universiti putra malaysia a study on diversity and …
TRANSCRIPT
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.