bio-dynamics and life cycle of shrimp ...psasir.upm.edu.my/544/1/1600396.pdfiv the present study was...

BIO-DYNAMICS AND LIFE CYCLE OF SHRIMP ( PENAEUS

SEMISULCATUS DE HAAN), IN BUSHEHR COASTAL

WATERS OF THE PERSIAN GULF

By

NASSIR NIAMAIMANDI

Thesis Submitted to the School of Graduate Studies, Universiti Putra Malaysia, in

Fulfilment of the Requirements for the Degree of Doctor of Philosophy

August 2006

ii

DEDICATION

This thesis is dedicated to

My wife,

Noushin Farhadi

Your motivation, sacrifice and support during the period of my academic

mission are appreciated.

iii

Abstract of thesis presented to the Senate of Universiti Putra Malaysia in

fulfilment of the requirement for the degree of Doctor of Philosophy

BIO-DYNAMICS AND LIFE CYCLE OF SHRIMP (PENAEUS

SEMISULCATUS DE HAAN), IN BUSHEHR COASTAL

WATERS OF THE PERSIAN GULF

By

NASSIR NIAMAIMANDI

August 2006

Chairman: Associate Professor Aziz bin Arshad, PhD

Faculty: Science

Shrimp fisheries in the Persian Gulf have a long history. The trawl fishery for

penaeid prawn developed along the mid-northern coasts of the Persian Gulf

(Bushehr waters) beginning in 1960. From 1998 to 2004 shrimp landings have

fluctuated between 480t and 2700t per year. The major species of penaeid prawn,

in terms of catch (>80%) and economic importance, in the Persian Gulf (including

Bushehr waters) is Penaeus semisulcatus, commonly known as the green tiger

prawn. More than 2000 vessels and 33000 fishermen from the Bushehr study area

have participated in the shrimp fishery each season (August-September) over the

last decade. In the Arabian side of the Persian Gulf, P. semisulcatus has been now

become the most important single component species. Despite of commercial

importance, there has been very little published research in the biology this species.

In the study area, Bushehr waters, shrimp fishery is only controlled during open

season. Because of lack of any data, there is no any prohibition and restricted areas

in the nursery and spawning grounds and also the fluctuations of shrimp resources

are unknown.

iv

The present study was carried out in Bushehr coastal waters during 2003-2005 and

describes the bio-dynamics and life history of P. semisulcatus. Monthly sampling of

shrimp was collected by R/V Lavar II for 15 months from January 2003 to March

2004 from 50 fixed stations. Carapace length (CL) frequency was used to estimate

growth, mortality and exploitation rates for males and females using FiSAT

statistical analyses software. The Von Bertalanffy growth parameters were

estimated as K = 1.6 Year-1

and 2.2 Year-1

and L∞ = 38 mm and 50.4 mm for males

and females, respectively. Estimated annual mortality rates for males were Z = 6.4,

F= 4.3, and M = 2.11. While females were Z = 8.2, F = 5.8, and M = 2.41.

Exploitation rates for males and females were E = 0.67 and E = 0.70, respectively.

Relatively high values of total mortality and exploitation rates were obtained,

suggesting this species is overexploited. The maximum ages calculated were 20

months for males and 15 months for females.

v

Gonad maturity results indicated that the major spawning activity for P.

semisulcatus within the study area occurs during winter and early spring, with an

additional minor peak evident in autumn. These periods coincided with seasonal

rainfall in the Bushehr study area. Size at 50% maturity (L50%) was 40.6 mm

carapace length (CL). Less than 5% of females were smaller than 24 mm CL.

Mature shrimps aggregated in the middle and southern regions of study area at

depths of 20-30 m. The areas of Imam Hassan, Genaveh and Helleh represented the

most important spawning grounds in Bushehr waters. A sex ratio of 0.84

(male/female) occurred over the study period.

A total of 2571 prawn were tagged and released using a green color polyethylene

steamer tag during the 15 month survey. From these, 68 recaptures were reported

during August-September of 2003-04, of which four were omitted from analysis

due to lack of information. Maximum recapture depths were 16-20 m and 26-30 m

for males and females, respectively. The majority of recaptures indicated that these

prawn moved at speeds of < 2 km/day. Maximum speed was recorded 3-4 km/day

for males and 2 km/day for females. Around 70% of the tagged shrimp were

recaptured within 20 km of their release locations, while approximately 10% of

them traveled a linear distance > 50 km. The maximum recorded distance was 127

km after 95 days of release for a female shrimp. The maximum time at-liberty was

381 days for a female that moved 20 km. Tag recapture data indicated shrimp

generally moved to the northern area (Bahrekan) where waters were greater in

depth.

vi

In the shallow study areas, post-spawning sampling of juvenile P. semisulcatus was

carried out during July, November and December 2003 and January, February,

April, June, July 2004 and February and March 2005. Some stations that recorded

a depth of less than 10 m were trawled using a small vessel equipped with a small

beam trawl. A total of 748 juvenile P. semisulcatus of less than 15 mm CL were

collected. These juveniles were observed in July and November 2003 and April and

June 2004. The CL of juveniles ranged from 3 mm to 15 mm and minimum of

mean of CL were 4.5 mm in November and maximum 9.6 mm in July. Observed

locations of juveniles were around vegetated sites at Mottaf (southern region) and

Helaileh (middle region), although a few juveniles were also collected in the non-

vegetated areas. Within these nursery grounds, 47% of marine fauna included

other marine species such as fishes of Sparidae and Platycephalidae family, crabs,

shells and other penaeid shrimp species such as Metapenaeus affinis and M.

stebbingi. Seasonal abundance of juveniles was greatest during the warmer months

(from April to November) and very few were captured during the colder seasons

(from December to May).

The study also used Random Amplified Polymorphic DNA (RAPD) analysis to

compare the genetic diversity of the population of prawn, P. semisulcatus. The

results showed that the genetic diversity is not significantly different and it was

confirmed that this species should be identified as one population in the Bushehr

waters.

vii

In the present study mature shrimp were dominant from December through March,

while juvenile stages were observed generally from April through July.

Exploitation rates of P. semisulcatus suggest that fishing activities should be

regulated during spawning periods and within the nursery grounds.

Abstrak tesis yang dikemukakan kepada Senat Universiti Putra Malaysia sebagai

memenuhi keperluan untuk ijazah Doktor Falsafah

BIODINAMIK DAN KITAR HIDUP UDANG (PENAEUS SEMISULCATUS DE

HAAN), DI KAWASAN PERAIRAN PANTAI BUSHEHR

TELUK PARSI

Oleh

NASSIR NIAMAIMANDI

Ogos 2006

Pengerusi: Profesor Madya Aziz bin Arshad, PhD

Fakulti: Sains

viii

Penangkapan udang di Teluk Parsi mempunyai sejarah yang panjang.

Penangkapan secara pukat tunda bagi udang penaeid telah dimulak dan

dibangunkan sepanjang kawasan pantai utara tengah Teluk Parsi (Pantai Bushehr)

pada 1960. Dari 1998 ke 2004, pendaratan udang berubah-ubah dalam lingkungan

480t dan 2700t setahun. Spesies utama udang penaeid dalam penangkapan (>80%)

dan mempunyai kepentingan ekonomi di Teluk Parsi (termasuk kawasan Pantai

Bushehr) ialah Penaeus semisulcatus, biasanya dikenali sebagai udang harimau

hijau. Lebih daripada 2000 bot tangkapan dan 33,000 nelayan dari kawasan kajian

di Bushehr telah menjalankan aktiviti penangkapan udang setiap musim (Ogos-

September) sejak beberapa abad yang lalu. Di sebelah perairan Arab Teluk Parsi,

P. semisulcatus telah menjadi satu-satunya spesies yang paling penting. Walaupun

ia mempunyai kepentingan komersil, tetapi hanya terdapat sedikit sahaja hasil

penyelidikan yang melaporkan tentang aspek biologi spesies ini. Dalam kawasan

kajian pantai Bushehr, penangkapan udang hanya dikawal semasa musim

tangkapan. Oleh sebab kekurangan maklumat, maka tidak terdapat sebarang

penyekatan dan kawasan larangan di tempat asuhan dan kawasan pembiakan dan

juga perubahan sumber udang juga adalah tidak diketahui.

Kajian ini dilakukan di kawasan Pantai Bushehr sepanjang tahun 2003-2005 dan

ianya menghuraikan tentang biodinamik dan kitar hidup udang P. semisulcatus.

Pensampelan udang dilakukan setiap bulan dengan menggunakan bot

penyelidikan R/V Lavar II dari Januari 2003 hingga Mac 2004 selama 15 bulan di

50 stesen tetap. Frekuensi panjang karapas (CL) digunakan untuk menganggar

ix

pertumbuhan , mortaliti dan kadar eksploitasi bagi udang jantan dan betina

dengan menggunakan perisian analisis statistik FiSAT. Parameter pertumbuhan

Von Bertalanffy telah masing-masing dianggarkan pada K = 1.6tahun-1

dan 2.2tahun-1

dan L∞ = 38 mm dan 50.4 mm bagi individu jantan dan betina. Kadar mortaliti

tahunan yang dianggarkan bagi individu jantan ialah Z = 6.4, F = 4.3 dan M = 2.11.

Manakala untuk individu betina ianya pada Z = 8.2, F = 5.8 dan M = 2.41. Kadar

eksploitasi bagi individu jantan dan betina adalah masing-masing E = 0.67 dan

E = 0.70. Keputusan juga mendapati jumlah nilai mortaliti dan kadar eksploitasi

yang tinggi, dan ini mengunjur yang spesies ini telah dieksploitasikan melebihi

tahap optimum. Umur maksimum yang dikira ialah 20 bulan untuk jantan dan 15

bulan untuk betina.

Kajian kematangan gonad telah menunjukkan bahawa aktiviti pembiakan utama

bagi P. semisulcatus berlaku semasa musim sejuk dan awal musim bunga, dan satu

puncak minor tambahan di musim luruh. Semua tempoh ini telah dikaitkan

dengan musim hujan di kawasan kajian Bushehr. Saiz kematangan 50% (L50%)

ialah pada 40.6 mm panjang karapas (CL). Didapati yang kurang daripada 5%

betina kecil daripada 24 mm CL.

Udang yang matang bertumpu di sektor tengah dan selatan kawasan kajian pada

kedalaman 20-30 m. Kawasan Imam hassan, Genavah dan Helleh adalah antara

kawasan pembiakan penting di Pantai Bushehr. Nisbah jantina 0.84 (jantan/betina)

didapati sepanjang waktu kajian.

x

Sejumlah 2571 udang telah ditanda dengan menggunakan tanda ‘steamer’

polietelina berwarna hijau dan dilepaskan di sepanjang 15 bulan tinjauan.

Daripada jumlah ini, 68 ekor telah di tangkap semula semasa bulan Ogos-

September 2003-04 dan empat telah diabaikan daripada analisis disebabkan

kekurangan maklumat. Kedalaman maksimum tangkapan semula masing-masing

adalah pada 16-20 m dan 26-30 m untuk individu jantan dan betina. Majoriti

daripada tangkapan semula menunjukkan bahawa udang ini bergerak pada

kelajuan <2 km/hari. Kelajuan maksimum telah dicatatkan pada 3-4 km/hari bagi

jantan dan 2 km/hari bagi betina. Lebih kurang 70% udang yang ditanda telah

ditangkap semula dalam lingkungan 20 km dari tempat yang dilepaskan,

manakala lebih kurang 10% daripada mereka bergerak dalam satu perjalanan

garis lurus >50 km. Jarak maksimum yang dicatatkan ialah 127 km selepas 95 hari

pelepasan oleh seekor udang betina. Masa bebas maksimum yang dicatatkan ialah

381 hari bagi satu betina yang bergerak sejauh 20 km. Data tangkap-tanda-lepas-

tangkap semula menunjukkan udang pada umumnya bergerak ke arah utara

(Bahrekan) di mana air adalah lebih dalam.

Di kawasan kajian yang berkedalaman cetek, pensampelan peringkat juvenil P.

semisulcatus telah dilakukan pada Julai, November dan Disember 2003 dan

Januari, Februari, April, Jun, Julai 2004 dan Februari dan Mac 2005. Stesen yang

dicatatkan mempunyai kedalaman yang kurang daripada 10 m telah dipukat

dengan menggunakan satu bot tangkapan yang dilengkapi dengan satu pukat

‘beam’. Sejumlah 748 juvenil P.semisulcatus yang kurang daripada 15 mm CL

telah dikumpulkan. Juvenil ini telah direkodkankan pada bulan Julai dan

xi

November 2003 serta April dan Jun 2004. Panjang karapas (CL) juvenil berjulat

antara 3 mm hingga 15 mm dan min CL yang minimum ialah 4.5 mm semasa

November dan maksimumnya pula 9.6 mm pada bulan Julai. Juvenil dicatatkan

pada tempat yang bervegetatif di Mottaf (bahagian selatan) dan Helaileh (bahagian

tengah), walaupun begitu terdapat juga juvenil yang dikumpulkan di tempat tidak

bervegetatif tetapi dengan jumlah yang sedikit.

Daripada kajian tempat asuhan, sebanyak 47% faunanya terdiri daripada spesies

marin lain seperti ikan daripada famili Sparidae dan Platycephalidae, ketam,

kerang-kerangan dan spesies udang penaeid lain seperti Metapenaeus affinis dan M.

stebbingi. Kelimpahan juvenil yang paling banyak adalah pada bulan-bulan yang

lebih panas (dari April ke November) dan jumlah yang sedikit dapat ditangkap

pada musim sejuk (dari Disember ke Mei).

Kajian ini juga menggunakan “Random Amplified Polymorphic DNA” (RAPD)

analisis untuk membandingkan kepelbagaian genetik dalam populasi udang P.

semisulcatus. Keputusan kepelbagaian genetik tidak menunjukkan perbezaan

bererti dan ini memastikan bahawa spesies ini patut dikenalpasti sebagai satu

populasi sahaja di kawasan pantai Bushehr.

Dalam kajian ini, udang matang didapati dominan pada Disember hingga Mac,

manakala fasa juvenil pada umumnya diperhatikan pada April hingga Julai.

Kadar eksploitasi P. semisulcatus mengesyorkan yang aktiviti penangkapan udang

harus dikawal terutamanya semasa waktu pembiakan dan juga di tempat asuhan.

xii

ACKNOWLEDGEMENTS

xiii

I would like to express my sincere thanks to Associate Prof. Dr Aziz Bin Arshad,

the chairman of my supervisory committee for his helpful advices, comments and

confidence in me throughout this study. I would not have been able to continue my

study without these supports and encouragements.

I appreciate the helps of my supervisory committee members, Associate Prof. Dr

Siti Khlijah Daud and Associate Prof. Dr. Che Roos Saad for their supports and

assistance. I am also grateful to Dr Bahram Hassanzadeh Kiabi for his help during

my research in Iran. This research would not have been possible without their

supports.

I also wish to thanks the Head of the Department of Biology in the Faculty of

Science and staff and also Head of the Graduate School Organization and staff for

their help during my study in Universiti Putra Malaysia.

Further thanks to the head and staff of the stock assessment division of Iran

Shrimp Research Center, especially to Mr. Kambozia Khorshidian, Abdol Rasool

Esmaeeli, Rajab Khodadadi, Mehrdad Hossaini and Yadolah Bayat for their help

in sampling, data collection and tagging program and also the cooperation and

assistance of captain Balal Zadeh and the crew of research vessel Lavar II are

gratefully acknowledged.

xiv

I thank those members and fishermen in Dayer, Ameri, Bushehr and Ganaveh who

have participated in the research through the return of tagged prawns. I also like to

acknowledge to Mr Farokh Ansari for his cooperation and assistance during the

research.

I further wish to thank Dr. John Paul Hoolihan for reading the manuscript and

making a number of helpful suggestions and corrections.

This research was funded and supported by Iranian Fisheries Research Institute

(IFRO). I wish to express my sincere thanks to Dr Sohrab Rezvani, the Director, Dr

Mohammad Afsharnasab, the Research Deputy and Mr. Rostam Pour, the

Financial Deputy of IFRO who have been supporting me throughout the period of

my study.

Last but not least, special thanks go to my wife Noushin Farhadi for her prayers,

encouragement, and spiritual support during my study and research.

xv

I certify that an Examination Committee has met on 25 August 2006 to conduct the

final examination of Nassir Niamaimandi on his Doctor of Philosophy thesis

entiteled " Bio-Dynamics and Life Cycle of Shrimp (Penaeus semisulcatus De

Haan) in Bushehr Coastal Waters of the Persian Gulf" in accordance with

Universiti Pertanian Malaysia (Higher Degree) Act 1980 and Universiti Pertanian

Malaysia (Higher Degree) Regulations 1981. The Committee recommends that the

candidate be awarded the relevent degree. Members of the Examination

Committee are as follows:

Siti Shapor Hj. Siraj, PhD

Associate Professor

Faculty of Science

Universiti Putra Malaysia

(Chairman)

Japar Sidik Bujang, PhD

Associate Professor

Faculty of Science


(Internal Examiner)

Mustafa Kamal Abd. Satar, PhD

Associate Professor

Faculty of Agriculture


(Internal Examiner)

Mohd Azmi Ambak, PhD

Professor

Institute of Tropical Aquaculture

Kolej Universiti Sains dan Teknologi Malaysia

(External Examiner)

HASANAH MOHD. GHAZALI, PhD

Professor/Deputy Dean

School of Graduate Studies

xvi


Date: 08 February 2007

This thesis submitted to the Senate of Universiti Putra Malaysia and has been

accepted as fulfilment of the requirement for the degree of Doctor of Philosophy.

The members of the Supervisory Committee are as follows:

Aziz Bin Arshad, PhD

Associate Professor

Faculty of Science


(Chairman)

Siti Khalijah Daud, PhD

Associate Professor

Faculty of Science


(Member)

Che Roos Bin Saad, PhD

Associate Professor

Faculty of Agriculture


(Member)

Bahram Hassanzadeh Kiabi, Ph.D.

Associate Professor

Faculty of Science

Shaheed Beheshti University

(Member)

AINI IDERIS, PhD

Professor/Dean

School of Graduate Studies


Date:

xvii

DECLARATION

I hereby declare that the thesis is based on my original work except for quotations

and citations which have been duly acknowledged. I also declare that it has not

been previously or currently submitted for any other degree at UPM or other

institutions.

NASSIR NIAMAIMANDI

Date: 24 August 06

xviii

TABLE OF CONTENTS

Page

DEDICATION ii

ABSTRACT iii

ABSTRAK vii

ACKNOWLEDGEMENTS xii

APROVAL xiv

DECLARATION xvi

LIST OF TABLE xix

LIST OF FIGURES xxi

LIST OF ABBREVIATION/NOTATION/GLOSSARY OF TERMS xiv

CHAPTER

I INTRODUCTION 1

The Shrimp Fishery in the Persian Gulf

2

The Shrimp Fishery in the Bushehr Province

4

Effort and Catch

5

Study Objectives

9

II LITERATURE REVIEW 11

Biology of Penaeus semisulcatus

11

Taxonomy and Distinctive Characters

11 Life Cycle

13 Feeding 17

Effects of Environmental Factors

18 Salinity and Temperature

18

Rainfall

20

Research Activities on the Shrimp

21

xix

Population Dynamics

22

Reproductive Biology

26

Nursery Grounds

34

Movement and Migration

41

Population Genetics

46

III POPULATION DYNAMICS 53

Introduction

53

Materials and Methods

55

Estimation of Growth Parameters (K, L∞)

59

Estimation of Natural Mortality (M), Total Mortality (Z),

60 Fishing Mortality (F), Exploitation Rate and

Maximum Age (Tmax)

Results and Discussion

62

Growth Parameters

62

Mortality Parameters and Maximum Age

70

Environmental parameters

71

IV REPRODUCTIVE BIOLOGY 84

Introduction

84


86


89

Spawning Seasons and Mature Size

89

Spawning Areas

94

Sex Ratio 94

V MOVEMENT AND MIGRATION 105

Introduction

105

xx


107


111

Release and Recapture

111

Component of Movement

112

VI NURSERY GROUNDS 125

Introduction

125


127

Study Area

127

Sampling Methods

127


131

Seasonal Distribution

131

Size Distribution

131

Nursery Areas

133

Non Prawn Organisms

136

Environmental Parameters

137

VII POPULATION GENETICS 146

Introduction

146


148

DNA Extraction

148

Measurement of DNA Concentration

150 Amplification Conditions and Electrophoresis

150

Results and Discussion 152

VIII GENERAL DISCUSSION AND CONCLUSION 159

REFERENCES/BIBLIOGRAPHY 172

xxi

APPENDICES 194

BIODATA OF THE AUTHOR 202

LIST OF TABLES

Table Page

1.1. Total landing of shrimp in Bushehr waters (1998-2004)

2

1.2. Percentage of shrimp landing in Bushehr waters according to species

8

(1998-2004)

2.2. The natural mortality rates of P. semisulcatus in Kuwait waters

25

Estimated by method developed by Xu et al. (1995)

3.1. Trawl sampling stations in Bushehr area, Persian Gulf (2003-2004).

58

3.2. Comparison of growth estimated for P. semisulcatus from different areas

78

of the Persian Gulf

4.1. Monthly percentage of gonad stages in females of P. semisulcatus in

92

Bushehr waters, Persian Gulf (2003-2004)

4.2. Numbers of females, P. semisulcatus with ripe ovaries expressed

93

as a proportion to total sample from Bushehr waters, Persian Gulf

(2003-2004)

4.3. Sex ratio (male/female) of P. semisulcatus in Bushehr waters,

97

Persian Gulf (2003-2004)

5.1. Number of tagged and recaptured of males and females P. semisulcatus in

111


xxii

5.2. Percentage of release and recaptured males and females P. semisulcatus in

113

different depths of Bushehr waters, Persian Gulf (2003-2004)

6.1. Sampling stations for nursery grounds of P. semisulcatus in Bushehr

129

coastal waters, Persian Gulf (2003-2004)

6.2. Mean carapace length of juvenile (<15 mm) of juvenile P. semisulcatus

133

collected along Bushehr coastal waters, Persian Gulf (2003-2004)

6.3. Percentage of species composition (by number) in the nursery

137

grounds of P. semisulcatus in Bushehr coastal waters,


6.4. Percentage of P. semisulcatus during nursery periods and other shrimp

137

and marine species in Bushehr coastal waters, Persian Gulf (2003-2004)

7.1. Sequence and operon codes of OPA primers used for Penaeus semisulcatus

153

in Bushehr waters, Persian Gulf (2004-2005)

7.2. Number of amplicons and number of polymorphic bands produced by

153

each RAPD primer for population of P. semisulcatus in the Bushehr

waters, Persian Gulf (2004-20050

xxiii

LIST OF FIGUERS

Figure Page

2.1. Schematic diagram of type 3 life cycle of penaeid shrimps 15

(Dall et al., 1990)

3.1. Location of study area and sampling station for P. semisulcatus

57

in Bushehr waters (Persian Gulf)

3.2. Length-frequency curve of males P. semisulcatus in Bushehr waters,

63


3.3. Length-frequency curve of female P. semisulcatus in Bushehr

63

waters, Persian Gulf (2003-2004)

3.4. Relationship of total length and carapace length in males

65

P. semisulcatus in Bushehr waters, Persian Gulf (2003-2004)

xxiv

3.5. Relationship of total length and carapace length in females

65


3.6. Relationship of total length and weight in males P. semisulcatus in

66


3.7. Relationship of total length and weight in females P. semisulcatus

66


3.8. Relationship of carapace length and weight in males P. semisulcatus

67


3.9. Relationship of carapace length and weight in females P. semisulcatus

67


3.10. Average carapace length of male and female P. semisulcatus in

69


3.11. The length-converted catch curve of male P. semisulcatus in Bushehr

72

waters, PersianGulf (2003-2004)

3.12. The length-converted catch curve of female P. semisulcatus in

72


3.13 Average of salinity (ppt), SST (°C) and rainfall (mm) during different

74

months in Bushehr waters, Persian Gulf (2003-2004)

4.1. Different sexes female and male of P. semisulcatus

88

4.2. Gonad stages of penaeid prawns

88

4.3. Percentage of mature females (3, 4 stages) P. semisulcatus in

91


xxv

4.4. Logestic relationship between the number of matured females and

95

carapace length (mm) in Bushehr waters, Persian Gulf (2003-2004)

4.5. Relationship between percentage of ovary maturation (stages 3 and 4) of

95

female P. semisulcatus and carapace length (mm) in Bushehr waters,


4.6. Spawning grounds of P. semisulcatus in Bushehr waters,

96


5.1. Different process of tagging of P. semisulcatus in Bushehr waters,

109


5.2. Comparison of released and recaptured depths of males, P. semisulcatus

112


5.3. Comparison of released and recaptured depths of females,

112


5.4. Comparison of movement speeds of males and females of recaptured

116

prawn, P. semisulcatus in Bushehr waters, Persian Gulf (2003-2004)

5.5. Percentage of movement of tagged prawn, P. semisulcatus in

116


5.6. Comparison of days at large for recaptured male and female

118


5.7. Movement of tagged shrimp, P. semisulcatus recovered along the

119

Bushehr coastal waters, Persian Gulf (2003-2004)

6.1. Shallow water sampling areas for nursery grounds of P. semisulcatus

126

in Bushehr coastal waters, Persian Gulf (2003-2004)

6.2. Percent of size distribution of juvenile P. semisulcatus in Bushehr

132

xxvi

coastal waters, Persian Gulf (2003-2005)

6.3. Nursery grounds of P. semisulcatus in Bushehr coastal waters,

134


6.4. Mean temperature (°C) and salinity (ppt) in nursery area of

139

P. semisulcatus in Bushehr waters, Persian Gulf (2003-2005).

7.1.Sampling areas for P. semisulcatus in Bushehr coastal waters,

149

Persian Gulf (2004)

7.2. PCR generated by primers OPA-04 (A) and OPA-09 (B) for

154

P. semisulcatus population in Bushehr waters, Persian Gulf (2004).

M contained DNA size in OPA-04 lanes 1-6 Daylam and lanes 7-10

Helaileh areas and in OPA-09 lanes 1-5 Daylam and lanes 6-11 Helaileh

areas.

OPA showed one polymorphism arrow

xxvii

LIST OF ABBREVIATIONS/SYMBOLS/GLOSSARY OF TERMS

R/V Research Vessel

K Growth Coefficient

L∞ Asymptotic Length

FiSAT Fish Stock Assessment Toll

F Fishing Mortality

M Natural Mortality

Z Total Mortality

E Exploitation Rate

FAO Food and Agriculture Organization

CPUE Catch Per Unit Effort

hp hours power

UNDP United Nation Development Program

ELEFAN Electronic Length Frequency Analysis

CL Carapace Length

TL Total Length

mtDNA mitochondrial DNA

RFLP Restriction Fragment Length Polymorphism

xxviii

PCR-SSCP Single Standard Conformation Polymorphism

EST Sea Surface water Temperature

ESP Explained Sum of Peaks

ASP Available Sum of Peaks

SST Sea Surface Water Temperature

Lat Latitude

Log Longitude

Lt Length at time t

Tmax Maximum age

C Amplitude of oscillation

WP Winter Point

W Weight

Ø fi prime

ppt part per thousand

L50% Length at 50% Maturity

X² Chi-Square

PFI Population Fecundity Index

GPS Global Positioning System

RFLP Restriction Fragment Length Polymorphism

% Percentage

∑ Summation

≈ Approximately

m Meter

cm Centimeter

xxix

mm Millimeter

ml Milliliter

g Gram

kg Kilogram

nm Nautical Mile

ph Hydrogen ion

e.g For example

hp/day Hource Power Per Day

cm/sec Centimeter Per Second

h Hour

< Less than

> More than

°C Degree Celsius

Per.com Personal communication

Rmp round per minute

EDTA Ethylane Diamine Tetracetic Acid

mm milimoul

SDS Sodium Dodecil Solphat

PCI Phenol, Chloroform, Isoamil alchohol

nm nanomolar

OD Optic Density

TBR Tris Br

ng/µl nanogram/microlitre

PCR Polymorase Change Reaction

xxx

bp base pair

min minute

µg microgram

DNA Deoxyribonucleic acid

V Volt

UV Ultra Violet

bio-dynamics and life cycle of shrimp ...psasir.upm.edu.my/544/1/1600396.pdfiv the present study was...

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