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UNIVERSITI PUTRA MALAYSIA A STUDY ON DIVERSITY AND DENSITY OF MARINE DINOFLAGELLATES IN THE STRAITS OF MALACCA AZURA ALIM FSAS 2001 11

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UNIVERSITI PUTRA MALAYSIA

A STUDY ON DIVERSITY AND DENSITY OF MARINE DINOFLAGELLATES IN THE STRAITS OF MALACCA

AZURA ALIM

FSAS 2001 11

A STUDY ON DIVERSITY AND DENSITY OF MARINE DINOFLAGELLATES IN THE STRAITS OF MALACCA

By

AZURA ALIM

Thesis Submitted in Fulfilment of the Requirements for the Degree of Master Science in the Faculty of Science and Environmental Studies

Universiti Putra Malaysia

March 2001

Abstract of thesis presented to the Senate ofUniversiti Putra Malaysia in fulfilment of the requirement for the degree of Master of Science

A STUDY ON DIVERSITY AND DENSITY OF MARINE DINOFLAGELLATES IN THE STRAITS OF MALACCA

By

AZURA ALIM

March 2001

Chairman Drs. Idris Abd. Ghani

Faculty Science and Environmental Studies

The objective of this study is to update and compare the species composition,

distribution and abundance of dinoflagellate in the Straits of Malacca . The

samples were collected during the JICA- MASDEC Straits of Malacca Expedition

on board RIV K. K. Mersuji from November 23 to December 2, 1 998 (northeast

monsoon season) and March 25 to April 4, 1 999 (pre southwest monsoon season)

by using "Van Dorm" water sampler. A total of 78 dinoflagellate species from 2 1

genus, 1 3 families, and six orders under Division Pyrrophyta were observed and

identified. During the northeast monsoon, Peridiniales was the dominant order

accounting for 87.87% of the total dinoflagellate density, followed by

Dinophysiales (5 .49%), Prorocentrales (2.98%), Noctilucales ( 1 . 85%),

Pyrocystales ( 1 .45%) and Gymnodiniales (0.36%). Peridiniales also constitute

of an important group during the pre southwest monsoon with an average 7 1 .84%

of the total dinoflagellate density, followed by Gymnodiniales (20 .5 1 %),

Dinophysiales (4.80%), Pyrocystales ( 1 .40%), Prorocentrales (0.89%) and

Noctilucales (0.57%). In terms of numbers, the total density of dinoflagellates

ii

varied from 2.64 cells/l at Station 24 to 2.93 x 1 03 cells/l at Station 1 3 and 1 8 .02

cells!l at Station 19 to 6 .24 x 102 cells!1 at Station 1 4 during the northeast

monsoon season and the pre southwest monsoon season respectively. The middle

zone of the Malacca Straits showed the highest density of dinoflagellate compared

to the northern and southern zone for both monsoon seasons. During the northeast

monsoon, the density of dinoflagellate in the offshore stations was higher than in

the inshore stations. In contrast, the inshore stations of the Straits showed the

highest density of dinoflagellate compared to the offshore stations during the pre

southwest monsoon. Ceratium furca was the most dominant species in the Straits

of Malacca during the northeast monsoon (maximum cell density reached 2 . 8 7 x

1 03 cells/l at Station 1 3) and dominated dinoflagellate assemblages almost at all

stations. Gymnodinium simplex, Protoperidinium ovum and Ceratium furca were

the three most dominant species in the Straits ofMalacca during the pre southwest

monsoon. ANOVA showed a significant difference (F = 2 1 .05, p < 0.05 and F =

1 1 .66, p < 0.05) in species diversity and species evenness between the northeast

monsoon and the pre southwest monsoon. However, there was no significant

difference (F = 2 .34, p > 0.05) in species richness between the northeast

monsoon and the pre southwest monsoon.

III

Abstrak tesis yang dikemukakan kepada Senat Universiti Putra Malaysia sebagai keperluan untuk ijazah Master Sains

KAJIAN KE ATAS DlVERSITI DAN DENSITI DINOFLAGELAT MARIN DI SELAT MELAKA

Oleh

AZURAALIM

Mac 2001

Pengerusi Drs. Idris Abd. Ghani

Fakulti Sains dan Pengajian Alam Sekitar

Objektif kajian adalah untuk mengemas kini dan membuat perbandingan

komposisi spesis, taburan serta kepadatan dinoflagelat di Selat Melaka. Sampel

fitoplankton dikumpulkan semasa Ekspedisi Selat Melaka JICA-MASDEC

menggunakan kapal RIV K. K. Mersuji dari 23 hb. November hingga 2 hb.

Disember, 1998 (monsun timur laut) dan dari 25 hb. Mac hingga 4 hb. April,

1999 (monsun pra barat daya) dengan menggunakan a1at pensampelan air "Van

Dorm". Sebanyak 78 spesis dinoflagelat daripada 21 genus, 13 famili dan en am

order daripada Divisi Pyrrophyta telah dikenalpasti. Semasa monsun timur laut,

Peridiniales merupakan order dominan dengan 87.87% daripada jumlah densiti

keseluruhan dinofiagelat, diikuti oleh Dinophysiales (5.49%), Prorocentrales

(2.98%), Noctilucales (1.85%), Pyrocystales (1.45%) dan Gymnodiniales

(0.36%). Peridiniales juga merupakan kumpulan penting semasa monsun pra barat

daya dengan purata 71.84% daripada jumlah keseluruhan densiti dinoflagelat,

diikuti oleh Gymnodiniales (20.51%), Dinophysiales (4.80%), Pyrocystales

(1 .40%), Prorocentrales (0.89%) dan Noctilucales (0.57%). Dari segi bilangan,

lV

jumlah keseluruhan densiti dinoflagelat dari 2.64 sel/l pada stesen 24 hingga 2.93

x 1 03 selll pada stesen 1 3 dan 1 8.02 selll hingga 6.24 x 102 sel/l pada stesen 14

semasa monsun timur laut dan monsun barat daya. Zon tengah Selat Melaka

menunjukkan densiti dinoflagelat yang lebih tinggi j ika dibandingkan dengan zon

utara dan zon selatan pada kedua-dua monsun. Semasa monsun timur laut, densiti

dinoflagelat lebih tinggi di stesen yang terletak j auh dari pantai berbanding stesen

yang terletak berhampiran dengan pantai . Berbeza dengan monsun pra barat daya

di mana densiti dinoflagelat lebih tinggi di stesen yang terletak berhampiran

pantai berbanding dengan stesen jauh dari pantai. Ceratium furca adalah spesis

dominan di Selat Melaka semasa monsun timur laut (densiti maksimum yang

dicapai sebanyak 2.87 x 1 03 selll pada stesen 1 3) dan mendominasi kumpulan

dinoflagelat hampir semua stesen. Gymnodinium simplex, Protoperidinium ovum

dan Ceratium furca adalah tiga jenis spes is dominan di Selat Melaka semasa

monsun pra barat daya. ANOVA menunjukkan perbezaan bererti (F = 2 1 .05 , P <

0.05 dan F = 1 1 .66, p < 0 .05) bagi diversiti spesis dan kesamarataan spesis antara

monsun timur laut dan monsun pra barat daya. Walau bagaimanapun, tiada

perbezaan bererti (F = 2.34, p > 0.05) bagi kekayaan spesis antara monsun timur

laut dan monsun barat daya.

v

ACKNOWLEDGEMENTS

In the name of ALLAH the most merciful and honorable.

I would like to express my appreciation to all those who help to contribute to this

research. I thank ALLAH for everything. I sincerely wish to express my

gratitude to my advisor Drs. Idris Abdul Ghani for his full support, guidance,

understanding and encouragement throughout the cause of this research.

I would like to express my sincere thanks to Dr. Hishamuddin Omar and Dr. Misri

Kusnan, for serving as my advisory committee, and providing their expertise and

other words of wisdom. Also, I am greatly indebted to them for their critical

review of the manuscript of my thesis.

A special thanks must go out to Dr. Hishamuddin for his expert guidance and

spent countless hours trying to help me understand the concept of ecology. I thank

everyone who has been part of the Aquatic Laboratory.

Finally, I dedicate this work to my lovely son Muhammad AI-Hadi, my beloved

husband Mohd. Harees Masian who have encouraged me to continue my study

and provided financial support so that I may pursue my study without any

difficulties and a special thank must go out to my parents for believing in me and

always being there.

VI

I certify that an Examination Committee met on 3 1 st March 2001 to conduct the final examination of Azura Binti Alim on her Master of S cience thesis entitled "A Study on Diversity and Density of Marine Dinoflagellates in the Straits of Malacca" in accordance with Universiti Pertanian Malaysia (Highe.r Degree) Act 1 980 and Universiti Pertanian Malaysia (Higher Degree) Regulation 1 98 1 . The Committee recommends that the candidate be awarded the relevant degree. Members of the Examination Committee are as follows:

AHMAD ISMAIL, Ph.D. Department of Biology, F acuIty of Science and Environmental Studies, Universiti Putra Malaysia, (Chairman)

IDRIS ABDUL GHANI, M. Sc. Department of Biology, Faculty of Science and Environmental Studies, Universiti Putra Malaysia, (Member)

HISHAMUDDIN OMAR, Ph.D. Department of Biology, Faculty of Science and Environmental Studies, Universiti Putra Malaysia, (Member)

MISRI KUSNAN, Ph.D. Department of Biology, Faculty of Science and Environmental Studies, Universiti Putra Malaysia, (Member)

MOHD. GHAZALI MOHAYIDIN, Ph.D. Professor Deputy Dean of Graduate School Universiti Putra Malaysia

Date: 1 1 SEP 2001

vii

This thesis submitted to the Senate of Universiti Putra Malaysia has been accepted as fulfilment of the requirement for the degree of Master o f Science.

AINI IDERIS, Ph.D. Professor Dean of Graduate School Universiti Putra Malaysia

Date: 0 8 NOV 2001

VIIl

DECLARATION

I hereby declare that the thesis is based on my original work except for quotations and citations, which have been duly acknowledged. I declare that this thesis has not been previously or concurrently submitted for any other degree at UP M or other institutions.

Date: 29th August 2001

IX

TABLE OF CONTENTS

Page

ABSTRACT . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ............ ............ .......... ........ 11

ABSTRAK . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . v ACKNOWLEDGEMENTS. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vlll APPROV AL SlIEETS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . " ............ ....... IX DECLARATION . . . . . . . . . . . . . . .. . . ,. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xl LIST OF TABLES . . . . . . . ...... .......... ...... .............. .... ............ .. .... ... XIV LIST OF FIGURES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . xv GLOSARY OF TERMS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . XVlll

CHAPTER

I INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . Research Objectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

II LITERATURE REVIEW . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . .. Classification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ' " . . . . . . . . . . . . . . . . . . . . . Morphological Characteristic . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Taxonomy of Dinoflagellates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Harmful Algae Report in South East Asian Waters . . . . . . . . . . . . . . . . . . . . . . . The Status of the Dinoflagellates Studies in Asia . . . . . . . . . . . . . . . . . . . . . . . . . . Dinoflagellates Study in Malaysia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Economic Impact of Dinoflagellatess . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F actors Affecting Growth of Dinoflagellates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Light . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . , . . . . . . . . Temperature . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Salinity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nutrients . . . . . . . . . . . . . . . . . . . ' " . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . .

Wind Flow in the Straits of Malacca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . , . . . . . . .

III MATERIALS AND METHODS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Study Area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sampling Techniques . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ' " . . . . . . . . . . . . . . . . . . . . . . . . FixationlPreservation and C oncentration of Algal Samples . . . . . . . . . . . . . . Quantitative Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . " . . . Identification of Dinoflagellate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . SEM Preparation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Statistical Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . ' " . . . . . . . . . . . . . . . . .. . . . . . . . . . . . .

IV RESULTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S

. C

't' pecles ompoSI Ion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Order Peridiniales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Order Dinophysiales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . .. . . " . . . . . . . . . . .

Order Gymnodiniales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Order Prorocentrales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

x

1 3

4 4 5 6 9 1 1 1 2 1 5 1 6 1 7 1 8 20 2 1 22

23 23 23 25 26 28 28 29

30 30 34 36 36 37

Order Pyrocystales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Order Noctilucales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . .

Northeast Monsoon Season (First Cruise) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pattern of Species Abundance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . S ' D'

. peCles lverslty . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Spatial Distribution of Dinoflagellates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Species Diversity, Evenness and Richness . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cluster of Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Percentage Cover of Dinoflagellate Orders . . . . . . . . . . " . . . . . . . . . . . . . . . . Percentage C over of Order Peridiniales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Physical and Chemical Parameters . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . .

Pre Southwest Monsoon (Second Cruise) . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . Pattern of Species Abundance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S ' D'

. peCles lverslty . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . .

Spatial Distribution o f Dinoflagellates . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . Species Diversity, Evenness and Richness . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cluster of Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Percentage Cover of Dinoflagellate Orders . . . . . . . . . . . . . . . . . . . . . . . . . . . . Percentage Cover of Order Peridiniales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Physical and Chemical Parameters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Comparison between Northeast and Pre Southwest Monsoon . . . . . . . . . . . Pattern of Species Abundance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Species Number . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spatial Distribution of Dinoflagellates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Species Diversity, Evenness and Richness . . . . . . . . . . . . . . . . . . . . . . . . . . . . Physical and Chemical Parameters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

V DISCUS SION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S

. C 't' pecles omposl lon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . � . . . . . . . .. .

Pattern of Species Abundance . . . . , . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spatial Distribution of Dinoflagellates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Species Diversity, Evenness and Richness . . . . . . . . . . . . . . . . . . . . . . . . . . . . Physical and Chemical Parameters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

VI SUMMARy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

37 37 38 38 40 42 44 46 49 5 1 52 56 5 6 59 5 9 6 1 62 66 69 70 70 70 73 73 74 76

78 78 84 86 88 89

9 1

BIBLIOGRAPHY . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94 APPENDIX . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 1 8 VITA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . .. . . . . . . . . . . . . . . . . . . . . . . . . . 1 73

xi

LIST OF TABLES

Tables Page

Location of the sampling stations . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . 24

2 Arbitrarily division of sampling stations in the Straits of Malacca. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... . . . . . . . . . . . . 25

3 Methods of quantification of algae. . . . . . . . . . . . . . . .. .. . .. . .. .. . . .. . . . 27

4 Species list of dinoflagellates in the Straits of Malacca during the northeast monsoon season (first cruise) and the pre southwest monsoon season ( second cruise) . . . . . . . . . . . . . . . . . . . . . . .. 3 1

5 List of the most frequent dinoflagellate encountered during the northeast monsoon season. . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . 4 1

6 List of the rare species of dinoflagellates encountered during the northeast monsoon season . . . . . . . . . . . . . . . . . . . . . . . . , . . . . . . . . . . . . . . . .

7 List of the most common dinoflagellate encountered during the pre southwest monsoon season . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

8 List of frequent species of dinoflagellates encountered during the pre southwest monsoon season . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . .

9 Correlation coefficients comparing total cell numbers (density) with physicochemical parameters . . . . . .. .. . . . . . . . . . . . . . . .

xii

42

57

58

77

Figure

LIST OF FIGURES

(A, B) Basic anatomy of gymnodinoid dinoflagellate, redrawn from Dodge (1982) . (C, D) Basic anatomy of peridinoid

Page

dinoflagellate. Modified after Evitt (1985) . . . . . . . . . .. .. . . .... . ... . .. 7

2 (A, B) Schematic of Gonyaulax showing ventral and dorsal surfaces (modified from Eaton, 1980). (C, D, E) Schematic of Gonyaulax with sulcal plates (redrawn from Balech, 1980) . . . . . .. . . . . . . .... .. . . .. . . .. . . . . . . . . . . . . . . . . . .. . . . . ... .. . . . . . . . . .. . . . . . . 10

3 Map showing the location of the sampling stations in the Straits of Malacca. . . . . . . . . . ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

4 Total density of dinoflagellates in the Straits of Malacca during the northeast monsoon season (first cruise). . .... . . ... . .... . . . . . . .. . 3 9

5 Dominant species of dinoflagellates in the Straits of Malacca during the northeast monsoon season (first cruise). .... .. . . .. . . . .. 3 9

6 Species number of dinoflagellates in the Straits of Malacca during the northeast monsoon season (first cruise). . . . .. . . . . . . . . .. 43

7 Changes in diversity index (H') of dinoflagellates in the Straits of Malacca during the northeast monsoon season (first cruise) and the pre southwest monsoon season (second cruise) . . . . . . . . . . . 43

8 Changes in species evenness (J') of dinoflagellates in the Straits of Malacca during the northeast monsoon season (first cruise) and the pre southwest monsoon season (second cruise).. 45

9 Changes in species richness (R) of dinoflagellates in the Straits of Malacca during the northeast monsoon season (first cruise) and the pre southwest monsoon season (second cruise)... . . .. . ... 45

10 Dendogram for the cluster analysis of 18 sampling units (first cruise) using Relative Euclidean Distance and flexible strategy. 47

11 Percentage cover of dinoflagellate orders in the Straits of Malaccaa during the northeast monsoon season (first cruise). . .. 50

12 Percentage cover of Order Peridiniales in the Straits of Malacca during the northeast monsoon season (first cruise). ... .. .. . . .. . . .. 50

xiii

13 Temperature variation (OC) in the Straits of Malacca during the northeast monsoon season (first cruise) and the pre southwest monsoon season (second cruise) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... . .. . .. 53

14 Salinity variation (ppt) in the Straits of Malacca during the northeast monsoon season (first cruise) and the pre southwest monsoon season (second cruise) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 3

15 Conductivity variation (mS/cm) in the Straits of Malacca during the northeast n,onsoon season (first cruise) and the pre southwest monsoon season (second cruise) . . . . . . . . .. . . . . . . . . . . . . . . . 54

16 Dissolved oxygen variation (mg/l) in the Straits of Malacca during the northeast monsoon season (first cruise) and the pre southwest monsoon season (second cruise) . . . . . . . . . . . . . . . . . . . . . . . .. 54

17 pH variation in the Straits of Malacca during the northeast monsoon season (first cruise) and the pre southwest monsoon season (second cruise) . . . . . . . . .. . .. . . . . . . .. .. . . . . . . . . . .. . . . . .. . . . . .. . .. 55

18 Total density of dinoflagellates in the Straits of Malacca during the pre southwest monsoon season (second cruise). . . . . . . . . . . .. . . . 5 5

19 Dominant species of dinoflagellates in the Straits of Malacca during the pre southwest monsoon season (second cruise). . . . . . . 60

20 Species number of dinoflagellates in the Straits of Malacca during the pre southwest monsoon season (second cruise) . . . . . . . 60

21 Dendogram for the cluster analysis of 20 sampling units (second cruise) using Relative Euclidean Distance and flexible strategy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63

22 Percentage cover of dinoflagellate orders in the Straits of Malacca during the pre southwest monsoon season (second cruise) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67

23 Percentage cover of Order Peridiniales in the Straits of Malacca during the pre southwest monsoon season (second cruise) . . . . . . . 67

24 Total density of dinoflagellates in the Straits of Malacca during the northeast monso:); 1 season (first cruise) and the pre southwest monsoon season (second cruise) . . . . . . . . . . . . . . . . . . . . . . . .. 72

25 Species number of dinoflagellates in the Straits of Malacca during the northeast monsoon season (first cruise) and the pre southwest monsoon season (second cruise) . . . . . . .. .. . . . . . . . . . . . . . . . 72

xiv

26 Variation of diversity index versus total density of dinoflagellates in the Straits of Malacca during the northeast monsoon season (first cruise) ... . .. . . . . .. . . . . ....... . .. .... .. .... .. . .. 75

27 Variation of diversity index versus total density of dinoflagellates in the Strait s of Mala cca during the pre southwest monsoon season (second cruise) . . . . . . .. . . . . .. . .. . .. . .. . . 75

xv

Antapex

Apex

Apical pore

Areola

Armored

Bioluminescent

Chain

Chlorophyll a

Cyst

Diarrhetic shellfish pOlsonmg

Dinoflagellates

Epitheca

Flagellum

GLOSARY

bottom end of the dinoflagellate cell.

top end of the dinoflagellate cell .

distinctive pore-like structure, present on top of some dinoflagellates cell.

the regularly repeated circular depressions m the cellulose wall of dinoflagellates.

dinoflagellate cells with a thick cellulose wall (theca) composed of rigid plates.

the emission of visible light by living organisms, such as the fireflies on the land and the dinoflagellates in the sea.

a group of micro-organisms which are connected in a linear arrangement.

the green colouring substance found m all photosynthetic algae and higher plants.

a thick-walled stage in the life-history of a dinoflagellate, also called resting spore. The cyst wall is often made up of resistant sporopollenin and may be ornamented with spines. Cysts are often produced at the onset of adverse conditions, and when conditions are suitable, they may germinated to seed new plankton blooms.

human illness caused by eating shellfish contaminated with certain dinoflagellate toxins (e.g. from Dinophysis acuminata).

single-celled golden-brown algae, characteristically having a large nucleus with clearly visible chromosomes, and with two flagella, protruding from the girdle groove and the sulcus groove, respectively.

the upper half of the dinoflagellate cell.

Whip-like extensions of the cells of certain mlcro­organisms. Mainly used for locomotion.

xvi

Girdle groove

Hom

Hypotheca

List

Micron

Naked

Neurotoxins

Paralytic shellfish pOlsonmg

Plankton bloom

Red tide

Scanning electron microscope

Spine

Sulcus

Symbiosis

Theca

the horizontal groove which encircles the dinoflagellate cell and which contains the transverse flagellum.

extension of the cellulose plates of a dinoflagellate cell .

the lower half of the dinoflagellate cell.

extension of the cellulose plates of dinoflagellate cells, usually around the girdle groove.

one thousandth of millimeter (/-lm).

Plankton cells bounded by a membranous covering only. These organisms are very easily damaged beyond recognition.

toxins that disrupt the normal functioning of the human nervous system.

human illness caused by eating shellfish which are contaminated with certain dinoflagellate toxins (e.g. Gymnodinium catenatum or Pyrodinium bahamense). In extreme cases this can lead to death through respiratory paralysis.

Prolific growth of plankton algae.

dens concentrations of plankton organisms (mostly dinoflagellates) that can colour the sea red or brown.

a type of electron microscope in which a beam of electrons scans the outer surface of objects previously coated with a thin layer of gold or platinum.

solid projection on the surface of the dinoflagellate cell wall.

the vertical groove on the dinoflagellate cell, extending onto epitheca and hypotheca, which guides the longitudinal flagellum.

the living together of two species of organisms with mutual advantages for both.

the complete cell covering of the dinoflagellate cell.

xvii

CHAPTER I

INTRODUCTION

Dinoflagellate is the second most abundant phytoplankton group followed by

diatoms. The name 'dinos' comes from the Greek word meaning 'whirling' as this

type of movement is characteristic of these organisms. Dinoflagellates is a unique

organisms because it posses two flagella or whiplike appendages that help them to

move up or down in the water column. They are a diverse group, comprising

approximately 1 1 5-1 3 1 genera (Sournia, 1986; Sournia et ai., 1 991 ), 2,000 living

species and 2,000 fossil species have been described (Fukuyo & Taylor, 1 989).

Dinoflagellates are mostly unicellular and range from 2 to 2,000 �m in diameter

(Hallegraef, 1988).

The dinoflagellates are very abundant throughout the world. These organisms are

important members of the phytoplankton in both fresh and marine waters,

although a much greater variety of this form is found in marine relatives. They are

found in all latitudes from the Artic and Antartic sea to the tropics. In warm

water, dinoflagellates are usually large and occurred in various forms. Lee

( 1 989) reported that dinoflagellate are more important in the warmer waters than

in colder polar waters. Many dinoflagellates also occur in marginal habitats.

According to Smetacek et al. ( 199 1 ) some are confined to near-shore or neritic

waters due to the richness of nutrient and lower salinity than the open sea.

2

Dinoflagellates are important food source for 'marine microzooplankton,

including fish larvae of which the p roduction is essential to marine fishery

(Zhen g et af., 1 989) . Their blooms are good source of carbon to the marine food

web (White, 1984). The organisms making up this group are very diverse in

nature, ranging from coral reef symbionts to free-living oceanic phytoplankton.

The best known associations are the "zooxanthellae" (Symbiodinium spp.) found

in corals, clams, radiolarians, acantharians and foraminiferans. In addition, there

are some parasitic and saprophytic species (Zheng et al., 1 989).

Dinoflagellate are probably best known as a principal cause of red tides and

paralytic shellfish poisoning (PSP). "Red tide" is the name given to the

phenomenon of red discoloration of marine and estuarine waters caused by a

bloom of phytoplankton, particularly dinoflagellates. The water becomes

noticeably coloured when concentrations reach to about 200,000-500,000 celVI,

and as the bloom develops, concentrations may exceed 108 cellll (Lalli and

Parsons, 1993). The colour of water can range from blood red to orange or brown

in daylight and is often visible as phosphorescence at night (Carey, 1 983). These

are features of coastal waters and more frequent in the tropics and subtropics than

temperate seas. Some species have attracted attention because of their beautiful

bioluminescent properties or their role in producing either toxic or non-toxic

blooms (Halle graeff, 1993).

3

The present study deals with dinoflagellate in the Straits o f Malacca. The Straits

of Malacca is one of the longest straits in the world and the busiest straits in the

ASEAN country. This straits is important for shipping and transportation, natural

resources, tourism, climatic and life support system o f the littoral states and the

world. Some 600 ships use it daily with more than 90% o f these vessels destined

for Japan, Taiwan, Korea, Singapore and Hong Kong (Ibrahim and Noro, 1 999).

The increase of economic activities in the region also resulted in an upsurge of

traffic in the Straits . The intensifie d world wide traffic may be responsi ble for the

spread of harmful phytoplankton species between re gions and continents

(Halle graef, 1 993) . Hallegraeff and Bolch ( 1992) confirmed the presence of

Gymnodinum catenatum cysts in four ballast water samples entering Australian

ports from both Korea and Japan. The organism is now also occurred in Sebatu,

Melaka (An n and Normawaty, 1 996; Ann et al. , 2000) .

Research Objectives

The primary purpose of the present study is to update the species compos it ion,

distribution and abundance of dinoflagellate in the Straits of Malacca. The second

objective is to compare the species composition, distribution and abundance of

dinofla gellates in the Straits durin g the northeast monsoon season and the pre

southwest monsoon season. JICA-MASDEC Straits o f Malacca Expedition

provides opportunity to do comprehensive study on dinofla gellate in t he Straits.

CHAPTER II

LITERATURE REVIEW

Classification

The dinoflagellates are members of Kingdom Protista under the phylum or

division Pyrrophyta or Dinophyta (Dinophycophyta) in botanical systems and

the order Dinoflagellida or Dinoflagellata in zoological systems. Until recently,

the group was generally divided into two classes or subdivisions, which were

essentially based on the position of the flagellar insertion. Some of the names for

the classes used by various authors;

Adiniferidea Diniferidea

Adina Diniferina

Desmokontae Dinophyceae

Desmokontae Dinokontae

Desmophyceae Dinophyceae

Lebour, 1 925

Levine et al. , 1 980

Schiller, 1 933, 1 937

Fritsch, 1 935

Leoblich, 1 970

In 1 973 Dodge and Bibby showed that there was no basic ultrastructural

distinction between the two groups and proposed that they should be

amalgamated, therefore Parke and Dixon ( 1 976) and Dodge ( 1 982), used only

one class, Dinophyceae. Following this scheme Leoblich ( 1 976) introduced a

new class, Syndiniophyceae, for the genera of intracellular parasitic

5

dinoflagellates. Based on Chretionnot-Dinet et al. ( 1 993) dinoflagellates were

classified into one class only, namely Dinophyceae, with 1 4 orders and 42

families.

Morphological Characteristics

Dinoflagellates are single-celled organlsms that have t wo distinctive flagella

during the least part of their life c ycle and a special type of nucleus known as

dinokaryon. The cells is divided by a transverse girdle groove into two parts, an

upper (epitheca) and a lower (hypotheca) region. Within the groove lies the

flattened, ribbon-like, transverse flagellum which encircles the cell. This may be

situated centrally, apically or posteriorly. The smooth longitudinal flagellum

extends to the base of the cell within a longitudinal furrow (sulcus). The

transverse flagellum causes the rotational motion of the cell, while the

longitudinal flagellum propels the cell forward.

Dinoflagellates, like diatoms, can exhi bit extensive morphological variations with

spines , horns and wings, especially in tropical species. These morphological

extensions are thought to aid flotation, protect against grazing by small

zooplankton, and enhance nutrient uptake through increased surface area and

rotational movements.

Dinoflagellates are described as armored or unarmored, according to the type of

cel l covering (theca or amphiesma), which always consists of several laye rs of

membranes. The thecate or armored group has a number of ce llulosic plates

6

which are thecal plates , or simply theca as a whole. Many thecal plates tightly

assembled to form a firm, strong peripheral skeleton. The number, shape and

arrangement of these plates form a distinctive geometry or topology known as

'plate tabulation', which is the main means for classification and they are a most

important taxonomic criterion (Taylor, 1980).

The number, arrangement and thickness of the thecal plates can vary

considerably. They may also be o rnamented with pores, depressions, spines,

ridges and reticulations (Vesk et ai, 1990). Scanning e lectron microscopy has

reveale d that ornamentation of the thecal plates and fine structure of the apical

pore, present on the top of many cells are also useful diagnostic characters

(Hallegraeff, 1 988). The other, athecate, unarmored, or nake d group lacks of the

plates.

Taxonomy of Dinoflagellates

Athecate species are identified principally by the size and shape o f their cells, i.e.

cell outline, position of girdle and sulcus groove, girdle displacement. In the

thecate group, genera are classified according to plate pattern, which is decided b y

the arrangement o f thecal plates. Species in each genus are identified by the s ize

and shape of the cell, similarly taxonomy of athecate forms. Therefore, for

critical taxonomy of thecate forms, staining or dissociation of thecal plates to

recognize their tabulation is necessary in addition to observation of cell shape

(Fukuyo and Taylor, 1 989).

epitheca

girdle

hypotheca

apex or anterior

chloroplast

flagella

nucleus

pusule

sulcus

antapex or posterior A B

ventral view

apex or anterior

�apical

..........- horn

7

clorsal view

epitlleca

PlateS } number and

utures arrangement = tabulation

hypotheca flagella

antapex or posterior antapical horns

c o ventral view dorsal view

Fi�ure 1: (A, B) Basic anatomy of gymnodinoid dinoflagellate, redrawn from Dodge (1982). (C, D) Basic anatomy of peridinoid dinoflagellate. Modified after Evitt, 1985.