ipes (Latrielle) and many of the
invertebrate group in the A vicennia wetland is the sesarmid crabs reproduced during May-June; the
243
3500 DenaHy
Table 4. Relative abundance of crustaceans in the A vicennia
wetland. 1, before flooding; 2, just after the flood receded; 3,
nine months later, +, rare, + + ,occasional; + + +, common;
+ + + +, abundant. Taxonomic authoritIes can be found in
Sasekumar (1974) and Lovett (1981).
Relative abundance
2 3
Cirripedia
Chthamalus malayensis +++ + ++
WeI Zona 6 Dry Zone 6 WeI zone 14 Dry Zone 14
Balanus sp. +++ + ++
-- • Praiiood _ JuS) aftor flood mE'ile mths, later
Isopoda
Limnoria sp. ++++ + +++
Sphaeroma sp. ++++ + +++
Decapoda
Penaeidae
Metapenaeus brevicomis ++ ++ ++
M. ensis ++ ++ ++
Parapenaeopsis hardwickii + + +
P. hungeifordi + + +
Penaeus merguiensis ++ ++ ++
P. indicus ++ ++ ++
P. monodon +++ +++ ++
P. semisulcatus ++ + +
Sergestidae
Acetes indicus + + +
A. erythreaus + + +
A. sibogae + + +
. Praflood _ Just after f lood EREI 9 mths. lalel Palaemonidae
Palaemon spp. ++ + +
Fig. 5. Density (no.m - 2) and percentage survival of Cer-
ithidea cingulata in the wet and dry zones of sampling sta-
Alpheidae
Alpheus sp. + + +
tions 6 and 14.
Anathus sp. + + +
Synalpheus spp. ++ ++ +
Portunidae
former have grown to 5-8 mm in carapace width Charybdis spp. ++ +++ +
in four months. Portunus pelagicus ++ +++ +
Thirty two species of fish (Table 5) have been P. sanguienolentus ++ ++ +
Scylla serrata +++ +++ +++
recorded from the stream which meanders
Thalamita spp. +++ +++ ++
through the A vicennia mangrove. Our observa- Grapsidae
tions indicate that many of the larger species such Chiromantes eumolpe ++++ + +++
as the Lethrinids, Carangids, Lutjanids, Serranids C. indiarum +++ + ++
and Mugilids are visitors and come in during high Clistocoeloma merguiensis ++++ + +++
Metaplax elegans ++ + +++
tides to feed. Others, such as the Sphyraenids,
M. tredecim ++ + +++
Lutjanids, Siganids, Theraponids and some Metapograpsus frontalis ++ ++ ++
Leiognathids, use the area as a nursery. The more Parasesarma plicatum ++++ + +++
permanent ichthyofauna include the Gobids, P. rutilimanum ++++ + ++
Toxotids, Scatophagids, Periophthalmids, Cen- Sarmatium spp. ++++ + +++
Leipocton sordidulum ++++ + +++
tropomids, Ariids, Plotosids, Theraponids and
Nanosesarma andersomi +++ + ++
some Leiognathids. Most of the large species are
244
Table 4. (Continued) Table 5. Fish collected from the A vicennia wetland stream.
Taxonomic authorities can be found in Matsuda, et al. (1984).
Relative abundance
Family Scientific name Local name
2 3
Rhinobatidae Rhynchobatus sp. pari
Ocypodldae Ariidae Arius spp. gagok, utek
Uea annulipes ++++ + ++++ Belonidae Hemiramphus sp.
U. rosea +++ + +++ Tylosurus sp.
U. triangularis +++ + +++ Carangidae Caranx spp. bamasa, menokok
Maerophthalmus erimtus +++ + ++ Centropomidae Chanda spp.
Xanthidae Toxotidae Toxotes jaculator
Heteropanope glabra ++++ + +++ Gobldae Hemigobius sp.
Coenobltidae Pseudogobius spp.
Coenobita sp. +++ +++ ++ Brachygobius doriae
Paguridae Lactariidae Lactaria sp. kelapa-kelapa
Clibanarius spp. ++ + +++ Leiognathidae Gazza minuta bilis pilaj au
Thalassinidae Leiognathus bindus bilis
Thalassina anomala +++ +++ ++ L. equulus pulup-pulut
L. sp/endens bilis
Secutor insidator bilis
Lethrinidae Le thrinus lentjan anduping
Lutjanidae Lut}anus spp. ketambak
regularly caught by the fishermen. During the L. argentimaeulatus merah
flood there was increased fishing activity and Mugilidae Mugil sp. balanak
Valamugil sp. balanak
initially the fish size was maintained but as the
Liza sp. kembura
flooding persisted the number of larger fish caught Periophthalmidae Periophthalmus sp.
declined. The species composition, however, re- Boleophthalmus sp.
mained similar throughout. No fish kills were ob- Platacidae Platax sp. bunda
ser·ved during the flood. Soon after the flood re- Plotosus sp. badukang
Scatophagidae Scatophagus argus kitang
ceded, gobies and mudskippers which appeared
Serranidae Epinephalus spp. kerapu
to have increased in abundance began to breed. Slganidae Siganus spp. belais
Juveniles grew to 13 ± 3 mm and 18 ± 3 mm In Sphyraenidae Sphyraena sp. linkoh
standard length, respectively in six months. Tetraodontidae Tetrodon sp. piasau-piasau
Theraponidae Therapon jarbua kirang-kirang
Donslly
l00.-~----------------------------------~
80
Discussion
80 Although there is substantial literature on human-

induced stresses to mangroves, very little is avail-
40 able on the effects of natural disturbances (Lugo
& Snedaker, 1974; Johns, 1986). What there is of
the latter mainly deals with either periodic or large
scale factors such as cyclones, tidal waves,
Seop. 1 Dey. 1 Uea 2 Sas. S Uee 3 Sos. 6 M
ol. 6
droughts, lightning strikes, etc. (Jimenez et al.,
_ Prafloed _ J usl . lar
f flood (j]j] 9 m lha. fatar 1985; Johns, 1986). These studies deal mainly
with immediate effects and the recovery dynam-
Fig. 6. Density (no.m - 2) of crabs at stations I, 2, 3 and 6 of
the mangrove. Scop., Seopimera bitympana; Ocy., Ocypode
ics of the mangrove ecosystem have been largely
cordimana; Uca, Uca spp.; Ses., sesarmids; Met., Metaplax ignored. Our study can therefore be discussed in
spp. the light of this deficiency.
245
The eight-week flooding of the Meragang wet- classes. A feature noted soon after the flood re-
land adversely affected populations of some of lease was the development of dense mats of black
the resident plant and animal species. Evidence fungal hyphae on the dead shrubs. There is a
based on the degree of disturbance and mortality possibility that fungal infection may have played
of the dominant species suggest that longer inun- a role in the mass mortality, a hypothesis that fits
dation could have resulted in the destruction of in well with blanket deaths in some areas where
the entire A vicennia marina ecosystem. Although root to root contact may have allowed the fungus
there have been some previous closures, none of to spread in the physiologically weakened plants
them lasted for more than a fortnight. The open- during the flood. However, we have no direct
ing of the mouth of the stream to allow the drain- evidence that such a pathogen was in fact in-
age of the floodwater and restore the original tidal volved. The fungus observed could have been a
patterns of marine and fluvial water exchange has, saprophyte that invaded the plants after they had
after a period of about nine months, allowed the died.
wetland to return close to the original community Unlike the other plants in the flooded area,
structure. However, there have been some A vicennia marina displayed adaptations which as-
changes in population densities and patterns of sisted its survival and re-establishment.
species distribution. In spite of the ring barking and necrosis of the
As yet unexplained is why one size class of outer wood of some A vicennia, the deeper inner
A. marina only displayed mass mortality. Few wood with its multiple cambia (Zamski, 1979)
large trees died and seedlings were, at the time of has been able to maintain sufficient conductance
the flood, present in insufficient numbers to be to allow the damaged plant to survive. Many have
considered. Almost all pneumatophores of large produced adventitious roots from just above the
and shrub-sized trees that were submerged died necrosed zone. These have grown rapidly towards
back from their tips as a result of the flooding. If, the soil forming Rhizophora-like prop roots. These
as reported in most treatises on mangroves, the new roots assist in supporting the damaged plants
pneumatophores have an important vent role for and supplement the conductive role of the rest of
respiratory gases (Chapman, 1976; Stewart & the root system.
Popp, 1987), then both size classes should have The ability of the pneumatophores to form lat-
died. The fact that the larger trees and some of the eral shoots below their flood-damaged tips is an-
shrubs survived means that A. marina plants can, other adaptation for recovery. Many Avicennia
for long periods survive without pneumatophores. are currently operating on 10-50% of their origi-
This brings into question the essential respiratory nal pneumatophores and most of these are in the
role of these structures (Tomlinson, 1986). Ma- form of recently produced lateral shoots.
nipulative investigations need to be undertaken to There has been an increase in the incidence of
determine the minimal pneumatopore require- flowering and fruiting and so large numbers of
ment for each size class of these mangroves. propagules have become available for reseeding
During the flood the salinity did not vary greatly the affected wetland. The result has been the ap-
from the normal situation (10-14 gl- 1) and there- pearance of high densities of A vicennia seedlings
fore the plants should not have been subjected to of the same age class in extensive areas of the
extreme levels of osmotic stress. wetland where, prior to the flood, they were un-
The majority of the shrub deaths were of trees common. Their survival, just after the flood re-
on ground away from the central creek. The large ceded, has been attributed to the timely tempo-
trees were either close to the creek or on raised rary decrease in the number of sesarmid and
islands associated with Thalassina mounds. The grapsid crabs which normally predate on the
differences in position or substrate may therefore newly fallen propagules (G. Maxwell, pers.
have either directly or indirectly had an influence comm.). Although crab populations have recov-
on the difference in survival between the two size ered, these seedlings are now well established and
246
beyond being susceptible to crab damage. How- flooded area would have been eventually fished
ever, very few new seedlings have since appeared. out had the stream mouth not been opened as
Thus, the result has been establishment of a single regular supply of young from the sea is required
age class of seedlings. Such age discontinuities to maintain species diversity and abundance.
(often despite continuous seedling production) in Since the mouth opened and the flood receded,
mangroves are not uncommon and may be ex- many juveniles of a variety of organisms includ-
plained in terms of changing patterns of predation ing fish, molluscs and crustaceans have been ob-
pressure (Osborne & Smith, 1991; Robertson, served within the mangrove channels and on the
1991). mudflats. Some of these were produced within
The disappearance of many species of the the area while others have been recruited from the
mudflat fauna as a result of the flood may be sea.
attributed to a single, or more likely, a combina- The ecological functions and significance of
tion offactors. These include the inability to carry mangrove ecosystems have been stressed on nu-
out sufficient gaseous exchange, problems in ionic merous occasions and is one of the highlighting
regulation, toxic chemicals, lack of food and pre- features of the ASEAN-ICLARM-US Coastal
dation. The high mortality of Cerithidea cingulata Resources Management Project (see Chua et al.,
may be due to the first factor. Although C. cingu- 1987). The mangroves of Brunei Darussalam are
lata has the ability to carry out aquatic as well as still well preserved but it is inevitable that some
aerial respiration, it seems better adapted for the areas will be lost to development of agriculture,
latter (Rao et al., 1986). Prolonged submergence aquaculture, industry and housing. The impact of
could have resulted in respiratory distress. Tere- such losses can be minimized by planned devel-
bralia and Telescopium are normally found in the opment and rational management. The proposed
wetter areas of the mangroves. They seem well National Mangrove Management Plan is a major
adapted for prolonged submergence as well as step towards this.
aerial exposure, provided the conditions are Most management strategies, however, con-
moist. They survived the flood. However, when sider only large tracts of mangroves and very often
the flood receded many of them were caught out small groves can disappear unnoticed. The rela-
in exposed areas which quickly became dry. These tively small A vicennia marina wetland around
died of desiccation. Most of the other molluscs Sungei Pemburongunan may succumb if it is not
may have suffered high predation from fish and managed specifically. Its closeness to the city and
swimming crabs which moved onto the mudflats its easy accessibility makes it very vulnerable. It
during the flood. could be easily made a target for conversion into
Grapsid (Grapsinae and Sesarminae) and other uses such as aquaculture pondification and
Ocypodid crabs, which virtually disappeared as a real estate development. Its survival is therefore
result of the flood, have recovered in number. under threat.
Metaplax spp. have colonised recently killed A vi- The first reaction regarding the conservation of
cennia areas which were previously inhabited by a small mangrove ecosystem such as this would
other sesarmids. It seems that these sesarmids be to declare it as a preserve. However, total
prefer covered areas while Metaplax spp. show preservation often leads to irregular monitoring
preference for more exposed ones. Changes in and non-management. Such an approach may
substrate quality and competition may have also not work as the mangroves here could die a natu-
contributed to this segregation. No fish kills were ral death, a result of the dynamic processes of
reported in the area during the flood. The flood ecosystems. The build-up of the sandbar across
water remained saline and oxygenated through- the mouth of the stream is a manifestation of this.
out the period (10-14 gl-l and 50-70% satu- Paradoxically, we feel that in this particular case
rated, respectively) and therefore tolerable to the only human intervention can help the mangrove
fish. Most fish, molluscs and crustaceans in the survive. It needs to be monitored and managed
247
closely. This can be effectively done if the area is Jimenez, J. A., A. E. Lugo & G. Cintron, 1985. Tree mortality
in mangrove forests. Biotropica 17: 177-185.
utilized in a non-destructive way.
Johns, R. J., 1986. Natural disturbances in mangrove ecosys-
A recommendation that the area be developed tems. In S. Cragg & N. Polunin (ed.), Report of the work-
into a recreational cum educational facility was shop on mangrove ecosystems dynamics. UNESCO/
made to the Department of Town and Country UNDP, New Delhi: 37-44.
Planning, Ministry of Development and an en- Lovett, D. L., 1981. A guide to the shrimps, prawns, lobsters
and crabs of Malaysia and Singapore. Universiti Pertanian
couraging response was received (Director of
Malaysia, Serdang, 156 pp.
Town & Country Planning, in litt.). Background Lugo, A. E. & S. C. Snedaker, 1974. The ecology of man-
information and details of our recommendation groves. Annu. Rev. Ecol. Syst. 5: 39-64.
are given in Choy & Booth (1991). The benefits Matsuda, H., K. Amaoko, C. Arasa, T. Uyeno & T. Yoshino,
of such a development will be that, besides cre- 1984. The fishes of the Japanese Archipelago. Tokai Uni-
ating greater public awareness of the natural heri- versity Press, 437 pp.
Osborne, K. & T. J. Smith III, 1990. Differential predation on
tage, the A vicennia mangrove and its associated mangrove propagules in open and closed canopy forest
flora and fauna at Meragang will be conserved habitats. Vegetatio 89: 1-6.
through closer monitoring and appropriate man- Rao, D. G. V. P., V. P. Rao, K. S. R. P. Rao, V. U. Devi &
agement. S. K. Patnaik, 1986. Studies on the respiration of cerithiids.
In M. F. Thompson, R. Sarojini & R. Nagabhushanam
(eds), Indian Ocean: Biology of Benthic Marine Organisms.
References A. A. Balkema, Rotterdam: 85-95.
Robertson, A. I., 1991. Plant-animal interactions and the
Chapman, V. J., 1976. Mangrove vegetation. J. Cram, Ger- structure and function of mangrove forest ecosystems. Aust.
many, 447 pp. J. Ecol. 16: 433-443.
Choy, S. C. & W. E. Booth, 1991. Ecology of the 'Api Api' Sasekumar, A., Distribution of macrofauna on a Malayan
(Avicennia marina) wetland around Sungai Pemburon- mangrove shore. J. animo Ecol. 43: 51-69.
gunan, Meragang with proposals for its utilization, conser- Stewart, G. R. & M. Popp, 1987. The ecophysiology of man-
vation and management. Document prepared for the De- groves. In R. M. M. Crawford (ed.), Plant life in aquatic
partment of Town and Country Planning, Ministry of and amphibious habitats. Blackwell Scientific, Palo Alto,
Development, Brunei Darussalam, 25 pp. CA: 333-345.
Chua, T. E., L. M. Chou & M. S. M. Sadorra, 1987. The Tomlinson, P. B., 1986. The botany of mangroves. Cambridge
coastal profile of Brunei Darussalam: resource assessment University Press, Cambridge, 413 pp.
and management issues. ICLARM Techn. Rept. 18, Fish- Zamski, E., 1979. The mode of secondary growth and the
eries Dept., Brunei Darussalam & ICLARM, Philippines, three dimensional structure of the phloem in Avicennia. Bo-
193 pp. tanical Gazette 140: 67-76.
Hydrobiologia 285: 249-255, 1994.
The remnant mangroves of Sei Keeil, Simpang Hilir, West Kalimantan,

Indonesia
Rochadi Abdulhadi & Suhardjono

Herbarium Bogoriense, Research and Development Center for Biology, J1. Juanda 22, Bogor 16122,
Indonesia
Key words: phytosociology, remnant mangrove
Abstract
Phytosociological analysis of the remnant mangrove forest was carried out in Sei Kecil, Telok Melano,
West Kalimantan, to describe the forest composition and structure, and to assess the regeneration
strategy of the important mangrove species. Six transect plots of 20 x 60 m, 20 x 80 m and 20 x 90 m,
with a total area of one hectare were established perpendicular to the coast line.
A total of 20 species of trees and saplings, two species of herbs and one species of nypa palm was
recorded. Three community types were recognized namely Rhizophora apiculata, Excoecaria agallocha-
Rhizophora apiculata and Excoecaria agallocha reflecting the zonation in this forest.
The number of trees, saplings and seedlings were 510, 1220 and 11085 per ha respectively. Rhizophora
apiculata was most successful in regeneration with 6231 seedlings per ha (56.2%) and followed by
Bruguiera parviflora with 3810 seedlings per ha (34.4 %). Seedlings of the first species were mostly growing
under gaps while the second species occurred mostly around the mother trees.
Introduction 980000 ha. Mangrove forests are one of the pri-

mary features of the coast of Kalimantan, how-
Indonesia has the largest area of mangrove forest ever, there have been no studies of the structure
in the world covering about 4.25 million hectares of these tidal forests.
(Silvius, 1989; Soemodihardjo & Soerianegara, The present study describes the structure, com-
1989). They are well developed in the estuaries of position and regeneration of a remnant mangrove
large rivers such as on the east coast of Sumatra, forest in West Kalimantan.
along the coasts of Kalimantan and South
Sulawesi and Irian Jaya.
Unfortunately, the extent of this forest is de- Study area
creasing due to pressure of human activities. Such
activities include the commercial exploitation for This study was conducted in Sei Kecil (10 14' S,
production of logs, wood chips and charcoal that 110 0 18' E), Simpang Hilir, Ketapang, West
is being carried out in Sumatra and Kalimantan. Kalimantan. It lies between Melano Bay and
Silvius (1989) reported that in Kalimantan, Kumbang Island (Fig. 1). The long coast line of
the area of mangrove forest was originally 3 km is covered by mangrove forests. In 1970, the
1801000 ha, but now there remains only landward part of this forest was converted into a
250
Pontianak District
1
Tonj ung sotoi
®
Remnant mangrove or tbe area
• Study Site
Fig I Map of the study site on the west coast of Kalimantan
coconut plantation, and thus the mangrove forests (Q) of 4.6 (Schmidt & Ferguson, 1951). The
were reduced to a width of 60-100 metres all average annual rainfall at the nearest station
along the coast. (Sukadana) is 3738 mm, and it is distributed
The climate is ever wet and belongs to Koppen rather evenly throughout the year. Soil in the area
type A with a ratio between dry and wet months is alluvial (Bakosurtanal, 1983a & b).
251
Methods Indonesia (Kartawinata et al., 1979). Although

this study site is a remnant mangrove forest, the
Six transect plots of 20 x 60, 20 x 80 and number of species recorded is comparable to the
20 x 90 m (with the total area of one ha) were other mangrove forests (Prawiroatmodjo et al.,
established perpendicular to the coast line. Each 1985; Mirmanto et al., 1989; Suhardjono &
transect was divided into 12-18 subplots of Budiman, 1990).
10 x 10 m. A cluster analysis using a Bray-Curtis dissimi-
Every tree (DBH> 10 cm), sapling (DBH larity measure and group average sorting strategy
2-10 cm) and seedling present within each plot (Clifford & Stephenson, 1975) showed three
was identified and recorded. The diameter of trees groups in the sample plots (Fig. 2), reflecting the
and saplings was also measured. The canopy of zonation within the mangrove forest. The three
each tree was mapped to construct the forest pro- zones, extending from the seaward to the land-
file. ward side are characterized by the following three
community types (Fig. 3):
1. Rhizophora apiculata community
Results and discussion
2. Excoecaria agallocha-Rhizophora apiculata
community
Twenty species were recorded within the six sam-
3. Excoecaria agallocha community
pling plots. These consisted of 17 tree species
(Table 1), two species of herbaceous weeds Acan-
thus ilicifolius and Acrostichum aureum and the 0.11"
palm Nypa fruticans.
It can be seen that 14 of 17 tree species (82.3 %)
are true mangrove species, and these represent
0.61"
36.8% of the true mangrove species found in
Table 1. List of species of trees, saplings and seedlings in six

line plots.
Family Species
0.31"
Apocynaceae 1. Cerbera manghas L.
Arecaceae 2. Nypa fruticans Wurmb. --,----
I
J 1
Bignoniaceae 3. Dolichandrone spathaceae (L.f.) Sch.
Combretaceae 4. Lumnitzera littorea (Jack) Vog. o .15"
Euphorbiaceae 5. Excoecaria agallocha L.
Lecythidaceae 6. Barringtonia racemosa (L.) Spreng.
Malvaceae 7. Hibiscus tiliaceus L.
Meliaceae 8. Xylocarpus granatum Koen.
Moraceae 9. Ficus microcarpa L.r.
Myrsinaceae 10. Aegiceras corniculatum (L.) Blanco
Rhizophoraceae 11. Bruguiera gymnorhiza Lamk.
12. Bruguiera parvilfora (Roxb.) W. & A.
ex Griff. o 0" 40 m Crom the seaward
13. Ceriops tagal (Perr.) C. B. Robins
14. Rhizophora apiculata Bl. 00 0 40" 80 m Crom the seaward
Rubiaceae 15. Scyphiphora hydrophylacea Gaertn. • • •• > 80 m Crom the seaward
Sonneratiaceae 16. Sonneratia caseolaris (L.) Engl.
Sterculiaceae 17. Heritiera littoralis Dryand. ex W. Ait Fig. 2. Sample plots grouped according to the species present,
Verbenaceae 18. Avicenia alba Bl. using a Bray-Curtis dissimilarity index and group average
sorting strategy.
252
M 20
C
B
15
!
j 10
~
Seaward
Fig. 3. Profile diagram of a remnant mangrove, showing the community changes from beach toward inland. A. Rhizophora apiculata
community, B. Excoecaria allagocha-R. apiculata community and C. E. allagocha community.
Abbreviations in canopy:
Ra = Rhizophora apiculata; Aa = A vicennia alba
Ll = Lumnitzera littorea; Ea = Excoecaria allagocha
Bp = Bruguiera parviflora; B = Bruguiera gymnorhiza
HI = Hibiscus tiliaceus
Rhizophora apiculata community Rhizophora apiculata. The other species occurring

This community on the seaward fringe had a in the community were Lumnitzera littorea,
width of 40 metres. The substrate consisted of Aegiceras corniculatum, A vicennia alba and
sandy mud with a pH of 5.5 ± 0.3. The commu- Excoecaria agallocha (Table 2).
nity was dominated by pure stands (90%) of This community was still young, with a homog-
Table 2. Number of indo ha - I (0), basal area ha - I (BA) and importance value (IV) of trees and saplings and seedlings in
Rhizophora apiculata community.
Species Trees Saplings Seedlings
0 BA IV 0 BA IV 0
Rhizophora apiculata 441 13104 238.95 868 1780 247.24 4695

Lumnitzera littorea 18 3008 24.30 25 18 7.48 36
Aegiceras corniculatum 10 564 13.42 40 67 13.85
A vicennia alba 10 437 9.50 42 77 17.03 869
Excoecaria agallocha 10 460 9.18
Heritiera littoralis 2 26 2.34 4 16 4.47
Bruguiera parviflora 2 40 2.35 4 6 3.94
Ceriops tagal 4 3 1.95
Scyphyphora hydrophyllacea 4 7 2.17
Xylocarpus granatum 2 I 1.87 8
Sonneratia caseolaris 2
Total 493 17639 300 993 1975 300 5610
253
enous structure. The density was very high, but bigger, with an average diameter of 32 cm. The
the basal area was relatively low. Trees were gen- big trees were mostly Lumnitzera littorea and
erally small with an average diameter of 11 cm. Excoecaria agallocha with average diameters of
Trees of 6 m height formed one layer (Fig. 3). 64 cm and 32 cm respectively. The largest tree
Saplings were also dense and composed mostly was Lumnitzera littorea with a diameter of75 cm.
of Rhizophora apiculata (87.4 %) and A vicennia
alba (4.2%). There was little undergrowth except Excoecaria agallocha community
for some seedlings of tree species of Rhizophora This community develops on better drained and
apiculata, A vicennia alba and Lumnitzera littorea. firm soils. The soil pH was low i.e., 4.3 0.6. Ex-
coecaria agallocha was the most dominant species
Excoecaria agallocha-Rhizophora apiculata with the highest importance value (Table 4). Its
community density was 70.2 % of the total density and the
The Excoecaria agallocha-Rhizophora apiculata basal area was 78.6% of the total basal area within
community type occurred behind the Rhizophora this community.
apiculata community. It comprised a community At least two canopy layers were recognized
with a more heterogenous floristic composition (Fig. 3). The top layer was dominated by Ex-
(Table 3), in which 10 species of trees and 14 coecaria agallocha which reached a height of 18 m.
species of saplings and seedlings were recorded. The second layer consisted of Bruguiera parvi-
The two dominant species were Excoecaria flora. Rhizophora apiculata, Heritiera littoralis and
agallocha and Rhizophora apiculata, with impor- Ficus microcarpa.
tance values of 108.6 and 92.2 respectively. The
density of this community was slightly lower than Regeneration of important mangrove species
that ofthe Rhizophora community. The basal area,
however, was more than twice that of the Rhizo- Regeneration in the three forest communities was
phora community. Trees in this community were good. The mean density of trees, saplings and
Table 3. Number of indo ha - 1 (0), basal area ha - 1 (BA) and importance value (IV) of trees and saplings and seedlings in
Excoecaria agallocha-Rhizophora apiculata community.
0 BA IV 0 BA IV 0
Excoecaria agallocha 191 15973 108.62 220 329 37.32 187

Lumnitzera lit/orea 42 13485 67.15 138 155 24.31 286
Bruguiera parviflora 9 851 7.66 96 110 20.33 1632
Scyphyphora hydrophyllacea 9 177 5.09 24 38 7.49
A vicennia alba 7 266 4.84 8 4 1.44 59
Aegiceras comiculatum 7 88 4.42 101 48 16.49 79
Heritiera lit/oralis 9 121 4.10 37 147 12.08 35
Dolichandrone spathacea 4 208 2.36
Hibiscus tilleaceus 4 86 2.06 13 18 3.55 7
Cerbera manghas 2 11 1.54
Bruguiera hymnorhiza 2 2 1.20
Xylocarpus granatum 2 1.16 121
Ceriops tagal 2 1.15 22
Nypa fruticans 195
Total 471 38384 300 1594 2606 300 11922
254
Table 4. Number of indo ha - I (D), basal area ha - I (BA) and importance value (IV) of trees and saplings and seedlings in
Excoecaria agallocha community.
D BA IV D BA IV D
Excoecaria agallocha 412 37227 182.10 400 539 77.64 212

Bruguiera parviflora 50 2199 35.37 250 228 45.59 9800
Heritiera littoralis 25 590 19.33 38 121 23.95
Ficus mucrocarpa 25 2830 21.37 62
A vicennia alba 12 2707 13.40 12 16 7.94 87
Aegiceras corniculatum 12 11 7.59
Xylocarpus granatum 12 5 7.18
Cerbera manghas 75 178 38.45
Nypa fruticans 775
Dolichandrone spathacea 88
Total 563 46694 300 1074 1447 300 15724
seedlings were 510, 1220 and 11 085 per ha re- parviflora with the mean number of seedlings of
spectively. This probably indicates that if there is 6936 (72.76%) and 1531 (16.06%) per ha respec-
no interference from human activities, this rem- tively. Seedlings of Rhizophora apiculata are com-
nant forest will be able to maintain itself. mon (F = 93 %); tending to favour the gaps in the
The most successful species in terms of regen- forest. Figure 4 shows that the percentage Foliage
eration were Rhizophora apiculata and Bruguiera Projective Cover (FPC) influenced the numbers
450
.... ..
e
-
0
300 .. ..
0
~
II)
en
.5 r = - 0.885
:0
c:.I
c:.I
(I) 150 " It
0:5
Z
"It , .. 1flI1t
It
'r '!I'>lliflrl ..
",.11.
a
a 30 45 60 75 '0 105
X
FPC
Fig. 4. Relationship between the percentage Foliage Projective Cover (FPC) and numbers of R. apiculata seedlings.
255
of seedlings. The number of seedlings decreased M. Tantra, 1979. Status pengetahuan hutan bakau di
Indonesia. In: Soemodihardjo, S. et al. (ed.), Pro siding
with an increase in the percentage of Foliage
Seminar I Ekosistem Mangrove, Panitia Program MAB
Cover (r = - 0.885); seedlings being densest Indonesia-LIPI, Jakarta: 21-37.
under the gaps. In contrast, seedlings of Bruguiera Mirmanto, E., K. Kartawinata & A. Suriadarma, 1989. Man-
parviflora were not evenly distributed (F - 17 %), grove and associated plant communities in the Barito
but clumped around the mother trees. River estuary and its vicinity, South Kalimantan. Eko!.
Indonesia I: 42-55.
Prawiroatmodjo, S., D. Sapulete, S. F. Pratignyo & A. Budi-
man, 1985. Structural analysis of mangrove vegetation
Conclusion in Elpaputih and Wailale, Ceram, Indonesia. In K. N.
Bardsley, J. D. S. Davie & C. D. Woodroffe (eds), Coasts
The remnant mangrove forest at Telok Melano, and tidal wetlands of the Australian monsoon region: 153-
Kalimantan contains a large number of mangrove 165.
Schmidt, F. H. & J. H. A. Ferguson, 1951. Rainfall types
species which showed good regenerative ability. based on wet and dry period ratios for Indonesia with
The functional importance of this mangrove forest western New Guinea. Kementerian Perhubungan, Djawa-
to the coastal environment should be recognised tan Meteorologi dan Geofisika, Jakarta. Verhandelingen,
and efforts should be made to conserve this rem- No. 42.
nant forest. Silvius, M. J., (1989). Indonesia. In D. A. Scot (ed.), A Di-
rectory of Asian Wetlands, IUCN, Gland, Switzerland:
981-1109.
Soemodihardjo, S. & I. Soerinagera, 1989. The status of man-
References grove forests in Indonesia. In I. Soerianegara, P M.
Zamora, K. Kartawinata, R. C. Umaly, S. Tjitrosoma,
Bakosurtanal, 1983a. Peta Tanah Eksplorasi: Atlas sumber- D. Sitompul & U. R. D. Syafii (eds), Symposium on Man-
daya regional Kalimantan Tengah. Lembar B. 2392. grove Management: its Ecological and Economic Consid-
Bakosurtanal, 1983b. Geologi: Atlas sumber daya regional erations. BIOTROP Special Pub!. No. 37: 73-114.
Kalimantan. Lembar B 0913. Suhardjono & A. Budiman, 1990. Structure of mangrove for-
Clifford, H. T. & W. Stephenson, 1975. An introduction to est at Tabobo, Halmahera, Indonesia. Presented Interna-
numerical classification. Academic Press, London. tional Congress of Ecology Yokohama, 23-30 August 1990,
Kartawinata, K., S. Adisoemarto, S. Soemodihardjo & I. G. 13 pp.
Hydrobiologia 285: 257-270, 1994.
Marine living resources management in the ASEAN region: lessons

learned and the integrated management approach
Chua Thia-Eng & Len R. Garces

International Center for Living Aquatic Resources Management, M.C.P.O. Box 2631, Makati, Metro
Manila, Philippines
Key words: resources management, ASEAN region
Abstract
This paper reviews the marine living resources management practices in the ASEAN region. Lessons
learned from past management of these resources are highlighted with an analysis of impacts. The lack
of an appropriate management policy at national and regional levels, inefficient enforcement measures,
the rapid population growth and unsustainable economic development have all contributed to the deg-
radation of the resource base in the marine environment of many ASEAN members.
The need to establish a holistic and integrated management approach as well as action plans to mitigate
the accelerated deterioration of the marine environmental quality and to promote sustainable use of
resources is strongly emphasized in this paper. Recent changes in government policies in ASEAN to-
wards sustainable development could provide valuable opportunities to arrest further resource depletion.
The ASEAN initiation of integrated management of marine resources is a step towards the right
direction.
Introduction 1940s and the 1960s), national policies in most

Southeast Asian nations were large-scale exploi-
The marine living resource base in the Associa- tation of primary resources, self-sufficiency in
tion of Southeast Asian Nations (ASEANl) re- food production and employment opportunity in
gion has undergone drastic changes over the last rural and urban areas. In the 1960s, an improved
50 years. After World War II, many ASEAN communication system was followed by a rapid
members embarked on very ambitious economic technological transfer that accelerated the exploi-
programs to reconstruct the war-torn economy tation of primary resources; for example, fishing
and to meet the demands for food and employ- became more efficient with the conversion of non-
ment. The richness of the primary resources, in- powered to powered fishing vessels, and the use
cluding the marine living resources, therefore be- of synthetic nettings and fishing gear such as
came the target of exploitation by these nations to trawls and purse seines. Promoted by the influx
supply raw material for foreign exchange. Hence, of capital into the region from bilateral and mul-
during the first 15 years after the war (between the tinational sources, fish production increased con-
siderably in a decade, resulting in an accelerated
I The term ASEAN includes Brunei Darussalam, Indonesia, upward trend in fish production since the 1960s
Malaysia, Philippines, Singapore and Thailand. (Pauly & Chua, 1988). The human population in
258
Southeast Asia also increased twofold over the This paper examines how these resources are
last half century of which a large percentage is utilized and managed in the marine environment
concentrated along the coast (Chua, 1991). in the ASEAN region, including useful manage-
Towards the 1980s, rapid industrialization in ment lessons learned under different cultural, eco-
some nations was towards marine transportation, nomic and sociopolitical conditions.
oil exploration, manufacturing and service. These
caused pollution in the form of industrial, agri- Marine living resources and their socioeconomic
cultural and domestic waste discharges which contributions
enter the river systems and coastal waters, thus
impairing the life-support systems in the marine The marine living resources that are of socioeco-
environment (Chua et al., 1989). nomic significance include fisheries (finfishes,
Realizing the non sustain able consequences of crustaceans, molluscs and seaweeds), coral reefs
economic development, ASEAN members began and seagrasses. Lesser resources include birds,
to take stock of their renewable resources, de- reptiles and mammals. The ASEAN region has a
velop regulatory measures to curb indiscriminate total coastline estimated at 85504 km (Table 1)
destruction of the environment and to initiate re- and its coastal zone is endowed with some of the
medial actions to protect and conserve the frag- world's richest habitats/ecosystems in terms of
ile ecosystems. biodiversity and productivity (Fig. 1). Roughly
The 1990s see an increasing environmental 30% of the world's coral reefs are found in the
awareness of the consequences of improperly ASEAN region (McManus, 1988), also about
planned, poorly managed economic development. one-quarter (26 %) of the world's total mangroves
National governments in the region have begun to (Chua, 1991).
include environmental concerns in formulating Among the living resources, the socioeconomic
economic development policies. This is evident contributions of fisheries are well-documented
from the establishment of the Ministry/Depart- and have received wide attention. Those of other
ment of Environment or related agencies in all resources such as corals, seagrasses and man-
ASEAN members. This may have come a little groves are comparatively less known.
bit too late as many of the renewable resources in Of the world marine fish production of 84.6
the region are already overexploited. On the other million mt, ASEAN members produced around
hand, it poses new challenges to improve the 6.8 million mt in 1988 (FAO Fish. Stat., 1988).
quality of the marine environment through greater Available information on the direct contribution
efforts in management, rehabilitation and conser- of fisheries to GOP in the region varies from
vation. 0.2% to 5% (Table 1). It is insignificant when
Table 1. Selected statistics relevant to fisheries in the ASEAN region.
Country Land area Coastline Population Population Per capita Marine Per capita Contribution
(km 2)8.< length (millions) growth rate GNP (US$) landing (mt) consumption of of fishing
(km)8 19908 .< (%) 1986-898 .< 1989< 1988b •d fish (kg) 1990c to GDP (%)d
Brunei Darussalam 5.8 161 0.25 nd nd 1,548 31.0 0.2 (1988)

Indonesia 1,904.6 54,716 182.65 2.1 490 2,169,557 14.0 1.9 (1988)
Malaysia 329.8 4,675 17.81 2.6 2,130 823,240 30.1 0.2 (1988)
Philippines 300.0 22,540 61.50 2.4 700 1,438,361 33.8 5.0 (1989)
Singapore 0.6 193 2.73 1.3 10,450 13,151 34.0 nil
Thailand 514.6 3,219 56.34 1.7 1,160 2,337,216 20.8 1.7 (1988)
TOTAL 85,504 321.28 6,783,073
Sources: a WRI (1990); b FAO (1990); c Infofish Trade News (1991); d SEAFDEC (1990);" ADB statistics.
Note: nd = no data/information; mt = metric tons.
,-:..r,",,\ ~
/~ " (,. .. > 'J
(' J <,~ , Probob&e erGO for nesting, feed ing
(.,..,.. .... .. ,.1.. - ..\ "'t,
~
mel $ho.-! m;gralion of seo lurtles
\
~ Known nesting sites of sea turtles
i.\ ...
-~
~ G Record of seogrOS5es
{
~ Dugong $i9hlings
. . t~f ·· . . ·
SOUlh China S80 ~ Co.-ol "",Is
OJ Mangrove forests
'.i/ ~
;
@ Andomon Sea
II;
C' ~$
I/;
'9
I) Pacific
~
~ o;
c;I:.
OC8on
@J I
5·
t
II
Indian OCtIOn
N "'"
o 200 400
, 600 800 1,000 km ~dt'
95· 100· 105· 110· t 15 0 IZO· 125'
FIg 1 DlstnbutlOn of mangroves, coral reefs, seagrasses and other manne hvmg resources m Southeast ASIa (ModIfied from Morgan & ValenCIa, 1983) tv
VI
\0
260
compared to nonrenewable resources such as oil lowlands and estuaries, the mangrove swamps
and marine mineral deposits or related activities provide natural protection against coastal ero-
such as shipping, port and harbor services and sion, contribute organic matter and nutrients to
coastal tourism. However, its contribution to em- the primary and secondary productivity of the
ployment opportunity has been very substantial. coastal ecosystems, supply forest products and
In 1978, close to 3 million people in Southeast serve as nursery ground for fish and sanctuary
Asia depended on fishing for livelihood (Josupeit, for wildlife (Paw & Chua, 1991). The estimated
1981). For example, Philippine fisheries produc- areal extent of mangroves in the ASEAN region
tion comprises about 5% of the GNP and em- is 5335 million ha or 26% of the world's man-
ploys over 1 million people or 5 % of the national groves (Chua, 1991). Considered as a wasteland
labor force (ACI, 1989). Moreover, almost all in many developing countries in Southeast Asia
the ASEAN members depend on fish as a main prior to the 1950s, mangroves are now valued
source of protein with a per capita annual at US$ 160-530 ha - I y- I (Dixon, 1989). In
consumption that ranges from 15 kg to 35 kg Malaysia, around US$ 604-8067 ha - I Y- I are
(Table 1), which is among the world's highest derived from charcoal, poles and firewood (Paw
(INFOFISH, 1991). & Chua, 1990).
Coral reefs have the highest primary produc- Like coral reefs which serve as nursery ground
tivity of any coastal ecosystem, contributing about and support a substantial fishery, seagrasses when
10-15 % of the total fisheries catch annually (Mc- found in abundance on the muddy bottom serve
Manus, 1988; Chua, 1991). In the Philippines, for as ideal nursery beds for young fishes, inverte-
example, the average harvest in coral reefs was brates, reptiles and mammals. They are also an
around 4-20 mt km - 2 Y- I (Munro & Williams, important food source of dugongs and green sea
1985). Reefs also play an important role in main- turtles (Fortes, 1989).
taining coastline stability by acting as an efficient The indirect contribution of marine living re-
physical barrier to tidal waves (White, 1987). sources to the development of secondary and ter-
Coral reefs are considered the 'rainforests' of the tiary industries is also substantial but difficult to
tropical seas. The contribution of coral reefs to quantify. In many ASEAN members, these re-
the pharmaceutical industry could be even greater sources also support the livelihood of a substan-
when knowledge and advanced technology are tial number of women and children gleaning over
available to exploit their potential fully. In addi- coral reefs, sandy and muddy beaches for shell-
tion, the aesthetic appeal, biological richness, fish, seaweeds and shells.
clear waters and relative accessibility of coral reefs Table 2 presents some of the socioeconomic
make them popular recreation areas for local and contributions of the resources, namely, food,
foreign tourists (White, 1987). tourism, foreign exchange earning and natural
Located in the intertidal zones and coastal products (e.g., timber).
Table 2. Socioeconomic contributions of marine living resources in the ASEAN region.
Contributions Fisheries Coral reefs Mangroves Seagrasses
1. Food
• • o o
2. Employment • * *
3. Coastline protection
• •o o
4. Tourism
• o
5. Timber/wood •o
6. Foreign exchange earning • o
Note: • = Very significant; 0 = Moderately significant; * = Significant; - = Not significant.

261
Exploitation, management and lessons learned intensified tin mining at Ranong, Phangnga and
Phuket provinces has led to heavy deposits of
Exploitation sediments to nearby mangrove habitats (Ak-
sornkoae, 1986). In Singapore, large tracts of
Over the last five decades, almost all renewable mangroves have been transformed into ponds for
marine resources in the ASEAN region have been shrimp cultivation and fattening of mud crabs. In
subjected to rampant exploitation and subsequent recent years, however, this activity has declined in
destruction by unregulated human activities for favor of reclamation for housing and industry
short-term economic gains (Pauly & Chua, 1988). (Corlett, 1986).
Almost 90 % of the fisheries resources in the re- The greatest contributors to reef stress are
gion come from the shallow continental shelf and human activities which include siltation, destruc-
coral reefs in the coastal zone (Fig. 2). Fishing is tive fishing methods, coral mining, tourism re-
most intense in the coastal waters within 7 miles lated activities and pollution (Table 3) (Gomez,
from the coast where majority of the artisanal 1988).
fishermen operate. Serious overfishing problems
occur in the coastal waters where trawls, purse
seines and other mechanical fishing gear are used. Management
Because of the development of these fisheries,
demersal fish resources in nearshore waters are The continued depletion of the marine resources
believed to be overexploited, and many pelagic and degradation of ecosystems suggest that the
fish resources in shallower waters are close to natural renewable resources in the ASEAN re-
overexploitation in recent years (Pauly & Chua, gion are inadequately managed. Table 4 shows
1988; Pauly et aI., 1989; Pauly, 1989). some of the management schemes that have been
With the growing population and increased implemented in the region to conserve or regulate
human activities in coastal areas, some critical the use of these resources.
habitats such as mangroves, coral reefs and sea- Fisheries constitute the largest and most im-
grass beds have also been heavily disturbed, caus- portant renewable resources in Southeast Asia.
ing rapid resource degradation or destruction The major management issues in capture fisher-
(Fortes, 1988; Gomez, 1988; McManus, 1988). ies are:
The decline in the areal extent of mangroves could
1. Too many fishermen. Fishing is a traditional
be attributed to clearfelling and conversion for
practice and fisheries resources are common
aquaculture or agriculture, tin mining and recla-
property resources, access to which is less restric-
mation for urban and industrial developments
tive; thus, many coastal waters are overcrowded
(Aksornkoae, 1986; Corlett, 1986; Primavera,
with artisanal fishermen. Fishing has become a
1991; Rao, 1986; Soemodiharjo, 1986).
convenient occupation for those who cannot find
Philippine mangroves have declined from
employment on land.
400000 ha in the 1920s to only 140000 ha at
present (Primavera, 1991). Sixty percent of this 2. Resource depletion. There is simply not
decrease is due to conversion into culture ponds enough fish available for the large fishing popu-
for milkfish and shrimps (Primavera, 1991). Simi- lation. This is worsened by the destruction of
larly, mangroves are also being converted to marine nursery grounds and coastal habitats.
brackishwater fishponds ('tambaks'), particularly
3. Loss of productivity. This is due to sacrifice of
for cultivating shrimp in Indonesia. The Five-
mangroves for 'tambak' (fishpond) development
Year (1984-1989) Development Plan ofIndone-
without taking mitigating measures.
sia marked the opening of thousands of square
kilometers of mangroves for conversion to fish- 4. Overcapitalization. In certain countries, espe-
ponds (Soemodihardjo, 1986). In Thailand, cially Thailand, Malaysia and the Philippines,
IV
0'\
IV
OemerlOI fishing grounds (pr... nt and pOllnticl)

Shollow-woter trowling, 'fish trepping ond
rrrrm oll"ter shallo..... -wafer demersol fisheries
Deep- water trawling t vertical longlining ond
~ other deep-water demersal fisheries
-'! "tI Pelagic fi5hing grounds (pro •• nt and potent ial)
'" . eP Shallow-weier purse sein ing. drift gillnetling and
~ other sha llow-woter pelagic fisherIes
Deep-waler purse 'Seining, pelagic longHning and
D other deep-water pelcQic f isheries
5°
0°
5°
N
j
o 200 400 600 800 1,000 km
95 0 100' 105 0 110' 11 5 0 120' 125 0 130' 135 0 1400 145'
Fig 2 PelagIc and demersal fishmg grounds m Southeast ASIa (ModIfied from Morgan & ValencIa, 1983)
263
Table 3. Human activities that contribute to destruction of marine resources in the ASEAN region.
Living resources and Brunei Indonesia Malaysia Philippines Singopore Thailand

human activities Darussalam
Fisheries
Overfishing * * *
* *
Deleterious fishing methods
* * * *
Environmental degradation * *
* * *
Coastal development
* * * * *
Coral reefs
Coastal development
* * * *
Sewage pollution
Agricultural pollution
* * * *
* * * *
Coral and sand mining * * * *
Overexploitation
Deleterious fishing methods
* * * *
* * * *
Intensive recreational use
* * * * *
Mangroves
Conversion for agriculture
* * * *
Conversion for aquaculture *
* * * *
Mining
* * *
Urbanization *
* * * * *
Conversion to salt ponds
*
Timber production
* * * *
Seagrasses
Industrial development *
* * *
Urban development
* * * *
Mining
*
* = Significant
- =Not significant.
Sources: Yap and Gomez (1985); Aksomkoae (1986); Corlett (1986); Rao (1986); Soemodiharjo (1986); Pauly and Chua (1988)
and UNEP/IUCN (1988).
commercial fisheries, especially the trawl fisher- have now earned better public understanding and
ies, are overcapitalized resulting in competition recognition. The establishment of marine parks/
for high-priced fish in nearshore waters in order reserves is a common management strategy to
to sustain profitability, thus creating resource use conserve or protect coral reefs in Southeast Asian
conflicts with inshore fishermen. countries (White, 1988). At present, there are
around 13 in Indonesia, 4 in Malaysia, 9 in the
5. Inadequate law enforcement. This is a com-
Philippines and 4 in Thailand (White, 1983).
mon problem in the region due partly to the lack
However, their effectiveness is often in question
of political will and financial and manpower re-
(Gomez & Yap, 1982). Local communities gen-
sources for law enforcement.
erally do not have jurisdiction over marine areas,
Over the last 50 years, almost none of the above and coral exploitation and deterioration are still
issues has been satisfactorily resolved. They still on the rise (White, 1988).
remain the predominant features of the coastal Other forms of reef management and conser-
fishing industry. vation, sometimes within the definition of marine
In the case of coral reef management, the situ- reserve include: (1) controls on fishing methods
ation is not any better. But the ecological function and rates of exploitation; (2) involvement and or-
and socioeconomic value of coral reef resources ganization of local fishing communities to man-
264
Table 4. Management of marine living resources in the ASEAN region.
Living resources and Brunei Indonesia Malaysia Philippines Singapore Thailand

management strategies Darussalam
Fisheries
Licenses
* * * * * *
Area closures
* *
Gear restrictions
* * * * * *
Coastal zone management planning 0 0 0 0 0 0
Coral reefs
Protected areas/marine parks 0
* * * *
Laws prohibiting use of destructive
fishing methods
* * * * *
Part of military reserve
*
Mangroves
Forest reserve
* * * * * *
Zonation/land use planning * *
* * *
Rehabilitation, reforestation
* * * * *
Aforestation *
* * *
National mangrove management plan *
0 0
* *
EIA for aquaculture development
*
Seagrasses
Rehabilitation * 0
National seagrass management program *
* *
* = Practiced.
o = Initiating.
- = Not practiced.
EIA - environmental impact assessment.
Sources: Yap and Gomez (1985); Aksornkoae (1986); Corlett (1986); Rao (1986); Soemodiharjo (1986); Pauly and Chua (1988)
and UNEP/IUCN (1988).
age their own reef resources; (3) education of the the rate and level of exploitation were not high.
coastal population; (4) municipal, provincial and This situation has changed, however, and a more
national legislation; and (5) various integrated holistic and systematic management of the man-
approaches to coastal resources management by grove forests is needed to reduce the present rate
national agencies (White, 1987). of depletion.
Moreover, the collection, sale and export of Various tools used in managing mangroves in-
corals are now banned in the Philippines reflect- clude setting up of management, conservation,
ing government resolve to protect and conserve conversion and/or reconstruction zones; integra-
the remaining reef resources (Gomez, 1989). An tion of forestry and fishery production; and use of
educational campaign was also launched in Thai- mangrove forests for agriculture and habitation
land to reduce the rate of coral destruction by (Aksornkoae, 1989).
tourists and local inhabitants. A comprehensive Perhaps, Malaysia has the most effectively
national program to conserve these resources has managed mangrove reserve in the Matang Forest
not yet been developed, although specific local- Reserve (Darus & Haron, 1989), but its success
ized management plans have been formulated has not been repeated in other parts of the coun-
(e.g., Coral Reef Management Plan for Phuket try or in other ASEAN members. Economic ben-
Island, Thailand). efits and political considerations have often led to
Traditional mangrove management for multiple the sacrifice of mangrove ecosystems. While the
resource use was effective prior to the 1970s when United Nations Educational, Scientific and
265
Cultural Organization (UNESCO) has been pro- and socioeconomic conditions within and among
moting sound management strategies (Vannucci, the nations. Here are some recommendations for
1988) and forming the National Mangrove Advi- a sound management of resources:
sory Committee, effective mangrove management
1. National policy on natural resources manage-
has yet to be demonstrated. Notwithstanding this,
ment. The lack of policy guidelines on the use and
the concerted efforts of scientists, conservation-
management of common property resources is
ists and environment-conscious organizations at
one of the major reasons for the resource
national and international levels have sensitized
managers' inability to effectively reduce the im-
large numbers of people at decisionmaking and
pact of environmental and resource degradation.
community levels to the urgent need for mangrove
Although the concept of sustainable development
management. The use of mangroves is more regu-
is widely accepted by policymakers in the region,
lated now.
it has not yet been translated into applicable ac-
Despite efforts through international and re-
tion programs that merit national interest and
gional conventions, e.g., Stockholm Conference
government support. A comprehensive national
(1971); London Dumping Convention (1972);
policy on natural resource management is there-
Law of the Sea Conference (1982); Montreal Pro-
fore essential.
tocol on Prevention of Land-based Pollution
(1985), serious management measures to prevent 2. Local government involvement. In many
and regulate massive pollution in some Southeast countries in Southeast Asia, the local government
Asian nations took place only in the last one to implements programs for the development and
two decades. These occurred after these countries management of marine resources. Therefore, the
established environmental departments, minis- early involvement and participation of local au-
tries or related agencies (e.g., National Environ- thorities in plan formulation and implementation
ment Board, Thailand). National regulatory mea- are crucial.
sures were formulated especially for land-based
pollution. However, with the exception of Sin- 3. Support from policymakers and community.
gapore and to a certain extent, Malaysia, effective Any program or plan designed to resolve the
enforcement has not been successful. This led to management problems of fisheries or other sec-
the worsening pollution problem in most coastal tors related to marine resources will need the full
waters in the ASEAN region. Some rivers and support and understanding of policymakers at
estuaries are already declared 'biologically dead' central, state/provincial and local levels as well as
by authorities because of organic and industrial the concerned coastal community. Their partici-
pollutants. Poor water quality does not only pose pation will be most useful in the pllm formulation
serious public health risks but also impairs the and implementation stage. Community consulta-
chemical processes that sustain marine and tion and participation should receive special at-
coastal ecosystems. tention as their livelihood may be greatly affected.
4. Multisectoral and interagency participation.
The involvement of different interest groups and
Lessons learned
concerned government and nongovernment agen-
cies is essential to generating greater cooperation
Many useful lessons could be learned from the
and better interagency linkage and collaboration
successes and failures of past and present man-
for resolving resource-use conflicts.
agement practices. Managing living resources in
the marine environment of developing nations in 5. Regulatory measures and public awareness.
Southeast Asia is a very difficult task, consider- Regulatory measures are effective and meaningful
ing the immense human pressure on the resources if fully enforced. To help reinforce implementa-
and the great differences in the political, cultural tion, efforts to inform the public of the issues are
266
needed. Community support for the protection ship of key elements in the environment (e.g.,
of the aquatic environment is also important. human population, physical processes, resources
A community can do a great deal if adequately and ecosystems) and their impacts. Research on
educated and organized. these areas, therefore, is important to solve the
problems in the marine environment. As shown
6. Alternative livelihood. Many of the manage-
in Fig. 3, planning and use of marine resources
ment issues of fisheries and other resources can-
range from local communities to national and in-
not be resolved by regulatory measures alone. In-
ternational levels, which require integration to
stead, incentives through the establishment of
avoid conflict and prevent degradation and loss
alternative livelihood are absolutely necessary
of sensitive ecosystems and habitat.
particularly in siphoning out excess fishermen to
Figure 4 illustrates the conceptual framework
reduce fishing effort or in preventing the use of
of integrated area management. The processes
destructive fishing gear.
are: (1) baseline data gathering, analysis and im-
pact assessment; (2) integration and decision-
making; and (3) generation of output.
The integrated approach to the management of Baseline data gathering provides information
marine resources on the environment of a study area. This includes
biogeophysical, sociocultural, economic and in-
The sectoral management of fisheries and other stitutional data including sectoral indicators (e.g.,
resources is inadequate to cope with these prob- optimum sustainable yield, target production).
lems: the increasing number of fishermen vis-a.-vis Data gathering covers the consolidation of sec-
dwindling resources, poverty, coastal pollution ondary information; and where data gaps exist,
and habitat degradation, inefficient law enforce- the generation of primary data. Also, updating
ment and existing institutional constraints, in ad- and/or verification is necessary to ensure reliabil-
dition to pressures of economic development and ity and viability of information required for deci-
human population. A holistic integrated manage- sionmaking.
ment program is necessary to address these. It Collected information is then analyzed for
will define natural resource allocation, develop trends and relationships among key elements in
policy reform and formulate management strate- the study area. Impact assessment, on the other
gies that guarantee sustainability, economic hand, provides the interactions of sectoral activi-
growth and protection of the environment. ties in the environment. The benefits and adverse
Therefore, management of fisheries, aquacul- impacts on the environment as well as conflicts
ture, mangrov~s, coral reefs and other resources, with other sectors can be identified in this pro-
will be more successfully implemented within the cess.
general framework of integrated coastal zone The integration and decisionmaking phase
management. A multi sectoral and interagency ap- evaluates all the information to come up with
proach to resource and environmental manage- mitigating measures. It involves expert decision
ment can be adopted with adequate action pro- and/or multivariate modeling to determine poli-
grams to address the various management issues. cies, strategies and actions needed. This can be
Institutional strengthening is thus programmed articulated through the formulation of an inte-
for the effective enforcement of regulatory mea- grated management plan for the study area.
sures. A feedback mechanism may be necessary to
Integrated management requires an interlink of monitor, validate and reassess the changes that
resource system research, policy analysis and so- will occur upon plan implementation. This leads
cioeconomic valuation. Thus, the success of this to a more practical and acceptable plan. Many
approach rests on the availability of baseline in- ASEAN policymakers, resource managers, sci-
formation and the understanding of the relation- entists and the public have been made more aware
267
TerrUaJ1a l Less Lessl

centrol exlfnslv& 90~emmenl
e'2 bylh.
coastal
col'llrallind
Ill' moO!
o naUcn conkol by ""
()
plf.." sec10,
- - - - - - - - - - - - - - - - ---'f--
Nallenal
Reglon.'
Intll!'rnallonat
• POri aoo
harbOur
• Aquacullure
• Coaslal
tourism
• Transponanon
- ---- ~
Tn, Inter·
Coa.tal "'I,nd or
uplandS nol'\CoaSlaJ
1 - 0 - - - - Conlln,n,,, ... r~ln - - - ----4-1
CS - The coastal basehne is a sene. 01 st,arght hnes Ihat .nlerconnect coastat headlands and promontones The CS IS the reference po.nt used to map
the oceanward boundary of both the temtona! sea and the exclusIVe econormc 20ne
Mod,f.ed lrom Sorensen and McCreary (1990) and Elder and Pemelta (1991)
Fzg 3 Extent of habItats/resources, econOffilC activIties, types of management and government control III the management of
manne and coastal resources and envIronments (after Chua, 1992)
tv
Steps 0\
00
1. Baseline data
gathering and
Impact assessment
Causes
• Policy failure
• Management failure
• Oalagaps
Sectoral activiUes Indicators Benefits Conllicts

(Optimum sustainable yield, (Human welfare) (waler resource a1localion,
target production) land use, environmental degradation)
• Political, sociocultural
and economic consideration
2. Integration and - nalional goal
declslonmaklng - employment
- foreign exchange
-food
• Carrying capacltyl
environmental capacity
• Environmental considerations
- Ecological cost
- Social cost
Output
Management
• Po§cy
3. Output generation plans
• Stralegy
• Action
Fig. 4. The conceptual framework of integrated area management in evaluating and optimizing multi-resource, multi-sector activities.
269
of the urgent need for a collective national and Proc. 30. Department of Science and Technology; Depart-
ment of Agriculture; Department of Environment and
regional effort in the integrated management of
Natural Resources; and Department of Tourism, Philip-
the marine environment. This view has been up- pines; and International Center for Living Aquatic Re-
held over the last few years by different countries sources Management, Philippines: 21-35.
in the region, as reflected in the Langkawi Dec- Chua, T.-E., 1992. Coastal area planning and management in
laration on the Environment (1989), Kuala Lum- developing countries: a synthesis of ASEAN experience.
Paper presented at the Regional Workshop on Integrated
pur Accord on Environment and Development
Coastal Zone Planning and Management in ASEAN: Les-
(1990), Baguio Resolution on Coastal Resources sons Learned. 28-30 April, 1992. Bandar Seri Begawan,
Management (1990), and Singapore Resolution Brunei Darussalam.
on Waste Management (1991). These agreements Chua, T.-E., l. N. Paw & F. Y. Guarin, 1989. The environ-
also show the increasing commitment of national mental impact of aquaculture and the effects of pollution on
leadership to the management of terrestrial and coastal aquaculture development in Southeast Asia. Mar.
Pollut. Bull. 20: 335-343.
marine resources to achieve the goals of sustain- Corlett, R., 1986. Singapore. In Mangroves of Asia and the
able development. Pacific: status and management. Technical Report of the
However, experiences in integrated coastal UNDP/UNESCO Research and Training Pilot Pro-
zone management in developing nations are still gramme on Mangrove Ecosystems in Asia and the Pacific
very limited. The six ASEAN member-nations (RAS/79/002): 211-218.
Dixon, l. A., 1989. Valuation of mangroves. Trop. Coast.
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grated management plans which are being con- ment of Matang mangrove forest reserves in Peninsular
sidered for implementation by their respective Malaysia. In T.-E. Chua & D. Pauly (eds), Coastal area
governments (Chua, 1991). management in Southeast Asia: policies, management,
strategies and case studies. ICLARM Conf. Proc. 19. Min-
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84.
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FAO (Food and Agriculture Organization of the United Na-
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Fortes, M. D., 1989. Seagrasses: a resource unknown in the
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the Pacific: status and management. Technical Report of ASEAN region. ICLARM Educ. Ser. 5. International Cen-
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gramme on Mangrove Ecosystems in Asia and the Pacific Philippines, 46 pp.
Gomez, E. D., 1988. Overview of environmental problems in
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Asian nations. In T.-E. Chua & D. Pauly (eds), Coastal ASEAN/US Coastal Resources Management Project. In
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Hydrobiologia 285: 271-276, 1994.
A. Sasekumar. N. Marshall & D. J. Macintosh (eds). Ecology and Conservation of Southeast Asian Marine and 271
Degradation of mangrove forests in South Sulawesi, Indonesia
Baharuddin Nurkin
Pusat Studi Lingkungan Hidup UNHAS, Ujung Pandang, South Sulawesi, Indonesia
Abstract
In South Sulawesi forests contain a large variety of genera and species of plants. These forests are
important as sources of timber, fuelwood, food and many other minor products. The major concern over
this important coastal resource is its increasing rate of exploitation. Prior to 1965 it was estimated that
there were at least 110 000 hectares of mangrove forests covering the coast of South Sulawesi. How-
ever, today about 80000 hectares have already been cleared for timber, fuelwood, and for conversion
to tambak (brackish water ponds). Tambak is one of the most common forms ofland use in the coastal
region of South Sulawesi, and has created a serious land use conflict. Conservation efforts involve coastal
revegetation projects and the establishment of a 200 metre wide 'green belt' all along the coast. The
implementation of these protective measures is discussed.
Introduction settlement has also caused mangrove degrada-

tion. However, conversion of mangroves to tam-
South Sulawesi is the most densely populated bak is by far the leading direct cause of mangrove
province in the eastern part of Indonesia. With a destruction.
population about 7 million (ratio ofland to popu-
lation about 1 ha/person) most of the arable land Distribution and composition of mangrove
is being utilized for agricultural production. In the
lowland regions, rice is the chief crop. South Su- It has been estimated that in the early 1950's
lawesi leads the other provinces (excluding Java) mangroves in South Sulawesi covered an area of
in the production of rice. Upland and mountain- about 110 000 hectares. Today, the remaining area
ous areas are also cultivated, for cloves, cocoa, is only 25 %of that original total. Based on ground
coffee, and horticultural products. surveys and aerial photo interpretation, Barkey
In coastal areas, tambak culture (brackish- (1990) estimated that total remaining area of
water fish ponds), has been developed extensively. mangrove comprises about 34000 hectares.
Most of tambaks are created by clearing man- The largest area of remaining mangroves are
grove forests. During the recent years the govern- prominent in Malangke and Malili, at the north-
ment has encouraged the expansion of tambak ern end of the Gulf of Bone and cover about
development by providing credits for farmers. 23000 hectares. The next largest area about
This is primarily part of the effort to increase the 8000 ha, lies in the northwest province in
quantity and value of export commodities to im- Lariang-Lumu, part of the Mamuju region
prove national income and provide foreign ex- (Fig. 1).
change. A study by PSL (Pusat Studi Lingkungan)
Over time, a combination of timber harvesting, UNHAS (1980, 1985) indicated that there are
fuelwood gathering, and clearing for human more than 30 species of mangroves in South Su-
272
SULAWESI
o"",_ _ _ _...1'60 km
GULF
OF
BONE
-
J
N-
~ MANGROVE
PANGKAJENE . . . . " TAMBAK
Fig. I. Distribution of remaining mangrove and tambak locations in South Sulawesi.
lawesi. These include trees, and shrubs along both Among the most important species are Bru-
the eastern and western coasts (Table 1). guiera gymnorhiza, B. cylindrica, Rhizophora api-
273
Table 1. List of mangrove species recorded along the eastern coastal region, especially in the southern part, has
(E) and western (W) coasts of South Sulawesi.
been developed for either tambak or dwelling
Scientific name Local name Coast areas.
Rhizophors apiculata BI. Lenro E,W

R. mucronata Bakko E,W Timber utilization
Bruguiera gymnorhiza L. Kaja-kajang E,W
B. parviflora (Roxb) W & A Gandi-gandi E
Ceriops tagal (Perr) C.B. Reb Tangere E,W Traditionally, in South Sulawesi the use of man-
Sonne ratio alba Vierh. Beroppa E,W grove as sources of timber and fuelwood has long
S. ovata Back Beroppa batu E been recognized. Bruguiera, Rhizophora, Xylocar-
S. acida L.F. Padada E,W pus and Intsia have strong, attractive, and durable
A vicennia marinaVierh. Lesse-Iesse E,W
woods for housing, boats, and other construction
Xylocarpus granatum Koen Buli cella E
X. moluccensis Roem Buli lotong E uses. Beside having a high caloric value, some of
Scyphiphora hydrophyllacea Giling-kiling E,W the species especially Rhizophora are highly pre-
Gaertn. ferred for household consumption because they
Morinda citrifolia L. Bangkudu E may improve the flavour of food when used for
Heritiera littoralis D. Talungan E,W
cooking.
Lumnitzera littorea Voigt Bunga-bunga E,W
Terminalia catappa L. Katapang E,W In Malangke area, the average volume per
Dolihendrone spathoceae Tui E hectare of three principal commercial species
K. Schum were: Rhizophora apiculata between 40.54-
Aegiceras corniculatum Utti-utti E,W 120.15 cu. m. (average 68.59 cu m), R. mucronata
(L.) Blanco
between 0.89-14.52 cu. m (average 3.86 cu. m),
Cerbera manghas L. Sali-saling E
Exocoecaria agallocha Buta-buta W and Bruguiera gymnorhiza typically of the order of
Cynometra sp. E 12.80-80.96 cu. m (average 35.63 cu. m). These
Intsia bijuga O.K. Ipi (Bayam) E figures are much higher than the average volume
Pongamia pinnata Merr. Langi-Iangi bokka E per hectare for mangroves overall in Indonesia,
Derris heterophylla Binre-binreng E
where Rhizophora and Bruguiera is estimated at
(Wild) Back
Serianthes minahassae Langi serutu E 45 cu. m per hectare (Wiroatmodjo & Judi, 1979;
Merr. et Perry Nurkin, 1980).
Ficus sp. Barana E In the region where a small portion of the man-
Gironniera sp. Taluku-Iuku E grove areas have remained relatively intact, i.e. in
Barringtonia sp. Alakkang E
Luwu and Mamuju, the direct uses of mangrove
Hibiscus tiliaceus L. Waru E
Nypa fruticans Wurmb. Nipa E,W for timber, fuelwood, and additional products
A canthus illicifolius L. EIIi-elli E,W such as fruits, fish, and crustaceans have pro-
Acrostichum aureum L. Lappio E,W vided livelihood and sustained the economy of
traditional communities of coastal villagers. Ap-
parently, this kind of small scale utilization has
culata, R. mucronata, Intsia sp. and Xylocarpus. had a negligible impact on the mangrove commu-
As shown in Table 1, mangroves on the eastern nity.
coast contain more species compared than along Commercial utilization of mangrove timbers
the western coast. This is particularly notable in were carried out in the early 1970's to supply
Malangke area where the mangrove community the Gowa-based paper industry. The felling
has developed on the great river delta of operations involving selection cutting, have been
Rongkong. The other reason for the poorer man- concentrated in Malili-Malangke area. In 1974, a
grove community along the western coast is be- total of 26 339 cu. m. of woodpulp was produced
cause the remaining stands have been subjected (Nurkin, 1979). It has been reported that a mix-
to over cutting for a longer period and most of the ture of mangrove wood fibers with bamboo chips
274
produce an improved quality of paper. Currently, prawns harvests between 92-205 kg ha - I y- I

due to the small total remaining area and under (136 kg on average). However, the yield per hect-
supply of desirable species, commercial logging are can be increased by greater management in-
has ceased. tensity including the application of fertilizers and
the provision of artificial feed for milkfish and
prawns. In such cases, yields have increased to
Tambak culture about 575 kg ha - I Y- I of milkfish and 209 kg
ha - I Y- I of prawns in the same ponds. Conse-
Tambak culture is one of the most common fea- quently, the overall incomes received by farmers
tures of land use in coastal region of South Su- generated from tambak culture are very much de-
lawesi. Tambak expansion has increased dramati- pendent on the level of tambak management.
cally during the last two decades. In 1969, the
Department of Agriculture reported that there
were about 37000 hectares of tambak culture in Regeneration and preservation
South Sulawesi province. Twenty two years later
this figure had doubled to 74286 hectares, which Study of natural regeneration in mangrove areas
account for approximately 30 percent of the total following logging operations has been conducted
tambak area in Indonesia. Through the program in Malangke and Mamuju (Nurkin, 1980; PSL
developed by the Fishery Extension Service, the UNHAS, 1985). On most suitable substrates
government has launched an ambitious plan to when the Rhizophora seeds are able to settle as
create more tambak in the next five to ten years. a result of favourable conditions, natural regen-
Based on the preliminary study conducted by this eration may become successfully established in
office the total tambak for the province will be less than a year.
reached and be maintained at a total of 150000 In Malangke forest, naturally regenerated
hectares. Because of the limited amount of man- Rhizophora and Bruguiera were recorded one
grove lands remaining, the new tambak to be de- year after clear-cutting ranged from 560-18900
veloped will include former rice fields especially in plants per hectare with an average height of 30 cm
the southwestern part of the region. Most farm- (range of 12 to 76 cm) and stem diameter of 0.6
ers prefer tambak to rice culture. They claim that to 2.1 cm. On other plots at six years of age, trees
tambak culture is attractive due to the strong mar- of both species reached heights ranging from 6 to
ket and higher revenues for milkfish and prawns. 11 m with stem diameter of 3.2 to 6.5 cm. All of
Baruadi (1990) and Cholik & Hanafi (1991) this regeneration came from seed trees remaining
have provided some data and information on on the next block of the clear cutting area.
socio-economic aspects of the tambaks in South In a survey made in Mamuju region, data re-
Sulawesi. In 1989, tambak production generated corded have shown that in areas where the old
an income of about Rp. 165,000 million, equiva- stands have been removed, the establishment of
lent to US$90 million (U.S. $1 = Rp. 1900), and a regeneration phase is easily identified and
provided job opportunities to about 27500 per- counted. In such areas, the average seedlings and
sons. saplings per hectare of Rhizophora were 840 and
Milkfish (Chanos chanos) and prawns (Penaeus 1600 respectively.
monodon and P. merguiensis) are the species In areas where tambaks have been developed,
widely stocked and are raised in tambaks ranging the destruction of mangrove has led to severe
in size from less than 1 to more than 10 hectares shortage of firewood. Farmers realized the need
as a polyculture system. Productivity of the tra- to restock these areas. Many villagers have raised
ditional tambak has been considered low, with mangrove trees on the boundaries of their tam-
milkfish harvests typically of the order of 240- baks. The planted trees also serve as a measure
535 kg ha - I y- I (average 391 kg ha - I y-l), and of protection for unstable riverbanks. After eight
275
to ten years plantations of Rhizophora in strips of fauna might rapidly disappear. Through LAND-
5 to 8 m width have formed a remarkable, two SAT and SPOT images, Barkey (1990) has pro-
storied stand. The largest trees then are regularly vided an estimate rate of mangrove degradation
cut for fuelwood. By doing this the smaller trees on the eastern coast ofthe province. In Malangke
underneath which originated from natural regen- and Bone-bone area the mangrove cover declined
eration are able to develop to form a new stand. from 5528 hectares to 4014 hectares between 1978
Efforts at larger scale artificial regeneration in- and 1986, or an estimated rate of degradation of
volve a coastal revegetation project and the es- about 189 hectares (3.4 percent) annually. These
tablishment of a 200 m wide prescribed 'green figures of destruction will probably increase in the
belt' along coastal lines and 50 m wide belt along future as the pressure continues for tambak de-
river banks. Initiated in 1989 this project is under velopment. Apparently the forces behind utiliza-
the coordination and planning of the Land Re- tion for mariculture remain strong, and restora-
habilitation and Soil Conservation Services. tion and preservation efforts are currently
Official data indicate that an area of about 1500 inadequate to reverse the loss of mangroves.
hectares has been replanted with Rhizophora Results from Whitten et al. (1987) and Giesen
seedlings, mostly in Sinjai, Jeneponto, Maros, (1990) underline the need for establishment of
Pangkep and Polmas. Since regeneration evalu- suitable mangrove protection aress. A reserve has
ation is still underway, no further data or infor- been proposed by Giesen, i.e. the Lariang area,
mation is available regarding seedling survival and on the north western coast of the province. The
establishment. area could be used to provide a source of reveg-
The effectiveness of a 200 m greenbelt along etation stock, and a dwelling for wildlife. In ad-
the coastal line for the purposes of mangrove mul- dition, as indicated by PSL (1980) on the eastern
tiple use was discussed by Whitten et al. (1987). coast, the Malangke and adjacent areas also
Moreover, it has been the subject of an inter- should be given priority for protection. The area
departmental debate in Indonesia for a decade. is unique and represents the richest community of
Fisheries, for example, is seeking a much wider mangrove remaining in South Sulawesi.
protected zone of 500 m for the benefit of inshore Regarding mangrove utilisation for aquaculture
and offshore fisheries. As suggested by Woodroffe development, PSL UNHAS (1985) and Giesen
(1985), cited by Whitten etal. (1987), green belt (1990) recommended intensification programmes
argument from just a forestry or fisheries perspec- for existing tambaks to increase their yields and
tive inevitably neglects the multiple use impor- values.
tance of mangroves.
Based on his recent study on wetlands, Giesen Conclusion
(1990) also emphasized the need to provide an
adequate, larger preservation area of mangroves Mangrove degradation is one of the most con-
in South Sulawesi. He pointed out that a narrow, spicuous results of coastal development in South
200 m width of green belt might not be sufficient Sulawesi. So far, within four decades about 70
to protect wildlife habitats. His observations in- percent of the total original area of mangrove
dicate that certain species depend on large tracts have been converted, mainly for tambak culture.
of relatively undisturbed mangrove for their ex- Recognizing the economic advantages of raising
istence, while others require at least some trees milkfish and prawns, tambak culture is likely to
for roosting and shelter. In addition, some species remain as the most important man made land use
need a larger area of relatively undisturbed man- practise in coastal areas.
groves for breeding purposes. The need to preserve and restore mangrove
With the currently rapid rate of degradation communities in South Sulawesi has been empha-
mainly due to tambak culture development, a sized in recent years. It has been urged that a
number of important species of both flora and much expanded effort should be undertaken in
276
order to ensure the protection of remaining man- Cholik, F. & A. Hanafi, 1991. Dampak penerapan teknologi
groves. This effort should include the following produksi budidaya perikanan pada peningkatan kesejahter-
aan petani tambak. Paper presented on Forum Peri-
measures: kanan II. Litbang Pertanian dan USAIDjFRDP.
1. Establishment of protected areas, namely in Giesen, W., 1990. The importance of wetlands in South Su-
lawesi. In M. Zierren, Y. Rusila Nur & PHPA (eds),
Lariang and Malangke regions. Proseding seminar keterpaduan antara konservasi dan
2. Revegetation and the maintenance of a 200 m tataguna lahan basah di Sulawesi Selatan. Ditjen PHP Aj
wide 'green belt' along the shore. Kanwil Dephut Prop. Sulse\.
3. Preventing conversion of the intact, remaining Nurkin, B., 1979. Beberapa catatan tentang aspek pengusa-
haan hutan mangrove di Sulawesi Selatan. In S. Soemidi-
mangroves by tambak expansion by focusing
hardjo, A. Nontji & A. Djamali (eds), Pro siding seminar
on intensification of existing tambaks to in- ekosistem hutan mangrove. LON - LIPI, Jakarta.
crease productivity. Nurkin, B., 1980. Hutan mangrove Malangke. Bagian kehu-
tanan, FakuItas Pertanian, Universitas Hasanuddin, Ujung
Pandang.
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ditinjau dari aspek konservasinya. In M. Zierren, Y. Rusila payau di Indonesia. In S. Soemidihardjo, A. Nontji &
A. Djamali (eds), Pro siding seminar ekosistem hutan man-
Nur & PHPA (eds)J Proseding seminar keterpaduan ant-
ara konservasi dan tataguna lahan basah di Sulawesi Se- grove. LON-LIPI, Jakarta.
latan. Ditjen PHPAjKanwil Dephut Prop. Sulse\.
Hydrobiologia 285: 277-282, 1994.
!£) 1994 Kluwer Academic Publishers.
Value of mangroves in coastal protection
Muhammad Akhir Othman

Coastal Engineer, Department of Irrigation and Drainage, lalan Sultan Salahuddin, 50626 Kuala
Lumpur, Malaysia
Abstract
Nearly 30% of the coastline of Malaysia is undergoing erosion. Many of these areas are coastal mud-
flats, fringed by mangroves. Behind the mangroves there are usually agricultural areas that are protected
by bunds from tidal inundation. These bunds are constructed by the Department of Irrigation and
Drainage and it is the policy of the department to maintain a strip of mangroves between the bunds and
the sea. Mangroves are known to reduce wave energy as waves travel through them. Thus, mangroves
are used to protect the bunds from eroding. However, mangroves themselves are susceptible to erosion.
Finding the best method in using this natural system of coastal protection is therefore important to the
Department of Irrigation and Drainage. This paper looks at the various methods of using the systems
developed to date.
Introduction mud and the plants take root. In this manner, the
trees can propagate further and further onto the
Mangroves occur along much of the west coast of mud-flats.
Peninsular Malaysia. The sheltering effect of the In order for A vicennia to take root, the wave
island of Sumatra provides relatively calm seas in conditions must be mild and the mud must be
the Straits of Malacca, compared to the South totally exposed for a certain time daily. For this
China Sea that abuts the east coast of the pen- reason, the trees are usually found above Mean
insular. Rivers that discharge into the straits carry Sea Level.
fine silt and clay into the straits. Due to the shel- When A vicennia has established itself, the
tering effect, the mud gets deposited along the waves that breaks on the trees and the rising tide
coast building the alluvial plains that characterise usually deposit debris and sediments behind the
much of the coastal areas of the west coast. On trees. This results in levees at the back of the
this mud the mangroves find the necessary sub- trees. These levees restrict the ebb flow of the tide
strate to establish themselves. causing permanent pools of water. Fresh water
Wave height along the west coast rarely ex- from land and rain reduces the salinity of the
ceeds 1.5 metres. During the calm seasons the water. Due to the constant inundation, the
waves are usually below 1.0 meters. This provides breathing roots of the A vicennia cannot take in
excellent conditions for mangroves to take root. oxygen and the trees cannot survive. New Avi-
Mangroves colonise the coastal mud-flats in suc- cennia trees also cannot take root as the seeds
cessive stages. The pioneer species is A vicennia. will remain floating in the water. Under these
Their seeds drop from the trees and float in the conditions, Rhizophora and the Bruguieria species
water. During low tide, the seeds get lodged in the take over.
278
DID bunding schemes Mud-trapping
As early as the 1950s until as late as the 1980s, Mangroves especially A vicennia slow down the
the Department of Irrigation and Drainage was currents with their roots systems. This water can
building bunds (earthen dykes) along the west be from two sources, either the tide or the fresh
coast within mangrove areas. The projects were water discharge from the hinterland. The water
implemented to create more agricultural lands by usually carries sediments, in the form of sus-
reclaiming the mangrove swamps. The criterion pended sediments and bed load. As the ability of
used in building the bunds was that there must be water to carry sediments depends on the velocity
at least a 200 metres mangrove belt along the of the flow, slowing the currents result in the sedi-
coast between the bunds and the sea to prevent ments settling. In this way, the mangroves con-
the waves from damaging the bunds. This is be- solidate the soil and build up land.
cause the bunds were constructed from earth and
are susceptible to wave damage.
During the construction of the bund, a 200 Retarding coastal erosion
metres width of mangrove belt was considered
sufficient to reduce the wave energy such that the Erosion of the muddy coast usually starts with
waves will not damage the bunds. Observations lowering of the mud-flats in front of the man-
done in Sungai Besar, Selangor, suggests that groves. This causes the roots of the mangroves
even a 50 metres wide belt of A vicennia is suffi- fringing the sea to be exposed. Eventually these
cient to reduce waves of 1 metre to a height less trees collapse and the erosion continues further
than 0.3 metre. However, work done using a com- into the mangrove belt (Fig. 1).
puter model provided by the consultants of the The cause of the lowering of the mud-flats is
National Coastal Erosion Study 1986 suggests still unknown. There are many possible reasons,
that 150 metres of mangrove width is needed to that can be accepted. Reduction of sediment
reduce the wave energy. supply to the coast can be a factor. The mud on
the mud-flats is dynamic due to constant agita-
tion by waves. When sediment supply from the
Mechanism of wave attenuation hinterland to the coast is reduced, there are not
enough sediments to replenish the mud that is lost
Mangroves attenuate waves (reduce wave energy) from the coast due to wave erosion.
by obstructing the waves with its roots and trunks. Clay particles tend to flocculate in saline water.
As a wave passes through the mangroves, the Thus, there is a greater tendency for clay particles
orbital motion of the water particles i.e. the to settle down in calm saline water than in fresh
mechanism transmitting the wave energy, is ob- water. Reduction of wave action, perhaps due to
structed by the roots and the trunks of the man- a series of calm seasons over a long period, can
groves. The closer the trees are together, the cause the clay particles to settle and thus build up
greater will be the attenuation of wave energy. the mud-flats. On the other hand, a series of ex-
Mangroves need not be very old before they are treme storms can cause erosion of the mud.
significant as wave attenuators. As long as the Observations along the west coast of Peninsu-
trees are sufficiently close together and are as high lar Malaysia suggests that mud-flats undergo a
as the in-coming wave, the trees will attenuate the cycle of accretion and erosion. In one site, Sungai
waves. It has been observed that a 5 year new Burong, Pulau Pinang, this cycle is about 20 years.
growth of A vicennia can act as an efficient wave Mangroves have an important role to play in the
attenuator. accretion and erosion cycle of the muddy coast.
The swamp, with its various species, act as a
system in retarding erosion.
279
However in the Rhizophora and the Bruguieria

zones, the combination of stronger soils and
deeper root system retards the erosion. When the
ORIGINAL CONDITION
next accretion cycle occurs, the A vicennia will
colonise the mud-flats again.
Damage of the system

INTERTIOAL SURFACE EROOE S
In much of the west coast of the peninsular, the
- - - - - - - - - .;;:--
complete mangrove system has been damaged.
Reclamation of the mangrove swamps for various
development projects usually means the destruc-
tion of the Rhizophora and Bruguieria zones. This
SURFACE CONT INUE S TO
is because these zones have stronger soils since
ERODE. SCARP FORMS
the soil have had more time to consolidate.
sz
--------~ Changes in drainage patterns can also affect
the system. Reduction of sediment supply to the
coast due to these changes will result in coastal
erosion. This reduction also means that there is
SCARP HEIGHT INCREASES.
no new build up of mud for the mangroves to
MANGROVE LINE RETREATS
colonise. Thus, in some cases, the coastline may
be under constant retreat.
Replanting of mangroves
Fig. 1. Mangrove retreat. Stages in erosion of a mud coast-
line (top to bottom). Erosion usually starts with lowering of The Drainage and Irrigation Department had
the mudflat (top). In the final stage collapse and erosion con- made several attempts at replanting of mangroves
tinues into the mangrove belt (bottom).
with varying degrees of success. In the 1970s, at
Jeram, Tanjung Karang and Sungai Besar in
Selangor, bamboo poles were split lengthwise and
A vicennia are usually the first to topple when stuck into the mudflat. It was hoped that these
the mud-flat lowers. Erosion under the trees easily bamboo poles would hold the mud-flat while the
exposes the roots as the roots are extended along mangroves established themselves. There was
the surface, but do not penetrate deep into the success in some places but in most places the
mud. The soil under the trees is also soft, being attempt resulted in failure. This was due to the
newly formed, and is also susceptible to erosion. mangroves being planted on a very low mud-flat
As erosion propagates into the Rhizophora and level, making the daily inundation period by sea
Bruguieria zones, the rate is slowed down. With water too long and the wave action too strong.
their prop roots that grow deep into the mud, the In Sungei Burong, Selangor, DID had a pilot
Rhizophora and the Bruguieria can anchor them- project replanting 10 hectares of mangroves. Six-
selves in the soft clay and withstand the natural teen thousand seedlings of Rhizophora were
forces. The soils in this zone, having had more planted upon advice from the Forest Research
time to consolidate are also usually firmer. Institute of Malaysia. Although during the early
Thus when the wave climate changes and larger years, the young mangroves suffered high mortal-
waves hit the coast, the mud-flats reduce in level ity due to attacks by caterpillars, 20% of the trees
and the A vicennia topple due to under scouring. survived and established themselves. A vicennia
280
also colonised the area in time due to large stands

of the species on both sides of the area. The seed- -d
a §
lings were brought into the area by littoral cur- z
« .n
rents and deposited in the area when the tide ...J
~
<I)
receded. =
<I)
The Sungei Burong experience reveals that; -5

~
u
B
(a) Avicennia species grows well in tidal mudflats 8c:>.
where the tidal flushing is most efficient, i.e. .9
during low tide the mud is completely ex- <-
VI
I<: ~
posed, and there is no stagnant water. This U
01<: '="
<I)
condition usually occurs at the fringe where O:u

cDi:
~
d
the mangrove belt meets the sea. u.ol- <I)
1;
(b) Dunes of crushed shells and sand usually °e
«,., ~
cr. oS
0
occur behind the A vicennia and prevent the
I!:)
'"oS
water from draining completely. This makes '"
~
~
the area more suitable for Rhizophora species. '"
<I)
The long seedlings of the Rhizophora can b<I)

penetrate the soil when they drop and estab-
OVI
u.o u.o 8
... > 0
ZO ~
lish themselves and are also long enough not <t cr-
...JI!:) ....0
to be completely covered by water. Avicennia ~z
u.o<t '"
<I)
seedlings will remain afloat in this condition crE ~\T >

0
and will not be able to establish them- L3~

c_-:J;-
6h
=
oS
selves. Furthermore, the breathing roots of <::J- ....08
the A vicennia will always be covered by water
a..
~
~-.
~
<l
and will not be able to take in air. ::I: f::J-~ .n
(c) Planting a mono crop i.e. one species of man-

5 <.~ «
....
groves will make the crop susceptible to pests 0
~
=
0
.;;;
0
and diseases. This is similar to a plantation VI <-
<I)
I c:>.
problem where a certain pest can destroy a .9
large area of crop in one attack. The best way '"
.9
to avoid this problem is to diversify the spe- c
<I)
@
cies. There are over 50 species of mangroves ~
~ '"
oS
0
to choose from and planting a mixture of spe- 0
cr
cr
u
<I)
cies will give a better chance of the regener- 0 >

0
CD
ated forest surviving. 6h
0
...J §
8
....0
0
I
=
0
Escarpment protection '"8

0
z
=>
~c:>.
cD
The Department of Irrigation and Drainage are c
.... '85
also experimenting with a combination of artifi- 0 c:>.
cial and natural methods of coastal protection. I til
u
The concept here is to protect the mangroves '"

"-l
so that the mangroves can in turn protect the

valuable agricultural behind. This method is called «
'~"
UJ
escarpment protection and was first employed Vl
281
successfully at Sungai Burong, Sabak Bern am, escarpment will also avoid loading that would
Selangor, to protect a stretch of 1.4 km of coast- endanger the bund. Should there be any failure of
line (Fig. 2). the structure due to localised weak soil, there
The soil underneath the mangrove forest is usu- would be no immediate danger to the bund and
ally firmer than that of the deposited mud on the flooding of agriculture land would not occur. The
mudflat. Thus, when erosion occurs and the mud- Drainage and Irrigation Department would have
flat lowers, the soil develops a vertical face ample time to react to the failure before the bund
underneath the mangrove line. The vertical face is threatened.
ranges from 0.5 to 1.0 m. This face is called the Four years after construction, this method have
escarpment. been proven successful in Sungai Burong. The
Mangrove trees have always been regarded as escarpment protection is preventing the soil under-
a wave attenuator, reducing wave energy as the neath the mangroves from eroding and the man-
wave travels through the swamp. Due to this, groves are reducing the wave energy before the
many efforts to replant mangroves on eroding waves reach the bund. With the success of this
mudflats have been tried in order to arrest method, this concept is being used along the
the erosion. However, mangroves cannot estab- adjacent coast to protect another 1.2 km of coast-
lish below Mean Sea Level (M.S.L.). Since line.
the levels of the eroded mudflats are generally
below M.S.L., the efforts were not successful.
Between M.S.L. and Mean High Water Spring Conclusions
(M.H.W.S.), mangroves grow very well and can
provide adequate protection to the bund. The key Mangrove swamps are important in retarding
is therefore to protect the soil beneath the man- coastal erosion and protecting the coast. The
groves, so that the trees will be able to do the rest swamps act as a system in which trees in all the
of the work in reducing the wave energy. Thus, zones have different roles to play. Damage to the
the escarpment protection concept was formu- system can result in irreversible coastal erosion.
lated. Mangrove replanting can be successful if the
The structure was constructed such that it will mangroves are planted in the right conditions. A
not have a detrimental effect on the mangrove lot of failures can be attributed to mistakes such
ecosystem. The ebb flow must be able to go over as planting on very low mudflats and where the
the structure freely so that there will be no stag- wave conditions are not favourable. Mono-
nant pools developing behind the structure. This cropping should be avoided to reduce damage
is because different mangrove species require dif- due to pests. Here again, if the objective is to
ferent salinities and disturbing the ebb flow could protect the coast, it is best to re-establish the
affect the salinity regime. system than just a certain species.
The structure will protect the escarpment from Where the lowering of the mudflats is unavoid-
erosion due to wave attack. Since the structure is able and further retreat of mangroves is unaccept-
low, waves will still penetrate during high tide. able, the escarpment protection is a feasible con-
Mangrove trees will then act as a wave attenua- cept. This method is still cheaper than the
tor and prevent the wave energy from reaching conventional method of protection using revet-
the bund. This will therefore prevent the bund ments as it uses less material. It also has the
from being eroded. A mangrove belt of about 100 added bonus of retaining the mangrove swamp
metres was provided to act as a wave attenuator. that can serve as a habitat for the fauna that
This is considered sufficient to reduce the wave depends on the swamp.
energy and protect the bund behind the belt.
A low structure has the advantage of being
easily constructed. Building the structure on the
282
References Othman, M. A., Sungei Burong Escarpment - A Combined

Structural and Natural Method of Coastal Protection, Pro-
ceedings of Conference on Coastal Engineering for Na-
Economic Planning Unit, Malaysia, National Coastal Ero-
tional Development, IEM-ICE, Kuala Lumpur, March
sion Study: Phase I, Report submitted by Stanley Consult-
1991.
ants, Inc., Jurutera Konsultant (S.E.A.) Pte. Ltd. and Mof-
fatt & Nichol, Engineers to the Government of Malaysia,
September 1985.
Hydrobiologia 285: 283-285, 1994.
A historical perspective of the resources and issues of Pak Phanang

Bay, southern Thailand
N. Srichai, S. Boromthanarat and B. Chaijaroenwatana

Coastal Resources Institute (CORIN), Prince of Songkla University, Hat Yai, Thailand 90100
Abstract
The Coastal Resources Institute (CORIN) at Prince of Songkla University worked with local commu-
nities to understand coastal issues and opportunities of the Pak Phanang Bay in Nakhon Si Thammarat,
Southern Thailand. Historically, the area has been an important agricultural field and trading town but
now it is characterized by slow population growth due to out migration and a decreased economic base.
Residents have noted major environmental degradation within their lifetimes. Through the 'Ecological
History', which means a history of the way people and nature have interacted over time, an initial
identification and assessment of the key environmental problems were made with the participation and
support of the people of Pak Phanang. The approach is based on the belief that for successful resources
management, it needs action by local people, supported by the government, that will safeguard and
restore local ecosystems.
Introduction the first step to achieve the goal. This paper sum-
marises a publication of CORIN, the history of
Southern Thailand is endowed with rich natural development of this important bay and the issues
resources along its 2600 km. of coastline. Ap- most affecting its management.
proximately 13 % of Thailand's population lives
in the 14 southern provinces. Throughout the
south there has been a rapid increase in the loss Objectives of the ecological history
of coastal resources as a consequence of growth
in tourism, fisheries, port development and shrimp The ecological history of Pak Phanang is a sum-
farming. In addition, economic growth and natu- mary and synthesis of data gathered which rep-
ral disasters caused by unprecedented rains in resents a new approach to how coastal problems
November of 1988, along with the destruction of and management issues should be identified and
mangroves, loss of fishery resources and the pol- analysed. In the process, the roots of today's
lution accompanying a rapidly increasing human problems are traced and the technical results of
population have created an urgent need for an research are combined with the knowledge, per-
integrated coastal resource management plan. It ceptions and values of the people who live in the
is the goal of the Coastal Resources Institute at area. The goal of the ecological history is to tell
Prince of Songkla's University, Thailand the story of the interaction of man and nature
(CORIN) to assist in developing a realistic man- over time with the following objectives: (1) to
agement plan for the sustainable use of Thailand's identify the key problems and understand which
coastal resources. The Ecological History of Pak of them are the most important issues to the resi-
Phanang Bay area in Nakhon Si Thammarat is dents and in need of coastal management strat-
284
egies; (2) to understand the context from which images, photos, government records, interviews
problems emerged and their interactions; (3) to with residents and government officials and com-
set the stage for further research and (4) to help munity meeting.
create public awareness and support of future Interviewing residents is a simple but effective
management. methodology to obtain resource-based informa-
tion. It involves the people who utilize coastal
resources and whose quality of life is affected by
The study area ecological and economic changes.
The district of Pak Phanang in N akhon Si Tham-

marat Province, was chosen as a study site. His- Results
torically it has been an important agricultural
area. Now, however, the province is character- In defining the issues and developing the Ecologi-
ized by slow population growth, partly due to cal History, a timeline of key events in the history
out-migration, and a decreased economic base. of the community was developed. This helped to
Most residents have noted major environmental identify past trends and put a perspective on
degradation within their lifetime. The Pak Pha- current issues.
nang Bay no longer produces thigh-sized fishes;
mangroves have been cut down for charcoal or
shrimp farming and the bay is much shallower
than it once was. Pak Phanang Bay Timeline
The study area incorporates approximately
300 km 2 of the coastal plain east of the N akhon 1900 Rice shipping is important
Si Thammarat mountain range. The Bay is a shal- 1940 Coastal shipping important; some foreign
low, elongated basin, approximately 14 km long trade.
and widening from 3 km at the mouth of the Pak Fish in the bay are big 'thigh sized'
Phanang river to nearly 10 km at the entrance of 1945 Purse seine introduced to Bay fishing.
the bay. Extensive tidal fiats and mangrove for- 1952 End of China Trade.
ests are found along the shoreline except on the 1957 First offshore trawler built.
north western side where they have been replaced 1960 Two-boat seine introduced.
by shrimp ponds. The area is made up of parts of 1962 Typhoon/depression.
Pak Phanang and Muang district and contains all 1964 River dredging.
or part of seven villages. In 1989, the total popu- 1965 Purse seining stopped, bay too shallow.
lation of the study area was less than 80000 people Mollusc production declined; shrimp and
and most of them were either rice farmers or fish- fish nets stopped; ghost boat fishing ended.
ermen. More recently, shrimp farming also has 1968 Rice trade by ship finished.
become an important occupation. In 1989, ap- 1970 Shrimp farming (extensive) started.
proximately 10 1 km 2 were under rice cultivation, 1972 River dredging.
62 km2 for shrimp aquaculture and 12 km 2 for 1975 Big fish gone. 'Thigh size' now 'arm size'.
orchards. Start of fish meal factories (1975).
1978 1100-1500 offshore trawlers in the fleet,
fishing fleet moved to Songkla to get better
Methodology market prices.
1980 Lift net fishery started, yielded 10 kg in
The History is the result of the compilation of a 1-2 hrs. Fishing with pesticides started.
variety of sources including published reports, 1984 River dredging.
books, newspapers, magazines, maps, satellite 1985 Mussels, cockles started to recover.
285
1988 River dredging. Intensive shrimp farming vestigated and will be addressed by a special area
started. management plan.
1990 River and bay dredging starts. Bay fishery Key issues include;
yielded in range of 2-5 kg d - 1. 70 % of Conservation and Allocation of Freshwater
trawler fleet landing at Songkla belong to Environmental Degradation ofPak Phanang
owners in Pak Phanang. Bay
Balancing Coastal Land Uses
Significant events in the history of the area were Breaking the Boom and Bust Cycle of Eco-
an era of prosperity based on rice trade and an nomic Development
active port, the disruption of World War II, the Other issues that are emerging, but are still
era of fisheries and the fish port, the devastation unexplored include protecting public health from
of the 1962 typhoon 'Harriet', the 1988 flood and waterborne disease and degrading water quality,
the droughts of 1989 and 1990. management of mangroves, sand mining and
flood control.
Management objectives need to be defined in
Discussion and conclusions close collaboration with Pak Phanang residents
and the provincial and national governments.
An integrated management plan for Pak Phanang Some of the issues will require further research.
is urgently needed. The preparation of the Eco- For example, the balance between fresh and salt
logical History (Boromthanarat et al., 1991) is water resources is not well understood and how
only the first step in a longer process of prepar- the change to shrimp farming has affected that
ing such a plan for the Pak Phanang area. Through balance must be determined. A more complete
the process of gathering data, reviewing literature, understanding of the complexities of the interre-
interviewing local residents and writing the Eco- lationships among local and national governmen-
logical History report, a clearer idea of the issues tal agencies, and the sometimes conflicting laws,
that face the people of Pak Phanang was devel- regulations and policies must be developed. Local
oped. A community workshop held to confirm the support for the concept of sustainable develop-
nature of the issues and extent of agreement about ment and for specific objectives must be built up.
them was organized in August 1991. Over 50 This requires a program to develop public aware-
people attended the workshop, including rice ness of the issues and support for the planning
farmers, shrimp farmers, teachers, businessmen, process. Finally, the development and implemen-
government officials, and members of voluntary, tation of a special area management plan will
non-governmental organizations. There was gen- complete the process.
eral agreement that the issues of water supply,
fisheries decline and shrimp farming were cur-
rently most important. Many people expressed
References
optimism about the future of Pak Phanang, and
suggested that further workshops of this nature
Boromthanarat, s. et at., 1991. Coastal management in Pak
could help to bridge the gap between local people Phanang: A historical perspective of the resources and is-
and government officials. Issues identified by sues. Coastal Resources Institute Publication, Prince of
CORIN and local residents must be further in- Songkla University, Hat Yai, Thailand, 96 pp.
Hydrobiologia 285: 287-302, 1994.
(£) 1994 Kluwer Academic Publishers.
Management of coastal ecosystems in eastern Sumatra: the case of

Berbak Wildlife Reserve, Jambi Province
Gordon Claridge
Asian Wetland Bureau - Indonesia, P,O. Box 254, Bogor 16001, Indonesia; James & Claridge
Environmental Management Consulting, 105 Honour Av., Chelmer, Brisbane, Q. 4068 Australia
Key words: peat swamp, management, forest products, settlement, transmigration, wildlife
Abstract
The eastern lowlands of Sumatra comprise about 88000 square kilometres, or approximately eighteen
percent of the island's total area. Most of these lowlands were originally peatswamp forest, freshwater
swamp forest and mangrove. A significant proportion of the area is subject to tidal influence.
The lowlands have experienced a variety of land uses, firstly by the native people, who were mainly
hunter-gatherers, later by the Buginese and Banjarese immigrants from Sulawesi and Kalimantan re-
spectively, and more recently by transmigrants from Java, Bali and Madura.
Land uses have involved principally agriculture, logging and fishing, and a wide range of associated
activities. As in most attempts to settle and convert wetlands to other uses, there have been many
problems. The Berbak Wildlife Reserve located on the southeast coast of the Province of J ambi pro-
vides a microcosm of these problems. The issues and impacts evident in Berbak include nearly all of
the issues and impacts affecting wetland areas elsewhere in the eastern lowlands and provide a useful
case study.
The Asian Wetland Bureau and the Indonesian Directorate General of Forest Protection and Nature
Conservation have carried out a two-year project which includes development of management of the
Reserve and the preparation of an Environmental Profile for the lowland wetlands of J ambi Province
as a background for regional planning. These components of the project are aimed at investigating and
resolving management problems affecting the Reserve, and may be extrapolated to the solution of similar
problems in other parts of the Sumatran lowlands.
The Eastern Lowlands of Sumatra deposits, frequently overlain by a layer of peat

that may reach depths of twenty meters (Scholz,
The Eastern Lowlands of Sumatra make up some 1983).
18 percent of the island (88000 square kilome- The majority of the Eastern Lowlands was
ters) in a strip running from Aceh in the north originally covered by forest, mainly peatswamp
to Lampung in the south (Fig. 1). They are at and freshwater swamp forest, with riverine forest
their widest in the central and southern parts along the levees bordering the larger rivers and
(Provinces of Riau, J ambi and Sumatra Selatan. extensive mangrove belts along the coast.
Most of the area consists of alluvial and marine
288
following an initiative by the Netherlands Indies

Society for Nature Conservation. At that time
it had an area of around 190000 hectares (de
Wulf & Rauf, 1982; Silvius et al., 1984; Resubun
et al., 1988). The original decree did not elaborate
on the purpose of the wildlife reserve, and used
a Dutch term 'wildreservaat' that in general was
used for reserves established for their faunistic
o·
values. Literally translated, this term means 'game
reserve', though this incorrectly carries connota-
tions of hunting. A better translation which fits
the original purpose of the reserve is 'wildlife
reserve' (Silvius, pers. comrn.). Since, in general,
Dutch laws continued in force after Indonesian
~
o ZOOKm
Independence, the Wildlife Reserve remained in
existence and its management became the respon-
sibility of the Indonesian Government.
Fig. 1. The Eastern Lowlands of Sumatra (after Scholz,
1983).
Representativeness
Berbak Wildlife Reserve as a case study
The Berbak Wildlife Reserve as originally pro-
Status
claimed was undoubtedly a good representative
of the Eastern Lowlands of Sumatra, with signif-
Berbak Wildlife Reserve in southeastern J ambi
icant areas of all the major habitat types occur-
Province was declared in 1935 by a decree of the
ring in that land unit.
Governor-General of the Netherlands Indies,
Today it is still one of the largest swamp for-
est reserves in Indonesia, has the largest expanse
of freshwater swamp forest of any reserve in
Sumatra (Petocz, 1987) and contains the most
extensive area of peat swamp forest in any reserve
in the Asia-Pacific region (MacKinnon & Artha,
1982). The only other large areas of peat swamp
to have been included in Sumatran reserves
(Fig. 2) are at:
• Kerumutan Barn in Riau, which was declared
a Nature Reserve in 1979 but is now in poor
condition, with all or most ecological values
lost as a result oflogging and settlement (F AO,
1977; Santiapillai pers. comm.);
• Padang Sugihan in South Sumatra which was
~
o ZOOKm
partially cleared and drained for a transmigra-
tion project before being declared a reserve
(Giesen, pers. comm.);
Fig. 2. Peatswamp Conservation Reserves in the Eastern • Siak Keeil in Riau Province which was selec-
Lowlands. tively logged before (and for a short time after)
289
being gazetted in 1983, and which has been recorded from the Reserve, including the Sumat-
subjected to illegal logging ever since; and ran Tiger (Panthera tigris sumatrae), Clouded
• Way Kambas in Lampung, large areas of which Leopard (Neofelis nebulosa), Tapir (Tapirus indi-
have been logged. cus) and Sumatran Rhinoceros (Dicerorhinus
sumatrensis), all of which are endangered. Tigers
Petocz (1987) listed Berbak among the priority
and Tapir exist in high densities, apparently in
conservation areas for management and protec-
many areas of the Reserve, while there is recent
tion in Sumatra.
evidence of Rhinoceros from two areas (Claridge,
1991).
Biological diversity Survey work in 1991 suggests that the Reserve
may be one of the remaining strongholds for the
According to MacKinnon & MacKinnon (1986) False Gavial (Tomistoma schlegelii) in Sumatra
Sumatra is one of the richest units in the Sundaic and perhaps for its entire range in the wild (Cox,
Sub-Region of the Indo-Malayan realm. Within pers. comm.). Saltwater crocodiles (Crocodylus
Sumatra, Berbak constitutes a good example of porosus) occur in at least one of the rivers of Ber-
the biological richness of the Eastern Lowlands. bak.
The biodiversity of the area is extremely high,
though its full extent has not yet been ascertained
because large areas of the Reserve have not been Regional benefits from the Reserve
visited by biologists. Despite limited survey, 260
species of woody plants have been recorded, in- The extensive peat domes of Berbak function as
cluding 23 species of palms (Arecaceae) and 10 a reservoir offresh water for the surrounding area,
species of pandanus (Pandanaceae) (Giesen, providing not only drinking water for local com-
1991a). The Reserve is richer in palm species than munities but also contributing to the prevention
any other known swamp forest (Dransfield, 1974; of saltwater intrusion into agricultural areas. They
Giesen, pers comm.). also help to regulate the runoff of rainwater from
Little is currently known of the genetic poten- the area, thus limiting flooding in the adjacent
tial of the plants of Berbak, though it may be very communities.
significant. A number of genera that have been
commercially exploited (eg. Artocarpus sp., Bac-
caurea sp., Calamus sp., Cyrtostachys sp., Durio Management
sp., Dyera sp., Ixora sp., Mangifera sp.) are found
there. The Berbak Wildlife Reserve is managed by the
More than 250 bird species (48 families) have Directorate-General for Forest Protection and
been observed. These include all of the kingfisher Nature Conservation (PHPA) of the Forestry
(Alcedinidae) species known from Sumatra and Department through a sub-section of the Pro-
nine of the ten hornbill (Bucerotidae) species (Sil- vince level Office of Natural Resources (KSDA)
vius et al., 1984). The White-winged wood Duck in Nipah Panjang.
(Cairina scutulata) which is one of the world's
rarest waterfowl (Lambert, 1988) has recently
been found in two of the Reserve's rivers, and is The impact of settlement and population growth
reported to breed there (HIMBIO, in prep.). The
threatened Storm's Stork (Ciconia stormi), Milky Early settlement
Stork (Mycteria cinerea) and Lesser Adjutant
(Leptoptilos javanicus) are all regularly recorded in Before the coming of the Dutch the few settle-
the Reserve. ments were mainly along the rivers, particularly
More than thirty species of mammal have been those which rose in the Barisan Range and the
290
Piedmont Zone to its east, and along the coast settled land on the Berbak coast. In this northern
(Furukawa, 1986; Scholz, 1983). The most sig- area they also obtained some type of permission,
nificant settlements were at the limits of tidal possibly from a government official, but possibly
inundation which, as a result of the low topo- from a traditional Buginese leader, to settle cer-
graphy, is often very far upstream. (Altitude sel- tain areas on the coast. They also paid (some-
dom exceeds ten meters above sea level). The city times in gold) for the right to cut down the for-
of Jambi, which is some 150 kilometers from the est in the new areas, though it is unclear who was
coast and at the limit of tidal movement, is an paid (pers. comm. Kepala Desa Cemara whose
example of such a settlement. father was the leader of one of these groups. This
For a long time, prior to the second half of the information was repeated by another Buginese
Seventeenth Century, the coasts were sparsely settler from Air Hitam Laut who was a part of
occupied by a mixture of Melayu, Arab and one of these early waves; see also Silvius et aI.,
Turkish settlers. Some of the Melayu people had 1984).
a tradition of seasonal movement, possibly fol- These post-1950 Buginese settlers constituted
lowing movements offish or prawns, planting rice the first major management problem for the Re-
in temporary fields, which sometimes had shallow serve, and their impacts in Berbak were repeated
canals for drainage of the swampy soil. Berbak over much of the tidal zone of the Eastern Low-
was apparently one of their traditional seasonal lands. The techniques that they used for clearing
occupation sites (Furukawa, 1986), though this and draining the coastal swamplands were im-
would have been on only a very small scale. In- provements on a system that has been used for
land, the indigenous Kubu people carried on a centuries by the Banjarese in Kalimantan to grow
hunter-gatherer lifestyle that shunned contact rice and coconuts on peat soils (Hanson & Koe-
with the outside world and appears to have made soebiono, 1979). They cleared the forest, then dug
very little impact on the natural resources. a geometric system of canals, designed both to
There has been a long history of Buginese mi- drain the land and to irrigate it using the power
gration from Sulawesi to what are now Riau and of the tides to push freshwater into the rice fields.
Jambi Provinces. In the latter half of the Seven- Crops were initially rice, gradually changing to
teenth Century, with the outbreak of the Dutch- coconut as problems with soil and drainage made
Makassar War, refugees from South Sulawesi fled the land unsuitable for rice.
to all the coasts of Southeast Asia, including In this way more than 11 000 ha of virgin man-
what is now northern J ambi (but not Berbak in grove and peatswamp forest was cleared through-
southern J ambi). out the coastal part of the Reserve, and water
levels in adjacent areas were lowered (de Wulf &
Later settlement Rauf, 1982). The clearing and draining apparently
stopped only when they reached peat that was
During the upheavals in southern Sulawesi in the considered too deep for farming (pers. comm.,
1950s another wave of Buginese moved to the village head, Cemara), usually several kilometers
eastern coast of Sumatra, including the area north inland. This cleared area, together with large
of the Berbak River between TanjungJabung and cleared areas to the north, has been excised from
Kuala Tungkal. They were followed by further the Reserve (Fig. 3) by the Department of For-
waves of immigrants in the mid-1960s and in estry.
1970. These settled along the coast from Lam-
pung to northern Riau. Transmigration
In the J ambi area most, if not all, of these
people came first to the northern coast (north of The methods developed by the Buginese were
the Berbak River), where they were advised by later adopted by the government, with less suc-
earlier Buginese migrants of the existence of un- cess, in official transmigration programs.
291
~
o 6KM
.".--~-'
,,' (;>"" / ' '--
eV' I --
~~ I \
'
I \
'-
I/ I
~/ \
/
I t
/ '\
\
,,
\
\
,"....
'.,"":... ~:;,
~~
"::"_... 0<:'0,
...., ~ .
" ,' "" <?~
'\
\ ....
\'.
\ ....
\ ",
\ "
\ ....
\
1.-_ .. --.- ....... ·
Original boun d aries

' (1935)
Current boundaries
Fig. 3. Berbak Wildlife Reserve.

292
Between 1969 and 1974 transmigration pest habitat. The result is the same - fires destroy
projects began to open up extensive areas of the or reduce the value and viability of swamp forest
swamps of the Eastern Lowlands, especially in habitats. In Berbak fires are not uncommon near
the provinces of Riau, J ambi and Sumatra Sela- the settled areas on the margin of the Reserve, at
tan. These official schemes are generally regarded least in part because the belief is prevalent among
as the major force in clearing the forests of the settlers that the edge of the forest is the bound-
East Sumatran swamps. However, the role of the ary of the Reserve. If they can push back the
Buginese and spontaneous transmigrants is also forest edge then their land is enlarged and the
significant. In the period 1969-1973 the Buginese Reserve decreased (YASIKA, pers. comm.).
settlers in the Musi-Banyuasin delta of Sumatra Only the fact that the peatswamp is usually satu-
Selatan opened as much forest area as the rated has limited the damage to this habitat. Nev-
government program, and they have operated in ertheless, in 1982 fires affected 3400 ha of partly
all the major deltas of the east coast (Hanson & undisturbed peatswamp forest (Silvius et al.,
Koesoebiono, 1979). 1984). A significant area was probably affected
Spontaneous transmigrants (unassisted sett- again in 1991 when fires were reported in the
lers, usually from Java or Bali, but sometimes southern part of the Reserve (Universitas Padj-
escaping from unsuccessful official transmigra- adjaran survey team, pers. comm).
tion schemes) opened up an area of between Apart from lowering the watertable and thereby
40 percent of that opened by official programs increasing the risk of fires by drying out the peat,
(Danielsen & Verheugt, 1990) and 300 percent drainage which is not properly regulated has other
(BCEOM, 1990). highly deleterious effects. For example:
Buginese settlement, official transmigration and
• peat tends to subside after being drained, with
spontaneous transmigration doubled the popula-
the result that the ground surface is lower and
tion of the Indragiri-Batanghari section of the
floods and tides have a greater influence;
Eastern Lowlands in the sixties and seventies
• irreversible shrinkage of peat adversely affects
(Scholz, 1983) and this is probably true for the
its ability to retain water;
whole of the lowlands.
• potential acid-sulphate soils are exposed to
air as a result of the lowered water table,
thus causing acidification and rendering the
Impacts of settlement
soils unfit for agriculture (Soepraptohardjo &
Driessen, 1976).
All settlers of the swamp areas have faced a num-
ber of challenges, both to become established and Such has been the scale of settlement and logging
to provide their daily needs, and these have cre- operations that by 1989 only thirty percent of the
ated management problems for those with respon- original lowland forest remained in Sumatra, and
sibility for the natural resources of the area. only eighteen percent of the freshwater swamp
forest. Much of this remaining area is disturbed
and in small, isolated blocks (Whitten, 1989) that
Clearing and drainage would not constitute viable conservation reserves
or wildlife habitat, particularly for the larger
As a part of initial clearing, and frequently as a mammals.
preliminary to opening an extension to a cleared
area, spontaneous transmigrants often start fires
at the end of the dry season to remove as much Pests
vegetation as possible. Sometimes also, fires es-
cape from farmed land, or from areas of scrub on Agricultural activities in areas of cleared forest in
cleared land where farmers are trying to eradicate the Eastern Lowlands generally lead to conflicts
293
between people and wildlife because certain spe- of Jambi as a whole the mangrove forest has been
cies become pests. Some species, eg pigs, rats and reduced from several tens of thousands of hect-
probably monkeys, increase in numbers after par- ares I to around five thousand hectares in 1989
tial clearing of an area, while others do not seem (based on interpretation of radar and SPOT im-
to increase but become pests when they come out agery). This can be attributed partly to cutting for
of the remaining forest patches to raid crop areas fuel, partly to the demand for construction mate-
or livestock. Deer, bears and tigers fall into this rial and in part to the reclamation of the rearward
latter category. areas for agriculture.
The problems with pigs and rats have often
been incorrectly attributed to the presence of rem-
nant stands of primary forest (for example, Construction material
BCEOM, 1990). However, it is clear that these
species actually do better in areas of secondary The mangrove and freshwater forests provide
regrowth on areas of abandoned or logged land materials for a wide range of construction pur-
than they do in primary forest (Whitten, 1989; poses and this places a heavy and sometimes un-
pers. comm. various village heads, Berbak area). sustainable pressure on the resource. For ex-
Tigers and bears, on the other hand, prefer ample, off the coast of the Berbak Reserve there
primary forest and move out of this to raid are currently (1991) 42 large fish traps which
farmers' lands (Whitten, 1989). The experience of would have required 12600 three-metre poles for
many transmigrants shows that with increasing initial construction and up to 4000 poles for
density of settlement the willingness of tigers and maintenance each year. Since these traps are in
bears to enter the area decreases (pers. comm. the process of being phased out in favour of trawl-
various village heads). However, the presence of ing it can be assumed that in earlier times the
abandoned land, and hence a lower density of demand for mangrove poles was even higher.
settlement, means that these animals will con- Mangrove timbers are also used for construction
tinue to be a problem throughout large areas of of houses, jetties and boats. Little wonder that the
the lowlands while any primary forest remains. formerly extensive mangrove forests have virtu-
Problems for Reserve and wildlife managers ally disappeared.
arise through both the resentment that builds up The nipah palm (Nypafruticans) which grows in
against remaining forest areas which are seen as the back areas of mangrove forest is used for
the source of pests, and the trapping, shooting or thatching of roofs and walls and there is a con-
poisoning of protected and sometimes endan- tinuing high demand that in the Berbak area is
gered species by farmers. mainly satisfied by the nipah stands within the
Reserve.
Nibung (Oncosperma tigillarium) is used for
Fuel house poles, floors and steps, as well as for road
surfaces and foundations, and for fish trap sup-
Mangrove timber has traditionally been used as ports. An average house requires 100 nibung poles
firewood in Indonesia, not only for daily cooking, five to eight meters long, and these need to be
but also for small scale industries such as pro- renewed every three to five years (Danielsen &
duction of coconut oil, palm sugar, and lime (Soe- Verheugt, 1990). Berbak has dense nibung zones
giarto & Soemodihardjo, 1985). On the coast ad-
jacent to the Berbak Reserve the formerly
1 Sumardja (1989) gives 60000 hectares as the area of man-
extensive mangrove zone has been almost com-
grove in J ambi Province, however this seems to be a signifi-
pletely removed. Along a distance of around fifty cant over-estimate, but not as great as that of MacKinnon &
kilometers of coast there is now not more than Artha (1982) who give an original area of 165000 ha reduced
500 hectares of mangrove, and for the Province to 90000 ha.
294
behind the nipah on the river banks, though these The impact of economic development
are becoming depleted from illegal harvesting
which constitutes a management problem. There Particularly in the early stages of settlement
are approximately 600 houses in Air Hitam Laut, schemes, settlers have low levels of economic and
the largest village in the vicinity of Berbak. Of social welfare until their agricultural production
these, at least two-thirds use nibung in their con- becomes established. In some instances, for ex-
struction which suggests an annual demand of ample where sites have been located on deep peat
10000 poles or around 3500 to 4500 trees for this or acid sulphate soils, this stage can persist for
one village. many years. Under such situations the settlers are
forced to seek alternative sources of income,
Freshwater either in neighboring towns, in more successful
agricultural areas or through exploitation offorest
Supplies of drinking water are generally inad- products.
equate in the near-coastal settlement areas of the Typically the forest products that are exploited
Eastern Lowlands (Anon, 1991). The marked dry are timber, nibung, nipah,jelutung, rotan, fish and
season in the area means that water from rivers, wildlife.
drainage canals and shallow aquifers becomes
brackish in the dry season. This is partly a natural
occurrence due to reduced flows at this time of Timber
year, but it is significantly increased in places by
reduction of outflow resulting from habitat alter- In many instances sawmills associated with log-
ation and increased saltwater intrusion following ging concessions are located close to settlements.
mangrove destruction. This deterioration of water Settlers are able to sell any logs that they can
quality is exacerbated in the canal systems by use extract from forest areas to the sawmills in order
of the canals as toilets, input of organic material to supplement their incomes. According to Scholz
(e.g. household wastes and agricultural chemi- (1983) the first few years of Buginese swamp
cals), and seepage of pesticides (BCEOM, 1990). agriculture are usually financed by the sale of
Settlers cannot rely on rainwater supplies, timber. Transmigrants have also used the income
partly because of the uncertainty of rainfall dur- from the timber resource in the remaining forest
ing the latter stages of the dry season and partly blocks to finance their departure from the trans-
because few households have sufficient water migration site (J ames, pers. comm.). The result of
storage to cope with more than a IS-day dry spell. this uncontrolled exploitation is that remaining
Those people living on the larger rivers collect forest areas are often rapidly depleted, with a
freshwater up to twenty kilometers upstream consequent loss of other economic and wildlife
during dry seasons, taking advantage of the water values.
storage and flow regulation functions of the peat- In the Berbak area there are well known stan-
swamp forests in the area (Hanson & Koesoe- dard prices which will be paid by nearby sawmills
biono, 1979). Those away from rivers sometimes or their agents for logs, either at the riverbank or
have to resort to using coconut milk for drinking at the mill.
for several months each year (Giesen, pers.
comm.).
In Berbak people from the coastal villages typi- Nibung
cally collect freshwater inside the Reserve in the
dry season. This leads to management problems, In addition to the cutting of nibung for local use,
since it is difficult to determine whether people on settlers sometimes become involved in commer-
the river are there solely for the purpose of col- cial harvesting of nibung poles for export to other
lecting water. areas of the country. This can amount to a
295
significant industry, with an estimated value in the income to the community usually comes only
South Sumatra alone of Rp 400 million in 1988 from 'taxes' which may be levied on the product
(Danielsen & Verheugt, 1990). Since the harvest- by the local village head.
ing is virtually unregulated this can have a signif-
icant impact on the viability of this vegetation
type and in many places it has virtually ceased to Fish
exist.
None of the major ethnic groups in the Eastern
Lowlands normally includes professional fresh-
Nipah
water fishermen. Some groups of Buginese have
a tradition of sea fishing, but are generally not
The production of tiles of nipah thatch can re-
experienced in the different techniques necessary
present an important income supplement for set-
for river fishing. As a result, while a proportion of
tlers. An average family can earn Rp 5000 per day
the population turns to fishing in the rivers and
from this activity (1988 prices), which can be un-
wetlands of the area for supplementary income,
dertaken in the dry season when there are no
they are not usually involved in large scale fresh-
pressing agricultural activities to be undertaken
water fishing operations, though cumulative catch
(Danielsen & Verheugt, 1990). Provided simple
figures can be high. A group of fewer than twenty
guidelines are followed, harvesting of nipah fronds
fishermen took 1.5 tons of fish from a three-
should not have any significant effects on the re-
kilometer stretch of one river in Berbak in four
source, however it is an activity that brings people
weeks in mid-1991. The majority of the catch was
into the Reserve thereby providing opportunities
tapah (Wallago sp.) which has been exterminated
for other, more environmentally damaging, activi-
in some of the major river systems in the Eastern
ties.
Lowlands and should be considered as endan-
gered (Muchtar Ahmad, 1991).
Jelutung Typically, groups from outside the coastal low-
lands who have fisheries backgrounds take ad-
J elutung is the name given to the peatswamp vantage of the access provided by the settlements
forest tree Dyera costulata and also to the latex to fish the rivers. Because these are often people
which is obtained from it. This latex has a variety who have not been successful in sustainable fish-
of uses, including high quality chewing gum and ing in their original location, they frequently ini-
bubble gum and for making certain types of car- tially need to be 'sponsored' by an agent who may
ton (Tinker et al., 1982). Tapping of trees fre- be a fish merchant or, sometimes, the local village
quently results in the death of the tree after two head. These groups frequently use such unsus-
to five years, but it is not clear whether this is an tainable methods as setting permanent traps
inevitable result of tapping, or whether it is be- completely across rivers, and do considerable
cause of the practice of trying to obtain the great- damage to large areas of vegetation in clearing
est yield in the shortest time, as has been sug- living areas and collecting material to make traps.
gested by some tappers (Jelutung tapper, Sungai In Berbak these groups of fishermen usually
Benu, pers. comm.). come from the Ogan Komering lebaks2*, where
Jelutung tapping sometimes provides an addi- there is a long tradition of fishing rivers and lakes,
tional source of income for settlers, though it is but where a combination of unsuitable fishing
more frequently carried out by organised teams practices and an annual ballot system for fishing
from areas outside the Eastern Lowlands, such as rights have combined to make the fishery unsus-
the Batak people from Tapanuli area who have tainable (Giesen, 199Ib). They usually attempt to
been gathering jelutung in the eastern J ambi area
for generations (Furukawa, 1986). In such cases *A 'lebak' is an overflow wetland, usually a sedge/grassland.
296
settle inside the Reserve, but if detected and ex- line, though income may not be regular. Virtually
pelled by management staff they may settle just all parts of the tiger can be sold. In some in-
outside the Reserve boundary and make forays stances skin dealers or others willing to deal in
into the Reserve when no management patrols tiger products offer rewards for reports of tiger
are in the area. footprints, with a higher reward for reports of
territorial urine-marking sites.
In Berbak there are no reports of tiger-hunting
Rotan inside the Reserve, though tiger hunting is carried
on in the vicinity of villages adjacent to the
Rotan (rattan) is the stem of climbing palms and
boundary. Other species are hunted in the Re-
is harvested for a variety of uses, but principally
serve from time to time.
for making furniture. The material harvested usu-
The Asian Bonytongue (Scleropages formosus)
ally comes from the genera Calamus or Korthalsia.
is a rare fish with two colour forms that occur in
Before the imposition of a ban on the export of
some of the rivers of the Eastern Lowlands. This
all but finished rotan products in 1989, rotan was
fish represents a source of high income for mini-
by far Indonesia's most important non-timber
mal effort to the settlers of the region. A fully
forest product (De Beer & McDermott, 1989).
grown red specimen was sold in Singapore in
Rotan harvesting does not require a high level
1986 for US$ 2700 (Giesen, 1987). There is an
of skill, and is a common source of supplemen-
increase of fishing for the Asian Bonytongue in
tary income among settlers in the Eastern Low-
the wet season when this species breeds in the
lands. In Berbak, rotan is harvested illegally by
shallow floodwater adjacent to the rivers.
transmigrants from the schemes to the north and
Hunting of wildlife for supplementary income
west of the Reserve, as well as by teams from
is an activity which has the potential for obvious
outside the area (pers. obs.). In general, harvest-
and unacceptable impacts on the conservation
ing is done without regard to sustainability, and
value of the Reserve.
fears have been expressed for the survival of the
more popular rotan genera in the Reserve (Drans-
field, 1974). Social and cultural constraints to sustainable
management
Wildlife products Lack of awareness of legal controls
Because the settlers are moving into a recently
Settlers in transmigration areas usually have no
cleared forest environment (or they clear the for-
more than primary school education, so that it is
est themselves) and there are frequently remnant
not surprising that they generally have a low level
blocks of forest in the area, they tend to have a
of awareness of conservation legislation. A survey
close experience of the local wildlife. In addition,
in the villages adjacent to the Berbak Reserve
because some wildlife, such as tigers and bears,
showed that more than sixty percent of the
tends to seek food in the new agricultural areas,
population has a low or very low knowledge of
the settlers have good reason to want at least
the controls imposed by environmental law
some of the animals removed from the area. The
(YASIKA, 1991).
combination of these two factors with the high
price commanded by some wildlife products such
as tiger, bear, rhinoceros, and certain bird and Buginese social structure and attitudes
fish species, leads to a widespread hunting of
wildlife. Some other wildlife is hunted for its value In the Buginese villages adjacent to Berbak the
as food, for example deer and mouse deer. village head is usually the person who was the
Tiger hunting in particular is a profitable side- leader of the group which first opened the area for
297
settlement, or the son of that leader. These origi- information that needs to be collected, and oflow
nal groups were usually extended family groups. cost techniques for data collection. The lack of
The combination of the special position of the discipline and motivation among field staff for
leader of the original settlers, the cohesiveness of travelling in remote areas under field conditions
Buginese family ties, the well known independent to collect data is also a significant problem. In this
character of the Buginese and the remoteness of regard Berbak is unusual in that at one point in
these areas from central or provincial government its history it was subjected to two high quality
can lead to problems for government officials who inventories of flora and fauna adjacent to the
seek to enforce controls on natural resource ex- lower reaches of two rivers at the instigation of
ploitation. Some prefer not to try. management staff (Dliyaurohman, 1983 & 1985).
In addition there have been a number of surveys
of parts of the area, particularly those parts ad-
Administrative constraints to sustainable jacent to the larger rivers (e.g., de Wulf & Rauf,
management 1982; Silvius et aI., 1984; Blouch, 1984; Santiapil-
lai, 1989). However, the greater part of the exist-
MacKinnon & MacKinnon (1986) rated only ing information about Berbak has been collected
three of Sumatra's reserves (not including Ber- adjacent to the lower reaches of the major rivers
bak) as reaching an average standard of manage- and along the coast, and is generally inventory
ment in the mid-eighties. The remainder were data only. Information about movements of spe-
scored as having poor management or no effec- cies or life histories is generally lacking.
tive protection. However the authors were of the
view that at that time levels of staff, equipment
and funding were high for Asia, and commented Management plans
that excellent management plans existed for most
of the major reserves. In their view, shared by Berbak has had a management plan, prepared by
Petocz (1987), any deficiencies in management an FAO team (de Wulf & Rauf, 1982) since 1982.
arose from poor discipline and motivation, con- It is written in English, a language which none of
centration of staff and facilities in town headquar- the current management staff in Berbak under-
ters, and the high rate of opening of new areas in stands, and, not surprisingly, they did not know
the countryside. To this I would add the lack of of its existence at the commencement of a man-
information about the values and qualities of the agement implementation assistance programme
area's natural resources, and inadequacy of leg- by the Asian Wetland Bureau in 1991. The ma-
islation, boundary identification and demarca- jority of the measures proposed in that manage-
tion, and cross-sectoral coordination. ment plan remain relevant, but for various rea-
It is worth looking at these factors in relation sons most have not yet been implemented. Silvius
to the Berbak Wildlife Reserve as an example of et al. (1984) offered an analysis of the manage-
conservation management in the Eastern Low- ment problems and conservation needs ofthe area
lands. and made a number of recommendations for
management. These were translated into Indone-
Information sian by the head of the J ambi office of PHPA,
but most of the recommendations have not been
There is a great lack of reliable information about acted on.
the remaining areas of natural habitat in the East- The majority of other conservation reserves in
ern Lowlands. To some extent this is a product the Eastern Lowlands (other than national parks)
of the lack of resources and expertise to under- do not have management plans, and therefore the
take surveys, but it is also due to a lack of management staff are lacking a program of action
understanding by managers, both of the type of that is tailored to the values of each area.
298
Manpower the Reserve. However, the funding of running and

repair costs for these vessels is a continuing con-
Staffing at Berbak has fluctuated since the early straint on management patrolling.
1970s from one person who was responsible for
the protection of all reserves in J ambi Province to
Funding
a high of twenty in 1983, and has now dropped
back to eight, mainly as a result of dismissals of Whitten (1989) estimated that nationally PHPA
staff without primary schooling and a freeze on needed a six-fold increase in their budget in order
recruitment. The number of staff would be barely to achieve basic conservation requirements. A
adequate to maintain the integrity of the Reserve substantial funding increase is still needed in 1991
by taking a strict law enforcement approach if if the management of the Berbak Reserve is to
they had sufficient training, discipline and moti- reach an effective level.
vation. However, because of the remoteness of
the location, low salary levels and the extent of
the area to be covered with a limited number of Legislation, boundary identification and boundary
staff, it is difficult to maintain discipline and mo- marking
tivation among field staff. Ideally the level of staff-
Partly because of the transition from Dutch law
ing should be twice the current figures.
to Indonesian at Independence, and partly be-
cause of the Indonesian system under which leg-
Training islation can be made by a decision of the relevant
Minister or Governor, it can be very difficult to
The nature conservation management field staff identify the specific legislation which applies to
in Sumatra generally have no more than second- any particular topic or reserve. The same diffi-
ary schooling, and do not necessarily have any culty applies to the boundaries of reserves, where
interest in biology or conservation before joining poor or inaccurate descriptions, over-generalised
the PHP A. Training of field staff currently re- maps attached to official documents, and inabil-
ceives a low priority, though there are plans to ity to locate official documents lead to ambiguity
remedy this situation in the future. Training and confusion (Petocz, 1987).
courses are available for a limited range of sub- When compounded with a frequent failure to
jects such as law enforcement, awareness and take into account the need for ecologically mean-
Panca Sila (the State ideology), but only a small ingful boundaries which are clearly visible in the
proportion of staff are able to undertake techni- field, the result is practical difficulties for field
cal courses. In some instances individuals de- managers who have the task of maintaining the
velop an interest in the work and undertake per- integrity of the boundaries. In Berbak less than
sonal study programs, however without good five percent of the current boundary follows an
supervision field staff often are hard pressed to ecological limit of any kind, and in the one case
understand the nature of their role, let alone iden- where the boundary follows a river, no govern-
tify and follow a program of personal study. ment agency can state whether the boundary is on
one of the banks or in the middle of the channel
- a crucial distinction for field managers.
Transport
Transportation has recently been improved in Cross sectoral communication in planning and
Berbak as a result of the joint PHP AIAWB management
Sumatra Wetland Project (see below). The PHPA
now (1991) has five motor launches and six The need for increased cross-sectoral participa-
wooden canoes for patrolling the waterways of tion in planning and management of land and
299
water resources in Indonesia has been pointed an across-the-board increase in funding of PHPA
out on a number of occasions over a long period in the near future. This is not to say that high
(for example, Hanson & Koesoebiono, 1979; value areas such as Berbak may not be singled
MacKinnon & MacKinnon, 1986). In a situation out for special treatment. However there is a
where development is taking place raf>idly and strong argument that, for areas of international
where illegal land use activities are not uncom- significance such as Berbak, the international
mon, all sectors of planning and management in community should take on some of the burden of
the government would benefit greatly from im- funding routine management. This would require
proved coordination. It is a matter of historical a change of policy for most of the major aid do-
fact that the major cause of loss of environmen- nors, but it is an approach that will have to re-
tal quality in Berbak after illegal settlement has ceive serious consideration in the near future if
been activities which have been permitted by one the values of the area are to be maintained.
area of government without adequate consulta- In the short-term the joint PHPA/AWB
tion with those responsible for the management of Sumatra Wetland Project with funding from the
the Reserve (e.g.: Dept. Pertanian, 1983; de Wulf Dutch Government is in the final stages of as-
& Rauf, 1982; Ichsanudin & Purwoko, 1990). sessing the management situation at Berbak and
making an initial appraisal of the values of the
area. By the end of 1991 a comprehensive back-
Discussion - The future ground study report and management plan will be
finalised. It will remain to implement the pro-
There are a number of pressing requirements for grams set out in the management plan, a task that
the improvement of the management of the Ber- will require several years and ongoing funding.
bak Wildlife Reserve and, by extension, of the Already, as a part of the above cooperative
natural resources of the Eastern Lowlands of Project, an assessment has been made of infra-
Sumatra. An examination of programs being un- structure needs and three motor launches, six per-
dertaken in the Berbak region may suggest ap- ahu and various items of equipment have been
proaches that may be applied in other parts of the supplied to the management team. Training needs
Eastern Lowlands to address the problems out- analysis revealed that the most urgent need was
lined above. These programs are being carried out for management patrol training of field staff, and
as part of the Project to Improve Wetland Man- a ten-day course has been run by an AWB wet-
agement and Conservation in Indonesia, Part I: land management trainer with ongoing follow-up
Sumatra (generally known as the PHPA/AWB by an on-site management expert.
Sumatra Wetland Project) with funding from the
governments of the Netherlands and Indonesia.
These can be summarised under the broad head- Cross-sectoral coordination
ings of: funding; cross-sectoral coordination;
clarification of legislation and boundaries; and The problem of cross-sectoral coordination has
law enforcement. been tackled by the preparation of a regional en-
vironmental profile for the wetlands of eastern
J ambi Province, including Berbak. This profile is
Funding being prepared in cooperation with the Provincial
Regional Planning Agency (BAPPEDA) and the
An improvement in the funding situation is es- Environmental Study Centre of the University of
sential if there is to be adequate information, J ambi. A key part of the profile will be a cross-
manpower, training, motivation and discipline. sectoral planning strategy for guiding future
As stated above, given the demands on the In- decision-making. A cross-sectoral planning work-
donesian public sector there seems little chance of shop was held in Jambi in mid-1991 as part of a
300
the development of the environmental profile and minimum of skills and knowledge to carry these
was also aimed at encouraging communication out there is a good chance that effective natural
between government agencies. The proceedings resource management will result.
of this workshop (Van Der Mijn & Djak Rah-
man, 1991) have been made available to relevant
government agencies as an aid to cross-sectoral Conclusion
communication.
The problem of availability of information has Berbak Wildlife Reserve provides a good case
been partly addressed through reconnaissance study of the problems of management of coastal
surveys of the Reserve and a program of man- ecosystems in the Eastern Lowlands of Sumatra.
agement patrols to collect data, but it is clear that In their nature and extent they are probably typi-
a well-planned inventory of major wildlife groups cal of the situation in many developing countries,
is needed over the whole Reserve. The develop- particularly in areas of fairly recent settlement.
ment of a computer wetland database for the The important requirements needed to over-
whole of Sumatra (later to be extended to the come these problems (funding; cross-sectoral co-
whole ofIndonesia) is a part of the same Project, ordination; clarification oflegislation and bound-
and this will not only assist in placing the Reserve aries; and law enforcement) are probably not
in a regional perspective, but will also be able to within the power of the Indonesian Government
be used as a management tool for the Reserve. to provide within a time scale that will allow the
important values of the area to be maintained.
Outside help will be needed.
Legislation and boundaries
Ongoing efforts are being made to clarify the situ- Postscript (June 1992)
ation with regard to boundaries, and a proposal
for adoption of ecologically meaningful bound- The Indonesian Government changed the status
aries will be part of the management plan. The of the Berbak Wildlife Reserve to National Park
Department of Forestry is making funds available in April 1992 (apparently to meet a target of a
in its annual budgets for the marking of bound- certain number of national parks in the country),
aries. contrary to a recommendation in the draft man-
Legislation concerning the management of agement plan that this not be done without care-
conservation areas and wildlife populations is ful study of the effects it would have on ability to
very much need of review and rewriting to remove manage the area for the conservation of wildlife.
inconsistencies and to fill gaps in the powers that Soon afterward (June 1992) the Indonesian
are available to managers. Government terminated all Dutch aid projects in
Indonesia, including the Sumatra Wetlands
Project, so that important follow-up to the mea-
Law enforcement sures which had been proposed for a second
phase of the project were not able to be under-
None of the above measures will be effective until taken.
there is a sure and consistent system of enforce-
ment of laws. This will require as a first step
substantial increases in the salary levels of all Acknowledgements
staff in management agencies to remove the need
for them to seek supplementary income. If this is The assistance of the following persons and or-
accompanied by basic training to ensure that staff ganisations is gratefully acknowledged: the gov-
understand the nature of their duties and have the ernments of the Netherlands and Indonesia for
301
funding the PHPA/AWB Sumatra Wetland FAO, 1977. Nature Conservation and Wildlife Management:
Indonesia. Interim Report. UNDP/FAO, Rome.
Project; AI DAB for funding the author to attend
Furukawa, H., 1986. Environment and agriculture in the Ba-
the Regional Seminar on Ecology and Conserva- tang Hari River Basin of Jambi. In T. Kato, M. Lutti &
tion of Southeast Asian Marine and Freshwater N. Maeda (eds), Environment, Agriculture and Society in
Environments Including Wetlands in Kuala the Malay World (Vol. 4). Interim Reports of the Project
Lumpur; Drs Ben Zech, the survey staff of Ya- 'Transformation of the Agricultural Landscape in Tropical
Archipelagos'. Centre for Southeast Asian Studies, Kyoto
yasan Indonesia Untuk Kemajuan Desa, the Bi-
University, Kyoto: 49-86.
ology Students Association of the Padjadjaran Giesen, W., 1987. Danau Sentarum Wildlife Reserve: Inven-
University, and Drs Wim Giesen for information tory, Ecology and Management Guidelines. WWF-Indo-
gathered in Berbak and surrounding areas; Drs- nesia, Bogor.
Marcel Silvius and Drs Wim Giesen for com- Giesen, W., 1991a. Berbak Wildlife Reserve, Jambi: Recon-
naissance Survey Report. PHPA/AWB Sumatra Wetland
menting on the draft of this paper; Dr Jim Davie
Project Report No. 13. Asian Wetland Bureau, Bogor.
for detailed review comments on the text and Giesen, W., 1991b. Conservation and Management of the
useful suggestions on restructuring of the draft; Ogan-Komering Lebaks. PHPA/AWB Sumatra Wetland
and the village heads of Sungai Benu, Cemara, Project Report No.8. Asian Wetland Bureau, Bogor.
Air Hitam Laut and Simpang Datuk for their Hanson, A. J. & Koesoebiono, 1979. Settling coastal swamp-
lands in Sumatra, In MacAndrews, A. & Chia Liu Lien
patience with this bule's poor Bahasa Indonesia
(eds), Developing Economies and the Environment.
and for their cooperation with all stages of the McGraw Hill, London: 121-175.
Project. Ichsanudin, A. & F. Purwoko, 1990. Report of the Forestry
Survey Team Concerning the Berbak Wildlife Reserve. Di-
rectorate General ofInventory and Forest Utilization, For-
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Hydrobiologia 285: 303-309, 1994.
Mangrove conservation in relation to overall environmental considerations
Nelson Marshall
Professor Emiritus of Oceanography and Marine Affairs, University of Rhode Island; Adjunct Professor
Horn Point Environmental Laboratory, University of Maryland, Cambridge, MD 21613, USA;
Postal address: P.O. Box 1056, St. Michaels, MD 21663, USA
Key words: alternate uses and environmental subsidies
Abstract
The role of mangroves, as nursery and feeding areas, in the enrichment of coastal waters, in the stabi-
lization of the shoreline, and in trapping silt and wastes from upland runoff, is repeatedly being threatened
by suggestions for reclamation, whether for aquaculture, agriculture, or development projects. Propos-
als for such alternatives should only be judged after taking into account the environmental subsidies
involved, and possible losses in energy transformation steps. Assurance is needed that renewable re-
sources and other environmental capital will not be sacrificed.
My approach in this presentation is to consider From the conservation advocate's point of view
alternative uses of mangrove areas as I judge these the first alternative is generally considered ideal.
from my travels about Malaysia and to a lesser The second may be even beUer. There are various
extent in Indonesia, Thailand and the Philippines. pros and cons, and unknowns, relative to convert-
These alternatives are: ing mangroves to aquaculture ponds and crop-
land. Reclamation for non-living resources has
1. leaving the habitats in their natural state so as been relatively localized yet sometimes constitutes
not to detract from their potential in the en- a serious loss.
richment of coastal waters, from their role in As to the role of mangroves in contributing to
the stabilization of the shoreline and river coastal productivity, the most convincing infor-
banks, from their effect in counteracting the mation involves the correlations between man-
stress of inland runoff, and from their role as grove area and fisheries harvests. Jothy (1984)
nursery and feeding areas, reports on the harvest of molluscs, crustacea and
2. managing mangrove forests to optimize the fish indicating that, with the area of mangroves
potential yields of forest products while effec- along the west coast of the Malaysian peninsula
tively sustaining the values realized when man- being about five times that of the east coast, the
groves are retained in their natural state, overall fisheries of the former are about twice
3. altering or reclaiming substantial areas of the those of the latter. This differential is a bit less for
mangrove swamps for aquaculture, species not associated with mangroves. Limiting
4. similarly reclaiming mangrove areas for agri- the comparisons to shrimp (prawns) Sasekumar
culture or grazing, & Chong (1987), in state by state summaries for
5. reclaiming mangrove areas for non-living Peninsular Malaysia, show that, where the man-
resource use, for example for ports, urban set- groves are most extensive the yields of shrimp are
tings and even tourist accommodations. highest (Table 1). The general association of
304
Table I. Area of mangrove forests and the yields of shrimps listed by states in Peninsular Malaysia
State Mangrove area Yield of shrimp (mean of 4 years Yields

in hectares 1980 to 1983) in tonnes (source, kg shrimp
Fisheries Statistics 1980-1983) ha-1yr- 1
Kedah, Perlis & Pulau Pinang 9,443 14,845 1572.1*

Perak 40,497 23,596 582.7
Selangor 22,522 15,284 678.6
Negeri Sembilan 1,355 58 42.8
Melaka 77 228 2961.0**
West 10hore 17,164 5,283 307.8
East 10hore 8,454 1,690 199.9
Kelantan 22 471 21409.1 **
Terengganu 976 2,735 2820.7**
Pahang 2,573 746 289.9
Data from Sasekumar and Chong (1987).

Yield per hectare calculations by N. Marshall.
* The harvest included catches from distant locales not included in the area column.
** Mangroves are very limited; contribution to yields probably outweighed by other factors.
mangrove area and shrimp yields is also discussed In contrast to the nursery/feeding area role of
on a trans-tropical basis by Turner (1977) and by mangroves, their role in the nutrient enrichment
Martosubroto & Naamin (1977) who report on of nearby coastal waters is not consistent. As
14 provinces in Indonesia. Twilley (1985) states: 'Problems associated with
In assuming that the association between man- the measurements of these tidal fluxes have re-
grove area and coastal fisheries yields involves sulted in a controversy surrounding the hypothe-
some cause and effect relationship, it is useful to sis that intertidal wetlands affect the productivity
consider what is known of the nursery/feeding and nutrient cycles of estuarine waters'. An ex-
role of the mangrove environment and the extent tensive literature indicates the large quantities of
to which outwelling of nutrients from the man- litter produced by mangrove forests. As Twilley's
grove might be important. Anyone who has ex- (1988) summarization indicates, for fringe and
plored the mangrove and adjacent mudflats can riverine mangroves, i.e. those strips next to the
attest to the abundance of juvenile shrimps and coastline or adjacent to waterways, much of this
fishes. Chong et al. (1990) have quantified this for litter is directly exported to the coastal waters by
mangrove and mudflat sites along the coast of the sweep of the tides. As to basin mangroves,
Selangor in Malaysia (Table 2). They point out appreciably behind the fringe, some of the litter is
that, while these grounds are the nursery for cer-
tain species of shrimp, they serve more as feed-
Table 2. Biomass and density of the major species groups as
ing than as nursery areas for many of the wide- sampled in four habitats on the Selangor coast of Peninsular
ranging species of fishes that frequent them. While Malaysia.
their data do not indicate annual production, the
large biomass reported clearly suggests a very Habitat Biomass/in kg ha - 1
large contribution to coastal harvests, particularly Fish Prawn Others Total
when growth (minus mortality of course) is taken
into account. In reports on estuarine populations Creeks/inlets 17.7 3.18 0.52 21.4
in mangroves of northern Australia, Blaber et al. Mud fiats 5.96 0.77 1.79 8.52
(1983) and Morton (1990) do not offer data for Near inshore 6.06 0.23 1.32 7.61
Far inshore 3.74 1.14 1.79 6.67
shrirpp but, for fishes, their biomass data are even
larger than those reported for Selangor. Data from Chong et al. (1990).
305
incorporated in the mangrove floor, yet in almost 22_~""

every case studied there is an excess of litter and
very fine particles that is flushed out. And there SANGGAR RIVER
0--0 MATANG MANGROVE
are impressive examples of dissolved organic car- FOREST
bon being exported. - - SELANGOR RIVER
The pattern is not that simple with respect to 18
dissolved inorganic nutrients. For example Boto
& Wellington (1988), in reporting on a mangrove
system in northern Australia, where they observed
substantial export of plant litter, found that over
the year there was little net transport of nitrate 14
and nitrite to or from the system while reactive
phosphate was imported. Further, in the area they
m
studied, there was not even a net export of dis- o
z
solved organic carbon. They suggest that nitrogen +
N10
and phosphorus are conserved within the man- o
z
grove system. This is also suggested by the mix- +
ing diagram observation of Nixon et al. (1984) '"""
:J:
Z
indicating no increase in dissolved N and P off
the mangroves of the Matang Forest in western 6
Malaysia (Fig. 1).
Another insight is gained from the sampling of
the various forms of dissolved nitrogen done by
Thong et al.(ms. in prep.) on the Sementa Besar,
a mangrove creek near Klang in western Malay- 2
sia. The values were high in the rivulets running
off the floor of the mangrove, and either high
amplitude tides or heavy rainfall augmenting the 10 20 30
SALINITY (%0)
drainage from the soils of the mangroves greatly
increased inorganic nutrient levels in the creek. Fig. 1. Mixing diagram of Nixon et at. (1984) showing no
Even so, for two 24 h tidal cycle observations, increase in NH 4 , N0 2 , N0 3 along a salinity gradient in the
Matang Mangrove Forest Reserve.
plus sampling of less duration, there was no evi-
dence of net export from the mouth. There had
not been any appreciable rainfall to augment grove detritus as found in their guts. Such a re-
flushing from the creek coincident with this sam- fractory state was also suggested by Alongi (1990)
pling, otherwise the results might have been dif- from chemical analyses of mangrove detritus in
ferent. coastal sediments.
In essence there is no question that, in macro- On the other hand it is not clear that dissolved
litter and in fine particles, carbon and the nutri- inorganic nutrients are exported, perhaps under
ents contained are exported from mangroves, some conditions 'yes'; under others 'no'. Twilley
more from the fringe areas than from those be- (1988) emphasizes that nutrient enrichment from
hind. This probably enhances coastal productiv- the mangrove to the coastal waters tends to be
ity though, by the time the detritus reaches the site specific, depending on the hydrology and geo-
coastal waters, some of it may be highly refrac- morphology of the setting.
tory. This was suggested by Rodelli et al. (1984) As this information is considered collectively,
who, using stable carbon isotope analyses, found one point seems obvious, namely the outward
that offshore shrimp did not assimilate the man- migration of the fisheries species that use the
306
mangrove environment as nursery and feeding As a first step in considering the alternative of
areas is far more important than outwelling in converting mangrove area to aquaculture it is use-
augmenting production realized in coastal waters. ful to ask what might be lost in terms of harvests
Second on the list of alternative practices are along adjacent coasts. Toward this end it is useful
the mangrove areas suitably managed for a sus- to discuss in more detail two papers with infor-
tained yield of forest products. For these there is mation relating to pond culture in Indonesia.
relatively little to add, especially since the Matang Martosubroto & Naamin (1977) compared
Forest, an excellent example discussed by Ong shrimp harvests with mangrove area for 14 prov-
(1982), contributes to the large fishery off Perak:. inces of Indonesia (Table 3). Looking at the vari-
If anything, the turnover in forest growth probably ability in the shrimp per hectare ratios and asking
constitutes an improved utilization of the sun's which yields represent the full fisheries potential,
energy and, what's more, a crop is realized for I am prone to rule out all but five, assuming that
poles and for charcoal. Obviously, where harvest- harvests far less than 100 kg ha - 1 y- 1 represent
ing is carried out without a systematic means to either undeveloped fisheries or provinces where
replenish the tress, as may occur in the wood chip the loss of mangroves has already adversely af-
industry in Sabah, this can amount to a substan- fected the yields.
tial loss. This reasoning is supported by the analysis of
Table 3. Area of mangrove forests and the yields of shrimp in 14 provinces of Indonesia.
Area/Province Mangrove area Annual yield of shrimp Yields

in hectares in tonnes* kg shrimp ha - 1 yr- 1
Sumatra
Aceh 129,375 1,643 34.1
North Sumatra (eastern part) 81,250 7,929 97.6**
Riau 348,750 11,875 34.0
Jambi 67,500 1,138 16.8
South Sumatra 102,500 710 6.9
Lampung 6,875 1,159 168.6**
Java
North & east 58,700 5,214 88.8**
South coast 26,250 4,308 164.1**
Kalimantan
West & central 181,250 7,457 41.1
South 68,125 1,805 26.5
East 447,500 2,675 6.0
Sulawesi
North 37,500 398 10.7
South 58,750 4,845 82.5**
Maluku
Aru 70,625 4,642 65.7
Irian Jaya
Irian Jaya 898,125 10,765 12.0
* Martosubroto & Naamin (1977) refer to various national and provincial fisheries statistics sources.
** Interpreted by N. Marshall as approximating the yields expected from a fully developed fishery - see text.
307
Turner (1977) who only used data from stabilized profitably, is provided by H. T. Odum & Arding
and developed fisheries to derive, on a trans- (1991). These authors point out that, when ad-
tropical basis, the formula; equately analyzed, such culture is not in the best
interests of the country's economic and ecologi-
j = 158.7e - 0.Q70(x),
cal well-being. They put considerable emphasis
where y is kg schrimps/hectare/year and x is de- on the subsidies necessary for success in such
grees of latitude between 0 and 35 and I sug-
0 0 advanced aquaculture including the obvious, i.e.,
gest that, for simplification, the exponent be ig- fertilizers, food, labor, etc., and the not-so-
nored inasmuch as the Indonesian sites are near obvious, particularly a wide range of environmen-
the equator. Turner's specific entries for Indone- tal inputs such as tapping natural sources for
sia, contributing to this derivation, were 83.2 and shrimp stock and nutrients and depleting the en-
86.8 kg ha - 1 for South Java and Sumatra respec- vironmental capital of the mangrove ecosystem.
tively. Since these are less than the formula would They compare the energy required to realize pro-
indicate, it seems possible that they represent har- duction in ponds with the more efficient energy
vests already adversely affected by the depletion transformations of the coastal fisheries that must
of mangroves. These comments and Turner's suffer from the depletion of mangroves and ac-
equation seem to suggest that the two high figures companying loss of natural habitat. As to the
of Martosubroto and N aamin, approximating export trade, which is the major stimulus for
160 kg ha - 1, come closer to representing the po- shrimp mariculture, they calculate that the prod-
tential coastal shrimp harvests in Indonesia. ucts are going overseas with much less real wealth
While this is high, one may wonder why the kg returning. Odum & Arding refer to their compre-
ha - 1 ratios for Selangor and Perak in Malaysia hensive approach as an emergy evaluation and,
(Table 1) are almost four times as great. This while for this purpose they are compelled to forge
might be accounted for by realizing that, in plac- ahead with inadequate data in some respects, they
ing all the emphasis on the mangrove relation- direct attention to many factors commonly over-
ship, other contributing factors are being overlooked in simplified cost/benefit analyses.
looked. Such an oversight is suggested in Table 1 Finally it must be noted that aquaculture in
wherein several states, having modest mangrove mangroves, though often successful in terms of
areas and small yields of shrimp, nevertheless immediate returns, can fail completely as the high
have a high kg ha - 1 ratio. organic and iron content in some mangrove soils,
For a comparison with aquaculture Turner and the ever present sulfate from tidal seawater,
(1977) points out that the yield from fish ponds results in acid sulfate conditions when oxidized
in Indonesia, about 287 kg ha - 1, is about 480 kg through exposure in pond construction and op-
less than the calculations he presents for the eration (see for example, Ong, 1982). In some
combined harvests of fish and shrimp in coastal areas it has been possible to counteract this with
waters that are enriched by the mangroves. the addition of lime and careful manipUlation of
Though in this paper Turner doesn't establish that flushing and water levels. Where this proves to be
the coastal yields are actually less where pond uneconomical, reclamation for pond culture has
culture is practiced extensively, Turner & Boesch either been abandoned or, more wisely, never at-
(1987) cite several cases where it is demonstrated tempted.
that reclaimed wetlands have resulted in declin- As to the agriculture alternative, such use often
ing shrimp yields. seems especially wasteful. The mangrove soils,
This brings us to the growing interest in pond with their high iron, sulfide and salinity, are un-
culture for shrimp, as contrasted with fish, and favorable to many crops (Law, 1984) and, unlike
the extensive use of subsidies in such culture. An aquaculture, the coastal physiographic features
evaluation of how this works out in Ecuador, are usually of little advantage. Since crops are
where practiced very widely and seemingly very primary producers, the yields per hectare are likely
308
to be greater than in pond culture for shrimp or It is noteworthy that the pressure to convert
fish; however, as Turner & May (1977) show for mangrove to pond culture or to cropland is great-
rice, the lesser protein content offsets this advan- est where the population is most dense, for ex-
tage. An even further alternative, reclaiming man- ample along the north coast of Java or in the strip
grove areas to use for grazing, seems even more between Klang and Selangor in Malaysia. If the
wasteful in view of the production inefficiency of sacrifice of mangroves is a net loss when all con-
the cattle, goats or sheep as the case may be. siderations are weighed, this may be looked upon
Reclamation for either aquaculture or agricul- as a serious example of the adverse impact of
ture raises a very practical question; namely how population pressure. This takes a toll not only on
much mangrove is needed to provide for the full the fisheries potential but also on the overall via-
carrying capacity and yield potential of the adja- bility of the environment, with more industrial
cent offshore waters. Just as the data available and domestic pollution, more CO 2 and acidity in
suggest that 'the more mangrove area the greater the atmosphere, less wooded area, etc. As
the yields', they might be interpreted as suggest- E. P. Odum (1983) points out, ' ... the urbanized
ing 'the less mangrove area the lower the yields'. or fabricated environment is 'parasitic' on the life-
One might suggest, for example, that were it not support environment (natural and domesticated)
for reclamation activities which have sacrificed for basic biological necessities .... Much more can
about 30% of the mangroves along the Selangor be done to ... reduce the stress of high energy and
coast, both the forested area and the total yields densely populated 'hot spots'. In applying this
would be closer to those of Perak; however, the concept to coastal management, my view is that
information needed, in terms of reliable catch mangroves are obviously natural life-support en-
records of the past, a knowledge of the compara- vironments. Aquaculture ponds and agriculture
tive fishable areas, the carrying capacities, and the generally are not equally supportive.
effects of fisheries management practices, are in- Since there is considerable variation in the set-
sufficient to extend this beyond mere speculation. tings and circumstances in which alternate uses
But it seems doubtful that the mangrove/yield re- are or may be practiced and since a number of
lationship is limitless. There may be some level at unknowns and assumptions are involved when
which the extent of the coastal mangroves is in comparisons are made, it is impossible to offer
excess, that is additional mangrove area will not simple answers to the management questions that
increase the yield of the adjacent fishery. arise. Inasmuch as the mudflats, inlets, creeks
The foregoing considerations lead us to dealing and fringe habitats associated with mangroves
with situations wherein we reclaim mangrove are obviously so valuable as nursery areas for
tracts in favor of port development, housing and shrimp and feeding areas for fish, the extent to
even tourist facilities. The development of Klang which any alternative detracts from this role
and Port Klang on the west coast of Malaysia amounts to a serious loss in coastal fisheries. Less
amounts to a substantial sacrifice of mangrove clear is the possible loss in coastal productivity
area; yet it may be that, with the vast expanse from a possible decrease in nutrient exports where
of mangroves seaward of Klang, the loss has mangroves are sacrificed. Where alternatives se-
not greatly diminished the nearby coastal riously undermine auxiliary functions of the man-
productivity. I hasten to add that this example groves, such as stabilization of the shoreline and
may lead to undue complacency. I would not as river banks, moderating inland runoff and even
casually dismiss, for example, the mangrove loss absorbing pollutants, destroying wildlife habitat,
in Singapore, where almost the entire coastline and decreasing biodiversity, they constitute a se-
has been converted to urban and associated de- rious loss. The composite of such effects, not
velopment, and along the east coast of Malaysia, elaborated upon directly in this discussion, con-
where there is so little mangrove yet some has tributes to what E. P. Odum (1983) refers to as
been sacrificed for harbors. the natural life-support environment, too readily
309
sacrificed due to development and population grove Ecosystem. Proc. Workshop on Productivity of the
pressures. Mangrove Ecosystem: Management Implications. Univer-
sity Sains Malaysia, Penang: 121-128.
Finally, in management planning and decision Martosubroto, P. & N. Naamin. 1977. Relationship between
making, it is important to weigh all factors. En- tidal forests (mangroves) and commercial shrimp produc-
vironmental subsidies and the relative efficiencies tion in Indonesia. Marine Res. in Indonesia 18: 81-86.
in energy utilization must be thoroughly consid- Morton, R M., 1990. Community structure, density and
ered. And there must be assurances that renew- standing crop of fishes in a subtropical Au~tralian man-
grove area. Mar. Bio!. 105: 385-394.
able resources and other environmental capital, Nixon, S. W., B. N. Furnas, V. Lee, N. Marshall, J. E. Ong,
as needed for the future, will not be sacrificed. C. H. Wong, W. K. Gong & A. Sasekumar, 1984. The role
Policy makers should be encouraged to recognize of mangroves in the carbon and nutrient dynamics of Ma-
costs and subsidies that they may tend to over- laysia estuaries. In E. Soepadmo, A. N. Rao & D. J. Ma-
look. cintosh (eds). Proc. Asian Symp. on the Mangrove Envi-
ronment - Research and Management. University of
I gather that the objective of this symposium is Malaya, Kuala Lumpur: 534-544.
to sharpen our approach to marine environmen- Odum, E. P., 1983. Basic Ecology. Saunders College Pub-
tal problems and I hope my discussion has con- lishing, Philadelphia.
tributed to that end. I also hope that it is appar- Odum, H. T. & J. E. Arding, 1991. Emergy analysis of shrimp
ent that the focus on alternate uses of mangroves mariculture in Ecuador. Prepared for the Coastal Resources
Center, Univ. of Rhode Island, 114 pp.
is representative of the considerations to be faced Ong, J. E., 1982. Mangroves and aquaculture. Ambio. 11:
in coping with coastal freshwater and wetlands 252-257.
management in general. Rodelli, M. R., J. N. Gearing, P. J. Gearing, N. Marshall &
A. Sasekumar, 1984. Stable isotope ratio as a tracer of
mangrove carbon in Malaysian ecosystems. Oecologia 61:
326-333.
References Sasekumar, A. & V. C. Chong, 1987. Mangroves and prawns:
further perspectives. In A. Sasekumar, S. M. Phang &
Alongi, D. M., 1990. Effect of mangrove detrital outwelling on E. L. Chong (eds). Proc. 10th Annual Seminar Malaysian
nutrient regeneration and oxygen fluxes in coastal sediments Society of Marine Sciences, University of Malaya, Kuala
of the Central Great Barrier Reef Lagoon. Estuar. coast. Lumpur: 10-22.
Shelf Sci. 31: 581-598. Thong, K .. L., A. Sasekumar & N. Marshall, 1992. Nitrogen
Blaber, S. J. M., D. T. Brewer & J. P. Salini, 1989. Species concentrations in a mangrove creek with a large tidal range,
composition and biomass of fishes in different habitats of Peninsular Malaysia. Hydrobiologia (in press).
a tropical northern Australian estuary: their occurrence in Turner, R. E., 1977. Intertidal vegetation and commercial
the adjoining sea and estuarine dependence. Estuar. coast. yields ofpenaeid shrimp. Trans. am. Fish. Soc. 106: 411-
Shelf Sci. 29: 509-531. 416.
Boto, K. G. & J. T. Wellington, 1988. Seasonal variations in Turner, R. E. & D. F. Boesch, 1987. Aquatic animal produc-
. concentrations and fluxes of dissolved organic and inor- tion and wetland relationships: insights gleaned following
ganic materials in a tropical, tidally-dominated, mangrove wetland loss or gain. In B. Hook (ed.), Ecology and Man-
waterway. Mar. Ecol. Prog. Ser. 50: 151-160. agement of Wetlands: 25-39.
Chong, V. C., A. Sasekumar, M. U. C. Leh & R. D'cruz, Turner, R E. & N. May, 1977. An alternative evaluation of
1990. The fish and prawn communities of a Malaysian the fisheries value of tropical and subtropical wetlands.
coastal mangrove system, with comparisons to adjacent Proc. Fourth Internat!. Symp. on Tropical Ecol. 3: 836-
mudflats and inshore waters. Estuar. coast. Shelf Sci. 31: 852.
703-722. Twilley, R R, 1985. The exchange of organic carbon in basin
Jothy, A. A., 1984. Capture fisheries and the mangrove eco- mangrove forests in a Southwest Florida estuary. Estuar.
system. In J. E. Ong & W. K. Gong (eds), Productivity of coast Shelf Sci. 20: 543-557.
the Mangrove Ecosystem. Proc. Workshop on Productiv- Twilley, R R., 1988. Coupling of mangroves to the produc-
ity of the Mangrove Ecosystem: Management Implications. tivity of estuarine and coastal waters. In B. O. Jansson
University Sains Malaysia, Penang: 129-141. (ed.), Coastal-offshore Ecosystem Interactions. Lecture
Law, W. M., 1984. Productivity of the mangrove ecosystems: Notes on Coastal and Estuarine Studies 22. Spring~r
management implications for agricultural crops. In Verlag, Berlin: 155-180.
J. E. Ong & W. K. Gong (eds), Productivity of the Man-
Hydrobiologia 285: 311-322, 1994.
Integrated planning and management of freshwater habitats, including

wetlands
P. R. Burbridge
Department of Environmental Science, University of Stirling, Stirling FK9 4LA, United Kingdom
Key words: wetlands, functions, planning, management, sustainable development
Abstract
Freshwater habitats playa very important role in sustaining human activities. Natural functions of
wetlands, and other freshwater habitats, generate a wide array of resources that directly or indirectly
support the economic and social welfare of diverse groups of people. This role is being seriously
weakened as a result of inappropriate planning and management approaches which fail to maintain the
functional integrity of the freshwater ecosystems with the result that the flow and quality of resources
is degraded. This paper illustrates some of the major functions of wetlands and presents a case for
developing integrated planning and management practices that protect the health and productivity of
freshwater wetlands and seek to optimise the sustainable use of the flows of resources they generate.
Introduction our fundamental approach to the development of

wetlands from one where we seek to convert them
Freshwater habitats, including lakes, rivers and to alternative uses, to one where we protect the
wetlands, are among the Earth's most complex functional integrity of their ecosystems and con-
environments. In a paper of this nature it is not serve the flow of resources they generate. This
possible to discuss the management of all fresh- means that we have to develop strong principles
water habitats. Instead, the author has chosen to and practical guidelines for the planning and
focus on wetlands and to discus the role inte- management of:
grated planning and management in optimising
1. Land and water uses surrounding wetlands to
the social and economic benefits that can be de-
protect environmental processes that are es-
rived from the flow of renewable resources gen-
sential to the maintenance of the health and
erated by freshwater ecosystems.
productivity of the wetland ecosystems; and
Developers often speak of enhancing the value
2. Activities that depend upon the flow of wet-
of wetlands by converting them from 'wastelands'
land resources for their sustained develop-
to 'productive lands' for the cultivation of crops
ment.
of plants (rice) or animals (shrimp). Experience
has shown us that this is a misguided approach These sum of all the biophysical economic and
because there can be very severe economic, bio- social values that can be derived from the func-
physical and social consequences associated with tions of wetland ecosystems and the sustainable
man's interference with wetland ecosystems. To use of the flow of renewable resources they gen-
avoid these adverse impacts, we need to change erate is generally far superior to the benefits that
312
can be gained from converting wetlands to maxi- Production

mize their use for alternative forms of develop-
ment. Unfortunately, most planners are never Primary and secondary production is a key func-
trained to manage natural systems. As a result, tion that sustains the populations of plants and
most development plans indicate the allocation of animals in wetlands and in associated ecosystems
land and water resources amongst different com- (Maltby, 1991). Wetlands are a major source of
peting activities and pay little attention to the food plants, such as Sago, and non-food plants,
optimization of the economic and social benefits such as reeds used for thatching or herbs used to
that could be gained from the integration of dif- prepare medicines. They provide major grazing
ferent activities that can be sustained through the areas for domesticated animals and habitats and
multiple use of renewable resources. food for waterfowl and other wild animals. They
This paper outlines four principal themes cen- also provide important feeding, spawning and
tral to the planning, management, and wise use of nursery areas for fin and shell fish that spend part
wetlands and other freshwater ecosystems, of their life cycle in lakes, rivers, coastal waters,
namely: or the marine environment.
1. A basic requirement of sustainable utilisation
of wetlands is the maintenance of the environ-
Storage
mental processes that maintain the health and
productivity of the wetland ecosystem;
Wetlands act as a permanent sink for materials
2. The management of all wetlands should be
such as sediments and organic carbon, and store
approached from the perspective of managing
other materials, such as water, on a temporary
the wetland as an ecosystem, not from the
basis. The value of these storage functions is in-
perspective of maximizing the use of selected
creasingly recognised as an important feature of
attributes for exclusive forms of activity;
hydrological functions of watersheds such as
3. The maintenance of the natural functions of
flood control, geomorphological processes such
wetlands must be a primary objective applied
as control of coastal erosion, accumulation of
to the planning and management of activities
organic carbon (Maltby, 1991; Immirizi et al.,
that could influence the health and productiv-
1992), and the maintenance of genetic resources
ity of wetlands;
(James, 1991).
4. Policies directed towards the wise utilisation
Weare only beginning to appreciate the full
of the flow of the renewable resources gener-
range and importance of the storage functions
ated by wetlands should promote multiple-use
provided by wetlands. Often the value of these
management of these resources to optimise the
functions is only realised once a wetland has been
benefits to society.
converted to another use, functions are lost, and
we are exposed to adverse environmental, eco-
nomic and social impacts such as increased floods
The case for managing wetlands as ecosystems to and water shortages.
sustain the resources they generate Our understanding of the importance of such
functions is constantly improving. Until 30 years
Wetlands perform a wide variety of functions or so ago, it was generally believed that peat did
that benefit man. These have been reviewed by a not form in the tropics. We now know this to be
number of authors (Gosselink & Turner, 1978; totally untrue and are beginning to realise the
Burbridge, 1984; Mitsch & Gosselink, 1986; organic soils and peats in wetlands form one of
Hollis & Maltby, 1987; Maltby, 1991). Some of the largest reservoirs of carbon on earth (Maltby,
the most important functions are summarized in 1991, Immirizi etal., 1992). With the conversion
Table 1 and in the following paragraphs: of temperate and tropical peat wetlands to agri-
313
Table 1. Summary of the most Important functions of weflands
Role Elements Function Importance to Unwise use

tum:J1B!
Producer
Production Food, materials Harvest of food Overgrazing
of plants and habitat timber thatch Overexploitation
for migratory and fuel. Science Drainage. Excess
species and grazing Recreation dry land or other
agricultural uses
Animal Fish shell-fish Harvest and overexploitation
production grazing and fur- farming Excess conversion
bearing animals Habitat degradation
Organic matter Nutrient cycling Plant growth Drainage
Methane production Fuel Desiccation
Store/sink
Lake deposits Sediment Raised soil Channelization
deltas deposition and fertility Excess reduction
floodplains detention Improved water of sediment
quality downstream throughput
Lakes swamps Bio-chemlcal Natural filter Destruction of

& marshes self-purification for contaminants the ecosystem
Nutrient Treatment of Over-loading
accumulation organic wastes of the system
pathogens
Rare,threatened Genetic diversity Gene pool Excessive or
or endangered Recolonization Science/Education uncontrolled
plant & animal source Tourism harvesting, damage
species andlor Recreation Removal of
communities Heritage Pollution
Peat Nutrient Fuel Paleo- Drainage

contaminant environmental data Harvesting faster
and energy store Horticultural use than accumulation
Habitat support Heritage Destruction
Water storage Medicinal products Lowering the
watertable
Pathway
Terrestrial Food chain support Food production Interruption or
nutrients water Water supply abnormal change
and detritus Habitat support Waste disposal ofOows
Pollution
Tidal exchange Food chain support Fish, shellfish

of water Habitat support and other food Barriers to
detritus and Nursery for production flow
nutrients aquatic organisms Waste disposal Dredge and flU
Animal Support for Harvest Overexploitation

populations migratory species Recreation Interruption of
Ineluding fish and Science migration routes
shellfish Obstruction
Habitat degradation
Buffer
Water bodies Flood attenuation Reduced damage Filling and
vegetation to property reduction of
soils and and crops storage capacity
depressions Detention and Food production Removal of
retention of Improved water vegetation
nutrients quality and Drainage
flood protection
Ground-water Water supply Reduction of
recharge and Habitat maintenance recharge rates
discharge Effluent dilution Overpumping
River fisheries Pollution
Navigation
Water bodies Local and global Equable climate for Desiccation
and peat climate stabilization agriculture and Drainage
people
Soutee: Modified from a table prepared by Maltby, 1991; Hollis and Maltby, 1987.
314
culture, there has been a shift in the balance of wetlands in controlling flood damage has been
carbon and this may have a major impact upon calculated at $US 1.2 million per year for one
global climate (Armentano & Menges, 1986; watershed in the United States (Sather & Smith,
Maltby, 1991). Concern over the potential effects 1984).
of releasing more carbon to the atmosphere is Coastal wetlands serve as a buffer to coastal
leading scientists to urge that a goal of develop- storm surges and winds (Hamilton & Snedaker,
ment planning is 'the maintenance or enhance- 1984). When these wetlands are destroyed, the
ment of wetland carbon storage' (Armentano & risk to human life and the economic welfare of
Menges, 1986). coastal communities increases dramatically. In
effect, poor planning and management can turn a
natural event into a human disaster (Wijkman &
Filtration and cleansing Timberlake, 1984).
The functions performed by wetlands are of
The biogeochemical dynamics of wetlands also importance to the health and welfare of local
playa major role in filtering and cleansing water, communities (McCormick, 1978), nations and
for example the removal of toxic material from the the earth itself. All too often we learn about what
water column (Richardson, 1985). These func- we have lost by poor planning and management
tions are considered of great potential economic of development associated with wetlands once
value in providing tertiary treatment of sewage irreversible changes have taken place and disas-
and reducing pollution from agricultural wastes ter strikes. Those changes that can be reversed
(Maltby, 1991). cost us valuable time and money that could
be better spent designing more sustainable uses
of the resources available to us to meet human
Pathways or linkages among ecosystems needs.
The movement of water, nutrients and organic

materials between wetlands and other ecosystems What is meant by integrated planning and
is essential to the maintenance of food chains, management of freshwater habitats, including
migration routes, other pathways, and environ- wetlands
mental linkages that support the productivity and
health of the ecosystems and renewable resources Some definitions to ensure that we are all talking
man depends upon. Recognition of the impor- about the same things:
tance of these pathways/linkages has lead to the
passage of the RAMSAR Convention to protect
wetlands throughout the globe. Integrate
Integrate means to bring different elements to-

Buffer gether into a whole. In every day life we integrate
things in a common sense manner. For example,
Wetlands provide a number of buffer functions when we cook a meal we integrate an appropri-
that help to protect life, property and the economy ate mix of ingredients in a logical sequence. There
oflocal communities and nations. Wetlands help is no reason why we cannot conduct private busi-
to regulate the rates of surface water flow and ness and public affairs in the same logical man-
groundwater recharge within catchment areas. ner. A plan to build a paper mill will require the
This reduces flood water peaks and regulates base integration of wood supplies, access to power
water flows in rivers. The value of these hydro- supplies, reliable supplies of unpolluted water, se-
logic buffer functions is very real and the value of lection of a site with good access to sea, rail and
315
road communications, and a source of labour in associated with mismanagement of wetlands also
an area free from major environmental hazards. need to be considered. For example, conversion
of wetlands can increase the incidence and sever-
ity of flooding downstream with direct costs im-
Integrated Planning and Management (IPM) posed in terms oflost agricultural production and
damage to property.
The term integrated planning and management 3. Socio-cultural factors, including the role of
means combining assessments of the resources wetlands and their resources in protecting the
available to meet stated objectives, formulation of welfare of different individuals or social groups,
a strategy or plan of action to use the resources for example protection from floods or mainte-
in a wise way, and the implementation of the nance of food security. Cultural issues can also
strategy in a orderly and efficient manner. refer to the historical or religious significance of
These tasks cannot be carried out effectively wetlands to the beliefs of different social groups.
without the interaction of a number of well trained
persons from different disciplines. It is very im- These three sets of factors are inter-related, and
portant that the people responsible for these tasks must be incorporated into the planning and man-
are properly trained in interdisciplinary ap- agement process whether the wetlands and other
proaches to planning and management. Each of freshwater habitats are the responsibility of the
the specialists from the different disciplines must public or private sector. Figure 1 demonstrates
also be given basic training in interdisciplinary two levels of interaction, namely:
approaches to resource assessment, environmen-
1. Interactions between two sets of factors, see
tal planning, environmental management and
the light shaded area. For example the eco-
other tasks which contribute to the IPM process.
nomic dependence of a rural community on
the capture of wild stocks of fin or shell fish
sustained by a mangrove.
What do we need to integrate?
2. Interactions among all three sets of factors.
The dark shaded area covering the points at
When we consider the sustainable utilization of
which the three sets of factors overlaps repre-
wetlands and other freshwater habitats we need
sents the need to integrate the factors and the
to think very clearly about what needs to be
dynamic relationships among factors into the
integrated into the planning and management
arrangements that govern their development.
There are three basic sets offactors that we must
consider, namely:
1. Bio-physical factors or ecological process that
are essential to the maintenance of the health and
productivity of the wetland ecosystem, including
the main flows of materials into the wetland, the
structure and interrelationships among compo-
nents of the ecosystem, the environmental func-
tions that the ecosystem performs and the flow of
materials and resources from the wetland
2. Economic factors, including the environmental
and economic goods and services generated by
the wetland, and the individuals and economic
groups that directly or indirectly benefit from the F,g. 1. Three baSIC sets of factors that must be consIdered in
sustained flow of those goods and services. Costs the sustamable utiilzatlOn of wetlands.
316
evaluation of the importance of wetlands. The and the fisherman represents the Market Value.
dark shaded area also demonstrates the range An example of an Environmental Service would
of interests that need to be considered when be the protection of a farmer's rice crop from
planning the development of a wetland or storm damage by the buffer function of a coastal
activities or areas where there will be an influ- forest. Putting a realistic value on this service
ence on the wetland system. presents a problem. Nobody can make the farmer
pay for the service, none the less the service has
Defining the essential factors within these three a value equal to at least the value of the crop that
groups which need to be brought together in an is saved from storm damage. The economist must
integrated plan and management strategy is a be alerted to the fact that the environmental ser-
complex task. To do it effectively, we need to vice exists by the ecologist, and then he/she must
bring together skilled people from different disci- then find a way of representing its value.
plines to extend the frontiers of traditional disci- Some of these goods and services are harvested
plines so that they provide better analyses of the On-site, others are harvested at some location
interrelationships among factors. well away from the wetland itself (Off-site). This
A good example is the improved explanation of demonstrates the ecological point about environ-
the economic value of a wetland that has been mental and economic linkages extending well be-
achieved through ecologists and economists yond the territorial limits of a wetland. It also
working together. Figure 2 demonstrates that a demonstrates to the economist that the boundary
wetland generates both direct and indirect mar- for the economic evaluation of a wetland and its
keted and non-marketed goods and services. resources has to be very broad in order to en-
Some of these are Environmental while others are compass the full set of social and economic groups
Economic goods and services. The basic differ- that benefits from the flow of wetland resources.
ence between environmental and economic goods The flow of economic and environmental goods
and services is whether they are exchanged via an and services can sustain a wide cross section of
established market (Marketed versus Non-Mar- activities. Some of these activities are entirely
keted). An example of a Marketed good would be dependent upon wetlands for their economic
a piece of wood cut from a forest and sold locally viability. Others can make use of alternative re-
in a market to a fisherman to make a fish trap sources from other environmental systems to sus-
pole. The price agreed between the wood cutter tain themselves. When we search for ways of di-
VALUATION ON-SITE OFF-SITE

OF GOODS
AND 1 2
SERVICES Capture of fish downstream
Fuelwood or food collected
from a wetland that were
MARKETED and sold in a local market
born in the wetland and then
sold in a local market
3 4
Food, medicinal herbs Flood mitigation downstream
NON or wood products collected maintenence of base water
MARKETED and used without payment, flows in river systems,
water storage, maintenence of bio-diversity
purification of water
Fig. 2. Location of goods and services.

317
versifying and expanding local and· national This brings us to the next issue, the integration
economies, it is critically important to assess the of policy, plans and management strategies
dependence of these existing activities on wet- affecting wetlands.
lands and the role they can play in development.
Through discrete improvements in management
many freshwater resource dependent activities Functional analysis - the practical basis for
can play a very important role in improving the integration
economic and social welfare. For example the
intensification of pond aquaculture. New activi- The fundamental starting point for developing an
ties can also be developed to make use of un- integrated approach to the planning and manage-
derutilised resources, or through the more effi- ment of wetland ecosystems is the assessment of
cient use of the flow of resources, maintaining the functions of different wetlands and the stream
their quality and then extending their use through of environmental goods and services they gener-
recycling. This form of multiple use of specific ate. These need to be very carefully evaluated
resources can be demonstrated by the use of the before any decisions are taken that could in any
same fresh water for hydro-electric generation, way alter environmental processes that maintain
ground water recharge, irrigation, industrial pro- the functions of the wetland. However, we con-
cessing, cooling of thermal power plants, and stantly come up against major contradictions
maintenance of the salinity regime of estuaries. among different policies, management strategies
All these activities can be sustained if we protect and incentives for implementing different activi-
the flow and quality of the water throughout the ties which largely ignore the functions of wet-
different stages of its use. lands. For example, in Europe one agency of the
Maintenance of the environmental services European Commission is busy giving grants for
provided by wetlands can also help to reduce the the drainage of wetlands under the Common
risk of natural hazards, such as floods and Agriculture Policy, another agency is working
droughts, and protect the economic viability of hard to have wetlands conserved under the
development. Maintaining the hydrologic func- European Commission Habitat Directive. Clearly
tions of wetlands is of critical importance to food we need to improve the integration of policies and
security, the supply of essential water resources development incentives. The most logical basis
for the expansion of economic activities, and for this integration is a clear understanding of the
the reduction of the financial burden on public alternatives available, and the positive and nega-
authorities in providing flood protection. This last tive features of different alternative uses of wet-
point was clearly demonstrated by the US Corps lands resources.
of Engineers in their study of the flood storage Interdisciplinary approaches to wetlands man-
functions of a watershed in Massachusetts (see agement can provide a rational basis for devel-
above). oping strategies for the allocation of wetland re-
Without cooperation among disciplines, the sources, plans for their utilisation, or management
ecologist will find it hard to translate hard science strategies to implement their sustainable develop-
concerning wetland functions and resource val- ment.
ues into understandable terms that administra-
tors, planners and managers will respond to. Rationalising our basic approach to the allocation
Similarly, without the ecologist's knowledge, the of wetlands and other freshwater habitat
economist will create an economic model repre- resources and the planning and management of
senting the resources of a wetland which is in- their use
complete. As a result, the model will represent
only a part of reality and could be very mislead- The transition from Bio-Physical considerations
ing in terms of the values calculated. to Policy formulation, Resources Allocation, and
318
then Planning and Management of activities is limits of authority between different administra-
by no means easy. We all have practical experi- tive units. Such divisions of authority generally
ence in trying to explain the complex nature of make it difficult to integrate different policies and
wetlands and their value to administrators and priorities of political entities which share com-
planners. While it is sensible to recognise these mon hydrologic systems.
problems, they should not stop us from trying to Figure 3c illustrates the division of the wetland
communicate more effectively and make people catchment area between two administrative units.
more aware of good wetlands management prin- Ifthe administrative units work together and their
ciples and practical procedures for improving jurisdictions continue to follow the course of the
plans for their sustainable utilisation. river system, then there is potential for control
One of the most important points we need to over activities upstream and downstream. It is
get across is that the sustainable development of also possible to promote the protection of the
wetlands resources can only be achieved through environmental linkages between the wetland and
the maintenance of the flow of goods and services other ecological systems downstream. For ex-
generated by the wetland system. To achieve this ample, an estuary. However, it is often very dif-
we must shift the emphasis in planning and man- ficult to get two different political/administrative
agement away from the allocation of wetlands units to adopt unified management plans for
and other habitats amongst competing land uses shared resources without some form of unified
and move towards the optimization of the mix of policies and management strategies which both
activities that can be sustained without unaccept- must adhere to.
able changes in the flow of resources. This re- To optimise the use of the flow of economic
quires an adjustment, but not radical change, in and environmental goods and services from a
the manner in which we approach the definition wetland, or any other freshwater habitat, a com-
of management boundaries for wetlands and prehensive management boundary needs to be
similar ecosystems. adopted that encompasses:
(a) the activities upstream which could influence
Rationalising planning and management the environmental processes that maintain the
boundaries functions, health and productivity of the wet-
land ecosystem, and
The rationalization of planning and management
(b) the activities downstream that depend upon
boundaries requires the integration of three sets
the flow of wetland resources.
of factors. These are: key environmental pro-
cesses, administrative units and activities which Figure 3d illustrates this more comprehensive
affect or are dependent on wetland resources. from of management boundary.
Figures 3a depicts a freshwater wetland and Having defined the basic management bound-
Figures 3b, c, and d illustrate different types of ary, we are still faced with the challenge of inte-
management boundary that can be adopted. grating the development objectives, policies, and
In Fig. 3b the wetland is shown with an up- management strategies of the private and public
stream management boundary defined by the interests within the administrative areas which
catchment area within which major land and overlap the boundary of the ecological system.
water use activities could have an impact on
environmental processes that govern the health
and productivity of the wetland ecosystem. Applying integrated planning and management
The catchment area concept is familiar to those (IPM) to wetlands and other freshwater habitats
involved in watershed management. However, for
political purposes river courses are often adopted The sectoral planning approaches that dominate
as convenient natural features to segregate the many economies have been shown to be effective
319
3a & 3b Ecological Management Boundary

a) b)
,
Water
Catchment -- /
/
/'.",.,.-- ......
, \
/ \
I \
I \
I
I
I
/
I
.... I
"'---- ......
3c Administrative Management Boundary
- •- Administrative
Boundary
Wetland
District' A'
3d Comprehensive Management Boundary
District 'B '
Water
Downstream area where the

environmental and
economic goods and
services generated by a
wetland may benefit
different interest groups
Fig 3a-d
320
for certain forms of activity. However, the lack of (e) improved food security through the reduction
integration of plans and investment strategies in storm damage to crops and continuity in
often leads to conflicts among different sectors, water supplies.
unnecessary competition for land and water re-
sources, poorly timed and uncoordinated provi-
sion of essential services, and negative economic Major elements in the IPM process for
and environmental externalities that decrease the freshwater habitats, including wetlands
potential returns from investment.
IPM can help to avoid many of these problems The basic starting point in the IPM process is the
and improve the performance of individual sec- objective appraisal of the resources generated by
tors of the economy by removing many of the wetlands and other freshwater habitats that can
constraints imposed by poorly integrated and co- be utilised to sustain development. A major fea-
ordinated development. IPM involves a number ture of the IPM process is that environmental,
of procedures that provide a comprehensive basis economic and socio-cultural information is used
for the identification of development opportuni- at different stages to determine the most effective
ties, selection of alternative forms and mixes of use of resources and to illustrate the relative mer-
development, planning these in an economically its of different development alternatives. The IPM
and environmentally sound manner, and manag- process is iterative, each stage helps to strengthen
ing their efficient implementation. Unlike tradi- successive stages, and plans and management
tional forms of project development that focus on measures are constantly monitored to assess their
maximizing the production of activities within in- effectiveness and to provide information with
dividual economic sectors, the basic objective of which to improve plans and management mea-
IPM is to identify the optimal mix of activities sures. The continuum of activities that provide
with which to fulfil development objectives given continuing support to the planning and manage-
the resources available to a society. ment process includes:
1. baseline studies to determine the key:
Benefits that can be derived from integrated
(a) biophysical processes that maintain the
planning and management
health and productivity, functions and
flow of economic and environmental
There is a wide range of benefits that can be
goods and services from the ecosystem;
gained from improvements in the management of
(b) socio-cultural and economic factors that
wetlands and other freshwater habitats and the
can contribute to, or constrain, the sus-
resources they generate, examples include:
tainable utilisation of these resources;
(a) reduction in the negative impact on people 2. evaluation of the different resources (eco-
and economic activities resulting from natu- nomic and environmental goods and services)
ral hazards whose incidence and severity of available to sustain social and economic de-
is exacerbated by the human induced disrup- velopment objectives;
tion of natural processes environmental con- 3. assessment of development opportunities
trol mechanisms; based upon the sustainable utilisation of the
(b) economic development sustained by the con- flow of resources;
tinuous supply of renewable resources; 4. appraisal of different forms of resource use to
(c) increased economic activity stimulated by the determine compatible and non-compatible
enhanced utilization of renewable resources; activities;
(d) increased social stability in rural communities 5. identification of the most appropriate mix of
resulting from the maintenance of access to activities and the formulation of guidelines
resources; for planning and managing development
321
activities to make efficient use of available the flow of renewable resources based upon pro-
resources; tection of the ecosystem, maintenance of its func-
6. assessment of the potential environmental, tions and the management of the uses of the re-
social and economic impacts associated with newable resources based upon a multiple-use
alternative forms of development; approach that optimizes the benefits to society
7. formulation of spatial plans for the location over time.
of selected activities;
8. formulation of management plans to imple-
ment the planned development; References
9. monitoring of development plans and man-
agement measures to determine their effec- Armentano, T. V. & E. S. Menges, 1986. Patterns of Change
tiveness and to identify unforeseen adverse in the carbon balance of organic soil-wetlands of the tem-
impacts on environmental processes and perate zone. J. Ecol. 74: 755-774.
Burbridge, P. R., 1984. The Planning and Management of
ecosystem functions so that they can be rec-
mangrove Resources in Asia. In Soepadmo (ed.), Proc.
tified before irreversible environmental, social Asian Symp. Mangrove Environment, Kuala Lumpur: 27-
or economic damage occurs; 42.
10. adaptation of plans and management mea- Burbridge, P. R., 1991. Zonation of Mangrove Ecosystems
sures to correct for unforeseen adverse im- for Integrated Multiple Use Management. In J. Kusler &
S. Daly (eds), Proc. Int. Symp. Wetlands and River Cor-
pacts and to improve the performance of the
ridors Management, Charleston: 99-101.
planned development; Gosselink, J. G. & R. E. Turner, 1978. The Role of Hydrol-
11. updating of baseline studies, resources as- ogy in Freshwater Wetlands. In R. E. Good, D. F.
sessments, appraisal of development options, Whigham & R. L. Simpson (eds), 1978. Freshwater Wet-
and refinement of planning and management lands: Ecological Processes and Management Potential,
Academic Press: 63-78.
guidelines based upon the information gath-
Hamilton, L. S. & S. C. Snedaker (eds), 1984. Handbook for
ered and lessons learned during the initial Mangrove Area Management, UNEP-East-West Center,
stages in the IPM process. IUCN, Honolulu, Hawaii, 123 pp.
Hollis, T. E. & E. Maltby, 1987, table prepared for work-
shop discussion at the Ramsar Convention, Regina
Saskatchewan, 1987.
Conclusions
Immirizi, C. P., E. Maltby & R. S. Clymo, 1992. The Global
Status of Peatlands and their Role in Carbon Cycling, a
Freshwater habitats, including wetlands generate report by the Wetlands Ecosystem Research Group for
far more complex and valuable renewable re- Friends of the Earth, London.
sources than is commonly recognised. Before James, R. F., 1991. The valuation of Wetlands: approaches,
methods and issues, PHPAjAWB Sumatra Wetland
projects are approved that could degrade the
Project Report no. 29, Asian Wetland Bureau, Bogor,
functional integrity of their ecosystems, very care- Indonesia, 94 pp.
ful assessments should be made of the true ben- McCormick, J., 1978. Ecology and the Regulation of Fresh-
efits and losses that will result. water Wetlands, In R. E. Good, D. F. Whigham & R. L.
Because of the great social and economic value Simpson (eds), 1978. Freshwater Wetlands: Ecological
of the environmental and economic goods and Processes and Management Potential, Academic Press:
41-356.
services derived from freshwater habitats such as Maltby, E., 1986. Waterlogged Wealth - why waste the
wetlands, we should not rely on Environmental world's wet places? Earthscan, London, 200 pp.
Impact Assessment and other forms of project Maltby, E., 1991. Wetlands - Their Status and Role in the
evaluation to guide their development in response Biosphere. In M. B. Jackson, D. D. Davies & H. Lambers
(eds), Plant life under oxygen deprivation, S and B. Aca-
to proposals for their exclusive use by the private
demic Publishing, The Hague: 3-21.
sector or by individual government agencies. A Mitsch, w. J. & J. G. Gosselink, 1986. Wetlands, Van Nor-
much more pro-active approach should be taken strand Reinhold, New York, 539 pp.
to actively plan for the sustainable utilization of Richardson, C. J., 1985. Mechanisms controlling phospho-
322
rous retention capacity in freshwater wetlands, Science 228: Wijkman, A. & L. Timberlake, 1984. Natural Disasters -
1424-7. acts of God or acts of Man?, Earthscan, London and
Sather, 1. M. & R. D. Smith, 1984. An overview of major Washington, 143 pp.
wetlands functions and values. U.S. Fish and Wildlife Ser-
vice Report FWS/OBS-84/18, Washington D.C., 68 pp.
Hydrobiologia 285: 323-325, 1994.
Recommendations & Resolutions
Recognising that population and development pressures are creating an increasing and serious threat
with considerable loss of habitat in the coastal, freshwater and wetlands of the ASEAN region, a group
of ecologists from the region and outside, attended a meeting on 4-6 November 1991 at the Institute
of Advanced Studies, University of Malaya. From the sessions of the seminar titled 'Ecology and
Conversation of Southeast Asian Marine and Freshwater Environments including Wetlands', the fol-
lowing expressions of concern and resolutions emerged.
I. REGIONAL COOPERATION AND RESEARCH Freshwater systems

PRIORITIES IN MARINE AND FRESHWATER
ECOLOGY OF SOUTHEAST ASIA Many of the peoples of the Southeast Asian re-
gion are dependent on rivers for their existence.
It has been demonstrated in the seminar that there
has been significant impact of development on
Environmental problems
river ecosystems, e.g. Siak river in Sumatra. De-
spite this little is known of the actual effects ex-
The environmental problems that challenge
cept for some economically important resources
human society are fundamentally ecological in
like fish. This is due to the fact that we know
nature. Among the research priorities identified in
extremely little of the ecology of these ecosystems.
relation to a sustainable biosphere are: (a) global
The only information available is on its water
climatic change, i.e. understanding how ecologi-
quality, where the limited number of parameters
cal processes affect local, regional and global cli-
measured tells us little of the health of this eco-
matic conditions, and how changes in these pat-
system.
terns of climate affect ecological processes. Many
There should be a concerted effort to obtain
interactions involving the atmosphere, soil and
basic biological information such as the range of
water should be considered. (b) maintaining bio-
organisms living in streams, and how they are
logical diversity and the relationship to total eco-
adapted to the dominant influences such as rain
systems, and (c) sustaining ecological systems, i.e.
spates and flooding, turbidity and trophic rela-
understanding when natural and managed eco-
tionships. This will complement chemical analy-
logical systems are stressed to the point they are
ses and provide the basis for assessing impacts
no longer capable of being sustained. Studies
due to development and polutants.
should be directed toward developing methods to
Without cooperation such a program of study
restore damaged systems.
will not be possible in the ASEAN region in the
face of limited resources and expertise. To opti-
Resolution - Planning for regional research co- mise the limited resources regional efforts are the
operation in the ASEAN countries should direct only solution.
attention, in an integrated manner, to the factors
that contribute to global climate change, the Resolution - The ASEAN region should begin
threat to biological diversity, and the need to sus- discussions to plan a cooperative study on fresh-
tain ecological systems. water ecosystems.
324
Microbial-Biogeochemical Problems in Coastal (G) National sanctuaries of sufficient size should

Southeast Asian Environments be established to preserve the major habitats.
These areas can also be used for scientific
Microbes (namely bacteria) are responsible for research to compare natural and degraded
the cycling of the elements (carbon, nitrogen, (un-preserved) systems.
phosphorus) essential for life in coastal waters (H) It is clear that with logging and the decline in
and sediments. At present, almost nothing is coastal and freshwater vegetation, erosion
known of these cycles or the organisms that run and runoff of soils into rivers and waterways
them. and their subsequent deposition in nearshore
coastal sediments needs to be investigated.
Resolution - It is recommended that graduate
What are the effects of increased sedimenta-
students, post-graduate fellows and/or other
tion and nutrient runoff on nearshore pelagic
scientists be educated abroad or in Southeast
and benthic food chains? What is the fate of
Asia on microbial techniques and the interpreta-
the material?
tion thereof. The methods involved are too diffi-
(I) Multidisciplinary studies need to be con-
cult to be understood simply by reading a manual.
ducted with scientists from developed coun-
tries to investigate problems such as point H
Programs of research to be pursued in relation to
above. Many such problems can only be un-
the resolution include:
derstood with the concerted effort of a physi-
(A) Estimate bacterial numbers and their activity cal oceanographer, sedimentary geologist
(respiration, production) in coastal and fresh and a microbiologist.
waters and sediments.
Resolution - An emphasis on process-oriented
(B) Study (temperature, measurements of or-
research should be encouraged further in South-
ganic carbon, nitrogen, dissolved nutrients
east Asia to understand the increase in the deg-
such as ammonium) that may relate to micro-
radation of aquatic habitats.
bial distribution, abundance and activity.
(C) Investigate the role of microbes in aquatic
food chains. This aspect can be studied by II. SUSTAINABLE UTILIZATION OF WETLAND
radioactive labelling of bacteria and micro- RESOURCES OF SOUTHEAST ASIA
algae and feeding them to important organ-
isms such as prawns and larval fishes. Ecology and Planning
(D) Carry out studies to relate a large number of
measurements usually taken (dissolved oxy- As scientists we need to take a more positive role.
gen, pH, temperature) by Southeast Asian So far as we see our role as defining and under-
scientists to the microbes which greatly affect standing the natural world. Scientists need to
them. overcome this 'show and tell' image and test our
(E) Determine the health of these environments insights in the real world where decisions are
through processes monitored at monthly made about resource use.
intervals. We spend much of our life learning and apply-
(F) Because of degradation of many Southeast ing scientific method, yet the problem of design-
Asian coastal and freshwater habitats and ing and making decisions for the use of resources
erosion, these microbial processes should be to solve human problems of development are
related (or attempt to be) to increased ero- based upon a different system of analysis, i.e. on
sion and subsequent changes in sediment a 'design method' involving creativity, trial and
characteristics such as the loss of clay and error and fine tuning as the many elements of the
silt, changes from reduced to oxidized sul- environmental problems are acknowledged and
phuric (as H 2 S04 ) conditions, etc. incorporated.
325
Rarely do scientists have the patience to get - the need to bring together the people and the
involved in these other processes where uncer- disciplines associated with each sectoral inter-
tainty is so prevalent and replication and verifi- est to allow not only multiple use planning but
cation is unlikely. What has emerged from the also for management.
present seminar is a belief that we do have a role.
However, unless we step out of our 'scientific Resolution - Recognising the importance of di-
culture' and learn to speak in the language of the verse and productive tropical environments and
planner and the bureaucrat, we effectively deny the need to introduce change to the way we
their access to our insights. It seems easier to approach planning and management of natural
make a scientist into a planner than to make a resources if sustainable development of these re-
planner into a scientist. Scientists need to become sources is to be achieved, it is resolved: that a
agents of positive change. concerted effort to be made by ASEAN scientists
The World Conservation Strategy is based on to playa positive role in planning and manage-
the explicit link between conservation and devel- ment processes affecting tropical environments.
opment. How do scientists respond to this? Much This role will involve emphasis on:
of our work in mangroves, and to a lesser extent
- greater integration of economic, social, cul-
other wetlands, in the past few decades has been
tural and ecological information,
concerned with identifying resources and func-
- positive management approaches to show the
tions within wetlands which, while not in the
social and economic benefits of utilizing the
mainstream of the national economy, neverthe-
flows of renewable resources,
less represent not only resources foregone but
- emphasis on communication and on conflict
also positive economic opportunities for local and
resolution among all interested parties.
regional economic and social development.
Optimal economic development has to be in-
tegrated with ecological understanding to enable
sustain ability to be achieved. But, rather than only Ecological Economics
seeking to limit levels of exploitation of existing
recognised resources to sustainable levels, we With the need to integrate ecology and econom-
should also be seeking - with all our knowledge ics it is essential that we develop new methods of
of alternative uses - to substitute other economi- interpretation so as to maximize all energy and
cally valuable resources into the economy (a re- environmental inputs and limit the losses thereof.
versal of the traditional economic development Scientists, economists, planners and policy mak-
model). ers accustomed to appraisals based on financial
What we must endeavour to do, and to be seen returns coupled with a consideration of environ-
by society to be doing, is to stimulate the diver- mental impacts (usually by simple substraction)
sification and expansion of the economy by in- need to become familiar with holistic analyses
volving ourselves in the development of better based largely on energy transformations with full
management procedures for existing recognised considerations of the resources involved.
resources and to also involve ourselves in devel-
oping new economically valuable resources from Resolution - Planning and the evaluation of
the natural systems which we wish to protect. present and past practices should take into ac-
If we can begin to do this I believe we can count the extent to which endeavours have or will
become a significant influence of change in two be optimal in terms of inherent energy transfor-
areas which have emerged time and again as im- mations, the environmental subsidies involved,
portant in this seminar. and the evaluation of the impact on renewable
- the need to manage the whole system and not resources and other environmental capital as
just one economic product. needed for the future.

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Developments in Hydrobiology 98

Reprinted from Hydrobiologia, vol. 285 (1994)

Springer Science+Business Media, B.V.

ISBN 978-94-010-4414-1 ISBN 978-94-011-0958-1 (eBook)

Printed on acid-free paper

All Rights Reserved

ECOLOGY OF MARINE AND FRESHWATERS INCLUDING WETLANDS

CONSERVATION AND POLLUTION

MANAGEMENT ISSUES, ECOLOGICAL ECONOMICS AND OTHER TOPICS

RECOMMENDATIONS AND RESOLUTIONS 323

Monitoring of sea surface temperature in the South China Sea

Introduction Malaysian EEZ, it is also responsible for the col-

Fig. 1a. Location of the Port Meteorological Offices in Malaysia.

to the Malaysian EEZ during the winter months. References

Neap-spring tidal effects on dissolved oxygen in two Malaysian estuaries

Bruce W. Nelson t, A. Sasekumar 2 & Zelina Z. Ibrahim 3

Key words: dissolved oxygen, Malaysia, estuaries, neap-spring tidal cycles

Introduction the dissolved oxygen distribution in two Malay-

Fzg. 1. Location Map and Land Use Patterns.

SUNGEI SELANGOR ESTUARY

Fig.2a. StatIOn Locations III the Selangor River Estuary.

DISTANCE FROM MOUTH, Kilometers

Station S·4.8 7 ·23 ·89 Stotion 5-12 7 ·24·89

... . . . .~~......-. -' o . - . .. .. -•. .. . . ..

DISSOLVED OXYGEN, m g./l. DISSOLVED OXYGEN, mo.!l.

S-II 8'11'89 A =1.1 m. S - 10 8 '11' 8 9 A = I. I m.

Fig. 50. Station Locations in the Klang River Estuary.

.,E5 "' . ------=:

Fig. 5b. Dissolved Oxygen Concentrations for a 1.4 m Neap Tide.

Dissolved 02. mg.!l.

Fig. 5c. Dissolved Oxygen Concentrations for a 4.2 m Spring Tide.

KLANG RIVER ESTUARY

The role of bacteria in nutrient recycling in tropical mangrove and other

Key words: bacteria, sediments, nutrients, mangroves, tropics

Introduction their bypro ducts (gases, etc.) leads to replenish-

Estuaries Bacteria Productivity Growth rate

• mangrove high Intertidal (station 1 ) • mangrove beach wrack (station 3)

A DNA SYNTHESIS (llgC.cm·3.h· l ) B. Il (h· l )

• o coral reef carbonates

• • •• • • seagrass muddy sands

subtidal y = 46.4 - 1.44x

Type of disturbance Microbial effect Ecosystem consequences

Clear-felling of vegetation Loss of biomass and leaching of Impoverishment of sediment, in-

References interface in a tropical mangrove system. Mar. Ecol. Prog.

Studies on the production of useful chemicals, especially fa'tty acids in

Wan-Loy Chu, Siew-Moi Phang* & Swee-Hock Goh

Introduction acids de novo. These fatty acids are accumulated

Chlorophyll-a (mg/l) Table 1. Chemical composition (% dry weight) of N. con-

Chemical Flask 1 Fermenter 2 Nitzschia"

of the fermenter culture was lower than the cul-

N. conspicua flask cultures ended by day 5. Mter

Chemical composition OL-----'----L---'----'------'----L---'----'-------'---·

Ash was the major chemical component of Days

Fatty acid Present study N. palea" N. alba b Reported values

N. conspicua * N. conspicua ** Nitzschia c N. angularis d Nitzschia d

Abbreviation: Tr. - Traces; -: not present.

study, maximal fatty acid content was only 3.6% o 2 4 6 8 10 12 14 16 18 20

g)20:4 (AA) h)20:5 (EPA)

44.0% total fatty acids) in relation to culture age References