ECOLOGICAL STUDY OF THE MOLLUSCAN FAUNA ON INTERTIDAL ARTIFICIAL ROCKY SHORE OF TARKWA BAY AREA, LAGOS, NIGERIA

BY

ERUTEYA, ONOSIVBAKPO JONATHAN

`

FEBRUARY 2014

II

ECOLOGICAL STUDY OF THE MOLLUSCAN FAUNA

ON INTERTIDAL ARTIFICIAL ROCKY SHORE OF

TARKWA BAY AREA, LAGOS, NIGERIA

A thesis submitted for the degree of Doctor of Philosophy (Marine Biology)

BY

ERUTEYA, ONOSIVBAKPO JONATHAN

B.Sc, M.Sc (Zoology) Lagos.

MATRIC NO: 830809031

DEPARTMENT OF MARINE SCIENCES

III

IV

DEDICATION

This project is dedicated to Almighty God for His divine guidance over my life, and members of my immediate family.

V

ACKNOWLEDGEMENTS

I am indeed grateful for the help and supports I received from numerous people in various institutions during the course of this project. Firstly, I would like to thank my supervisors:

Associate Professor Clement Aghatise Edokpayi for his kindness, thoughtfulness, sense of humour and generous support, which made the work for this study more pleasurable. His sharing of ideas has been and will, hopefully, continue to be one of my most enjoyable experiences.

Again, I would also like to thank my second supervisor, Associate Professor J. K Saliu, who also supervised my M.Sc. thesis. Both of them have supported me throughout, and provided me with sound advice and opportunities that broaden my horizon.

I would like to thank Professors: K. K Ikusemiju, D. I. Nwankwo, Rosemary Egonmwan, and

L. O. Chukwu for their generous supports, for editing of my thesis, award of Graduate Fellow

Scholarship and provisions of scientific journals. They have also continually showed immense optimism and unswerving faith in my abilities.

Thanks to Dr (Mrs) A.O. Osibona, Dr S.O. Ayoola, Dr A.A Otitoloju, Dr A. Z. Aderolu, Dr

(Mrs) A. O. Lawal-Are, Dr M. P. Kuton, Dr I. C. Onyema, Dr Efe Uwadia, Dr M O Lawal and Dr Segun for their friendly supports and encouragement. I would also like to thank the non-teaching staff: Mrs Olutimilehin, Mr Lawal, Mr Coker (Marine Sciences) and Mr

Bengusa (Chemistry Department), Mr Uncle B, a retired Laboratory Technologist (Zoology

Department).

I, also extend my appreciation to my fellow Ph.D students: especially Mr Joseph Nkwoji, and Mr Uzoh (Marine sciences) for their moral support.

VI

Finally, I would also like to extend my appreciation to all members of my family, more importantly my children (Emmanuel, Josephine and Deborah), they are wonderful team. I cherish their faith in me. I also cherish the love and support of my wife, Mrs Eunice Esther

Eruteya, during the course of this project. This thesis is dedicated to her and my children.

Above all, I thank the Almighty God for His strength and Mercy upon my life without which

I can do nothing (Luke 1:37).

VII

TABLE OF CONTENTS

CONTENTS PAGES

CERTIFICATION………………………………………………………………………. .II

DEDICATION…………………………………………………………………………... 11I

ACKNOWLEDGEMENTS……………………………………………………………….1V

TABLE OF CONTENTS………………………………………………………………….V1

LIST OF TABLES……………………………………………………………………...... XIV

LIST OF FIGURES…………………………………………………………………….XV1II

LIST OF PLATES……………………………………………………………………..XX111

LIST OF APPENDICES…………………………………………………………………XXV

OPERATIONAL DEFINITION OF TERMS…………………………………………XXXII

LIST OF ABBREVIATIONS…………………………………………………………XXXIII

ABSTRACT…………………………………………………………………………...XXXIV

CHAPTER ONE

I.0 INTRODUCTION…………………………………………………………………..1

1. I BACKGROUND OF THE STUDY…………………………………………..1

1. 2 STATEMENT OF PROBLEM……………………………………………….5

1. 3 AIM AND SPECIFIC OBJECTIVES OF STUDY……………………...... 6

1. 4 SIGNIFICANCE OF THE STUDY…………………………………………..7

1. 5 RESEARCH QUESTIONS………………………………………………...... 8

VIII

CHAPTER TWO

2. 0 LITERATURE REVIEW………………………………………………………….9

2. 1 PHYSICO-CHEMICAL CHARACTERISTICS……………………………..9

2. 2 ROCKY SHORES: ENVIRONMENTAL CONDITIONS………………….11

2. 3 MOLLUSCAN COMMUNITIES AND ADAPTATIONS……………….....11

2. 4 INVERTEBRATE COMMUNITIES OF LAGOS HABOUR AND

GULF OF GUINEA………………………………………………………….12

2.5 BIOLOGY OF INTERTIDAL MOLLUSCS………………………………..13

2.5.1 Feeding………………………………………………….….…….13

2.5.2 Gut Content Analyses………………………………………...... 13

2.5.3 Growth of Intertidal Molluscs…………………………………...14

2.5.4 Heavy metal concentrations: seawaters, seaweeds and

and selected molluscs……………………………………………16

2.5.5 Reproduction in intertidal molluscs…………………………….18

2.6 EFFECT OF PHYSICAL AND PHYSIOLOGICAL STRESS ON

THE DISTRIBUTION OF INTERTIDAL MOLLUSCS…………………...19

2.6.1 Effect of sea surface temperature (SST) on intertidal

molluscs………………………………………………………….21

2.6.2 Ecological Implications of shore Height on molluscs…….……..22

2.6.3 Ecological Effects of Shell Size and Shape of Intertidal

molluscs……………………………………………………...... 24

2.6.4 Ecological Implications of molluscan faunal shell

IX

colours…………………………………………………………...25

2.6.5 Economic Importance of Intertidal Molluscs……………………26

2.6.6 Strategies for molluscan conservation and sustainable

utilization………………………………………………………….29

CHAPTER THREE

3.0 MATERIALS AND METHODS……………………………………………………31

3.1 DESCRIPTION OF THE STUDY AREA……………………………………..31

3.1.1 West mole………………………………………………………………33

3.1.2 Training mole……………………………………...... 35

3.1.3 East mole………………………………………………………………37

3.2 SAMPLING DESIGN………………………………………………………….39

3.2.1 Site selections………………………………………………………...39

3.2.2 Collection of seawater………………………………………………...44

3.2.3 Collection of algal samples…………………………………………...44

3.2.4 Collection of molluscan samples……………………………………..44

3.2.5 Choice of study species………………………………………………..45

3.2.6 Sample preservations………………………………………………….46

3.3 SAMPLE ANALYSES……………………………………………………………46

3.3.1 Water analyses………………………………………………………....46

3.3.2 Air and Water Temperatures…………………………………….…….46

X

3.3.3 Rock Temperature…………………………………………………...... 46

3.3.4 Salinity……………………………………………………………...... 46

3.3.5 Dissolved Oxygen (DO)…………………………….………………….46

3.3.6 Rainfall Data…………………………………………………………...47

3.4 LABORATORY ANALYSIS………………………..…………………………..47

3.4.1 Nutrient Analyses…………………………………………………....47

3.4.2 Heavy metal analysis………………………………………………..47

3.5 STUDIES ON THE BIOLOGY OF INTERTIDAL MOLLUSCS…………. …48

3.5.1 Feeding in selected intertidal molluscan species…………. ………...48

3.5.2 Foraging behaviours of intertidal molluscs………… ………………...48

3.5.3 Gut Content Analysis……………… ………………………………....49

3.5.4 Comparative studies on growth of selected molluscan species at three

different tidal zones…………………………………………………...49

3.5.5 Determination of wet, dry biomass and Ash-free Dry-

Weight (AFDW)………………………………………………………50

3.5.6 Effects of interspecific competition on molluscs’ growth rate

in the laboratory………………………………………………………50

3.5.7 Studies on shell dimensions of selected molluscs……………………51

3.5.8 Studies on the reproductive biology of selected molluscs………….....51

3.5.8.1 Gonadosomatic Index (GSI)……………………….....52

3.5.8.2 Reproductive Effort (R.F)……………….…………….53

3.5.8.3 Histology of the gonads of selected molluscs….. …53

XI

3.5.8.4 Sex Ratios…………………………………………….54

3.5.8.5 Size at first sexual maturity…………………………..54

3.6 Statistical Analysis……………………………………………………54

3.6.1 Molluscan Data analysis………………………………………………..55

CHAPTER FOUR

4. 0 RESULTS……………………………………………………………………………...57

4.1 PHYSICO-CHEMICAL CHARACTERISTICS OF TARKWA BAY…..57

4.1.1 Temperature……………………………………………………………………57

4.1.2 Salinity and Rainfall Patterns………………………………………………....67

4.1.3 Hydrogen Ion Concentration (pH)…………………………………..……….69

4.1.4 Dissolved Oxygen (DO)……………………………………………………..70

4.1.5 Biochemical Oxygen Demand (BOD)5………………………………… ….72

4.1.6 Chemical Oxygen Demand………………………………………………….74

4.1.7 Conductivity…………………………………………………………………76

4.1.8 Nutrients………………………….…………………………………………77

4.1.9 Heavy metals……………………………………………………………….80

4.2.0 ENVIRONMENTAL FLUCTUATIONS IN INTERTIDAL

ROCK POOLS OF TARKWA BAY……………………………………..………82

4.2.I Thermohaline fluctuations in three different intertidal

rock pools at Tarkwa Bay………………………….……...... 82

4.2.2 Thermal variations in air, rock surface and limpet´s

XII

foot…………………………………………………………..85

4.3.0 HEAVY METAL STUDIES……………………………………………………...86

4.3.1 Heavy metal concentrations in seawaters, seaweeds,

and selected molluscs…………………….…………………...87

4.3.2 Heavy metal concentrations in different body parts of

molluscs……………………………………………..……….90

4.4.0 COMPOSITION, DISTRIBUTION, ABUNDANCE, DIVERSITY AND

SEASONALITY OF MOLLUSCAN COMMUNITIES OF TARKWA

BAY INTERTIDAL ROCKY SHORE……………………………………..…….108

4.5.0 UNIVARIATE ANALYSIS OF MOLLUSCAN ECOLOGY AT THREE

MOLES OF TARKWA BAY………...... 129

4.5.1 Shannon´s Index (H’)………………………………………………...... 129

4.5.2 Pielou’ Evenness (J’)………………………………………….…...... 132

4.5.3 Simpson’ Dominance Index…………………………………………...... 135

4.5.4 Size- frequency distributions of selected molluscs…..…………………138

4.5.4.1 Patella safiana…………………………………………….....138

4.5.4.2 Nerita senegalensis………………………………………...... 138

4.5.4.3 callifera…………………………………………………139

4.5.4.3 Buccinum undatum…………………………………………..139

4.6.0 BIOLOGY OF INTERTIDAL MOLLUSCS……………………………….…….145

4.6.1 Studies on shell dimensions on selected molluscs……………………145

4.6.2 Gut Content Analysis………………………………………………….168

XIII

4.6.3 Foraging behaviours of selected intertidal molluscs………………….170

4.6.4 Studies on molluscs’ growth at different tidal zones……. ………….172

4.6.5. Effects of intra-and interspecific competitions on growth

rate of selected species at different environments………. ………..174

4.6.6 Growth rates between males, females and in comparison

to a mixed sex molluscan populations……………………… .. …….175

4.7.0 BIOMASS STUDIES ON INTERTIDAL MOLLUSC OF

ECONOMIC INTEREST……………………………………………………177

4.7.1 West mole: Wet Biomass studies………………………………….177

4.7.2 West mole: Dry Biomass studies………………………………. …178

4.7.3 Training mole: Wet Biomass studies……………………………...179

4.7.4 Training mole: Dry Biomass studies…………...... 179

4.7.5 East mole: Wet biomass studies………………………………….180

4.7.6 East mole: Dry biomass studies………………...... 181

4.7.7 Comparison of wet and dry biomass of the three moles…………182

4.8.0 REPRODUCTION IN SELECTED INTERTIDAL MOLLUSCS……………….189

4.8.1 Gonado somatic Index (GSI) in Patella safiana…………………189

4.8.2 Sex Ratio in Patella safiana………………………………………190

4.8.3 Reproductive efforts (RE)……………….……………………...... 191

4.8.4 Gonadosomatic Index (GSI) in Nerita senegalensis……………...193

4.8.5 Sex Ratio in Nerita senegalensis………………………………….194

XIV

4.8.6 Reproductive Effort (R.E) in Nerita senegalensis………………..196

4.8.7 Size at sexual maturity……………………………………………..198

CHAPTER FIVE

5.0 DISCUSSION…………………………………………………………………. …....199

CHAPTER SIX

6.0 SUMMARY AND CONCLUSION……………………………………………… …215

6.1 SUMMARY OF RESEARCH FINDINGS…………………………………...215

6.2 RECOMMENDATIONS…………………………………………………….216

6.3 CONTRIBUTIONS TO KNOWLEDGE………………….…………………218

6.4 CONCLUSION……….………………………………………………………219

REFERENCES…………………………………………………………………………….221

APPENDICES……………………………………………………………………………..246

LIST OF TABLES

CONTENTS PAGES

Table 1: GPS locations of 12 sampling stations and their co-ordinates at the three

moles of Tarkwa Bay……………………………………………………………40

Table 2: Ecological features of the sampling stations at the three moles XV

of Tarkwa Bay……………………………………………………………………..41

Table 3: Temporal variations in physico-chemical parameters investigated in the dry

season at the three moles of Tarkwa Bay…………………………………………59

Table 4: Temporal variations in physico-chemical parameters investigated in rainy

season at the three moles of Tarkwa Bay ………………………………………..60

Table 5: Correlation between physico-chemical parameters of the study stations

of the West mole of Tarkwa Bay……………………………………………….61

Table 6: Correlation between physico-chemical parameters of the study stations

of the Training mole of Tarkwa Bay………………………………………….62

Table 7: Correlation between physico-chemical parameters of the study stations

of the East mole of Tarkwa Bay……………………………………………….63

Table 8: Mean seasonal concentration of heavy metals in seawater, seaweed

and Patella safiana at the Training and West mole of Tarkwa Bay………88.

Table 9: Concentrations of heavy metals in different body parts of the

selected molluscan species investigated during the rainy season………….106

Table 10: Concentrations of heavy metals in different body parts of the

selected molluscs during the dry season…………………………………..107

Table 11: Composition, distribution, abundance, diversity and seasonality of

XVI

molluscan communities of the three moles of Tarkwa Bay……………….120

Table 12 Summary of seasonal variations in the number of individuals

recorded at the three moles of Tarkwa Bay………………………………..126

Table 13: Variations in the number of species recorded during the dry season at

the 3 moles of Tarkwa Bay…………………………………………………127

Table 14: Variation in the number of individuals recorded during the rainy season at the

3 moles of Tarkwa Bay………………………………………………………128

Table 15: Description of shell variables for Nerita senegalensis, Thais callifera

Buccinum .undatum…………………………………………………………145

Table 16: Descriptive statistics for Ash-free-dry Weight (AFDW)

and corresponding Shell length of Buccinum undatum (N=70)………….149

Table 17: Correlation between Log-transformed values of Ash-Free-Dry-Weight

(AFDW) against Shell Length (SL) for Buccinum undatum. Re-

Transformed value of AFDW and SL are provided in the form of a

conventional Allometric equation: W =aLb………………………………149

Table 18: Correlation between Log-transformed values of Ash-Free –Dry-Weight

(AFDW) against shell-length (SL) for Thais callifera. Retransformed

values of AFDW and SL are provided in the form of a conventional

XVII

allometric Equation: W = aLb……………………………………………155

Table 19: Correlation between Log-transformed values of Ash-Free-Dry-Weight

(AFDW) against shell length (SL) for Nerita senegalensis. Re-transformed

values of AFDW and SL are provided in the form of a

conventional Allometric equation W =aLb……………………………….161

Table 20: Laboratory observations on the feeding rates of two

herbivorous limpet……………………………………………………167

Table 21: Summary of mean values of foraging parameters for parameters for

Patella safiana during the dry and rainy seasons………………………..171

Table 22: Temporal variations in wet biomass (gm/m2) of ten dominant species of

molluscan fauna for laboratory analyses from the West mole……………183

Table 23: Temporal variations in dry biomass (gm/m2) of ten dominant species of

molluscan fauna for laboratory analyses from the West mole……………184

Table 24: Temporal variations in wet biomass (gm/m2) of ten dominant species of

molluscan fauna for laboratory analyses from the Training mole………185

Table 25: Temporal variations in dry biomass (gm/m2) of ten dominant species of

molluscan fauna for laboratory analyses from the Training mole …….186

Table 26: Temporal variations in wet biomass (gm/m2) of ten dominant species of

XVIII

molluscan fauna for laboratory analyses from the East mole……………187

Table 27: Temporal variations in dry biomass (gm/m2) of ten dominant species of

molluscan species fauna for laboratory analyses from the East mole…….188

Table 28: The estimated proportion of male and female Patella safiana in monthly

samples from the West mole of Tarkwa Bay ……………………………190

Table 29: Monthly reproductive effort of Patella safiana…………………………..192

Table 30: The estimated proportions of male and female Nerita senegalensis in

monthly samples from the West mole of Tarkwa Bay…………………..195

Table 31: Mean reproductive effort (RE) for male and female Nerita senegalensis

from the West mole of Tarkwa Bay……………………...... 197

LIST OF FIGURES

CONTENTS PAGE

Figure 1: Map of the study areas showing the three moles of Tarkwa Bay…………….32

Figure 2: Temporal variations in Air temperature at the three moles of Tarkwa bay ……64

Figure 3: Temporal variations in Water temperature at the three moles of Tarkwa Bay...65

Figure 4: Temporal variations in Rock temperature at the three moles of Tarkwa Bay …66

Figure 5: Variation of molluscan density with rainfall and salinity………………………68

XIX

Figure 6: Temporal variations in salinity at the three moles of Tarkwa Bay……………..69

Figure 7: Temporal variations in pH at the three artificial breakwaters of Tarkwa Bay…..70

Figure 8: Temporal variations in Dissolved Oxygen (DO) at the three moles

of Tarkwa Bay………………………………………………………………….71

Figure 9: Temporal variations in Biochemical Oxygen Demand (BOD5) at the three

moles of Tarkwa Bay ………………………………………………………...73

Figure 10: Temporal variations in Chemical Oxygen Demand (COD) at the three

moles of Tarkwa Bay………………………………………………………….75

Figure 11: Temporal variations in Conductivity (µS/cm) at the three moles of

Tarkwa Bay ……………………………………………………………76

2= Figure 12: Temporal variations in Nutrient (Sulphate-SO4 ) at the three moles

of Tarkwa Bay……………………………………………………………….79

Figure 13: Temporal variations in nutrients (Phosphate-Phosphorus: PO4-P) at the

three moles of Tarkwa Bay………………………………………………….80

Figure 14: Temporal variations in Nutrients (Nitrate: NO3-N) at the three moles

of Tarkwa Bay (April, 2008- March, 2010)………………………………81

Figure-15 Diurnal variations in temperature at three Rock pools at 3 different tidal

zones in the West mole of Tarkwa Bay…………………… …………………83

XX

Figure-16 Diurnal variations in salinity at three rock pools at 3 tidal zones

at the West mole of Tarkwa Bay……………………………………………..84

Figure-17 Temporal variations in Air, rock pool, rock surface and limpet temperature

at the tidal zones at the West mole of Tarkwa Bay………………………..86

Figure - 18 Mean seasonal variations of heavy metals in different body parts of

Nerita senegalensis……………………………………………………….91

Figure-19 Mean seasonal variations of lead in different body parts of

Nerita senegalensis……………………………………………………………92

Figure-20 Mean seasonal variations of zinc in different body parts of

Nerita senegalensis ………………………………………………………93

Figure-21 Mean seasonal variations of Manganese in different body parts of

Thais callifera……………………………………………………………….94

Figure-22 Mean seasonal variations of lead in different body parts of

Thais callifera………………………………………………………………95

Figure-23 Mean seasonal variations of chromium in different body parts of Thais

Callifera…………………………………………………………………………….96

Figure-24 Mean seasonal variations of Iron in different body parts of

XXI

Thais callifera………………………………………………………………………97

Figure-25 Mean seasonal concentrations of zinc in different body parts of

Thais callifera………………………………………………………………...98

Figure-26 Mean seasonal concentrations of copper in different body parts of

Thais callifera……………………………………………………………….99

Figure-27 Mean seasonal concentrations of Manganese in different body parts of

Buccinum undatum …………………………………………………………100

Figure-28 Mean seasonal concentrations of lead in different body parts of

Buccinium undatum………………………………………………………………..101

Figure-29 Mean seasonal concentrations of chromium in different body parts of

Buccinium undatum……………………………………………………102

Figure-30 Mean seasonal concentrations of Iron in different body parts of

Buccinium undatum……………………………………………………103

Figure-31 Mean seasonal variations of zinc in different body parts of

Buccinum undatum…………………………………………………….104

Figure-32 Mean seasonal concentrations of copper in different body parts of

Buccinium undatum……………………………………………………..105

Figure-33 Effects of rainfall on salinity at the three moles of

XXII

Tarkwa Bay……………………………………………………………..109

Figure-34 Mean number of molluscan species /m2 recorded at the three moles

of Tarkwa Bay…………………………………………………………110

Figure-35 Mean number of molluscan species/m2 at the Training mole …………111

Figure-36 Mean number of molluscan species/m2 of East mole …………………..112

Figure-37 Mean number of molluscan individual/m2 at West mole ……………….113

Figure-38 Mean number of molluscan individuals/m2 at the Training mole………..114

Figure-39 Mean number of molluscan individuals/m2 at the East mole …………..115

Figure: 40 Temporal variations in Shannon Wiener Index (H1) computed for

the three moles of Tarkwa Bay…………………………………………116

Figure -41 Temporal variations in Dominance Index (C) computed for the three

moles of Tarkwa Bay……………………………………………………117

Figure-42 Temporal variations in Evenness Index (J1) computed for the three

of Tarkwa Bay …………………………………………………………….118

Figure-43 Temporal variations in Shannon’s Index at 12 sampled stations of the three

moles of Tarkwa Bay ……………………………………………………..131

Figure-44 Temporal variations in Evenness Index computed for12 sampled

stations at the three moles of Tarkwa Bay………………………………..134

XXIII

Figure-45 Temporal variations in Simpson’s Index at 12 sampled stations for the

three moles of Tarkwa Bay ……………………………………………..137

Figure-46 Size-frequency distribution of Patella safiana at the three

moles of Tarkwa Bay…………………………………………………...140

Figure-47 Size-frequency distributions of Nerita senegalensis at the three

moles of Tarkwa Bay…………………………………………………….141

Figure-48 Size-frequency distribution of Thais callifera at the three

moles of Tarkwa Bay………………… ………………………………..143

Figure-49 Size-frequency distributions of Buccinum undatum at the three

of Tarkwa Bay………………………………………………………….....144

Figure-50 Shell dimensions of Buccinum undatum…………………………………..145

Figure-51 Shell dimensions for Thais callifera……………………………………....146

Figure-52 Shell dimensions for Nerita senegalensis………………………………….147

Figure-53 Relationship between Log Ash-Free-Dry-Weight (AFDW) and Log Shell

Length (SL) in Buccinum undatum……………………………………….150

Figure-54 Shell variables relationships for shell length Vs. shell width for

Buccinum undatum ……………………………………………………..151

Figure-55 Shell variable relationships for shell height Vs. in Buccinum undatum…..152

XXIV

Figure-56 Shell variable relationships for shell length Vs. Aperture length in

Buccinum undatum……………………………………………………….153

Figure-57 Shell variable relationships for aperture length Vs aperture width in

Buccinum undatum……………………………………………………….154

Figure-58 Relationship between Log Ash- Free-Dry-Weight (AFDW) and Log Shell

Length (SH) in Thais callifera…………………………………………….155

Figure-59 Shell variable relationship for shell length Vs shell width for

Thais callifera…………………………… ……………………………….156

Figure-60 Shell variable relationships for shell length Vs aperture width in

Thais callifera…………………………………………………………….157

Figure-61 Shell variable relationships for shell length Vs Aperture length for

Thais callifera………………………………………………………………158

Figure-62 Shell variable relationships for shell length Vs shell height in

Thais callifera………………………………………………………………………..159

Figure-63 Shell variable relationships for Aperture length Vs aperture width in

Thais callifera………………………………………………………………160

Figure-64 Relationship between Log Ash-Free-Dry-Weight (AFDW) and Log Shell

Length (SL) in Nerita senegalensis……………………………………….161

XXV

Figure-65 Shell variables relationships shell length Vs shell height for

Nerita senegalensis………………………………………………………..162

Figure-66 Shell variable relationships for shell length Vs shell height for

Nerita senegalensis………………………………………………………………….163

Figure-67 Shell variable relationships for shell length Vs shell width for

Nerita senegalensis…………………………………………………………..164

Figure-68 Shell variable relationships for shell length Vs aperture width in

Nerita senegalensis………………………………………………………..165

Figure-69 Shell variable relationships for shell length Vs aperture width for

Nerita senegalensis……………………………………………………….166

Figure-70 Mean Percentage of different algae and detritus found in the gut content of

Limpet species……………………………………………………………...168

Figure-71 Box-plot showing the minimum, mean and maximum percentage of gut

contents of limpet species………………………………………………….169

Figure- 72 Mean monthly increment in Shell Length (SL) in cohort of

Patella safiana……………………………………………..……………….172

Figure-73 Mean monthly increment in Shell Length (SL) in cohort of

Nerita senegalensis...... 173

XXVI

Figure-74 Variations in Shell Length (SL) increment in Tank 1 Patella safiana and

Tank 2 (Patella safiana+ Nerita senegalensis)………………………….174

Figure-75 Mean shell length increment in population of Patella safiana………….176

Figure-76 Mean wet weight increment in population of Patella safiana…………….177

Figure-77 Temporal variations GSI values for male and female of Patella safiana….189

Figure-78 Temporal variations in GSI values for male and female of

Nerita senegalensis………………………………………………………194

LIST OF PLATES

CONTENTS PAGES

Plate-1 West mole of Tarkwa Bay…………….……………………………………….34

Plate-2 Training mole of Tarkwa Bay………………………………………………….36

Plate-3 East mole of Tarkwa Bay………………………………………………………38

Plate-4 Station 1 at the West mole of Tarkwa Bay……………………………………..41

Plate-5 Station 2 at the West mole of Tarkwa Bay……………………………………41

Plate-6 Station 3 at the West mole of Tarkwa Bay……………………………………...41

XXVII

Plate-7 Station 4 at the West mole of Tarkwa Bay…………………………………….. 41

Plate-8 Station 1 at the Training mole of Tarkwa Bay… ……………………………...42

Plate-9 Station 2 at the Training mole of Tarkwa Bay………………………………….42

Plate-10 Station 3 at the Training mole of Tarkwa Bay………………………………….42

Plate-11 Station 4 at the Training mole of Tarkwa Bay………………………………….42

Plate-12 Station 1 at the East mole of Tarkwa Bay………………………………………43

Plate-13 Station 2 at the East mole of Tarkwa Bay……………………………………….43

Plate-14 Station 3 at the East mole of Tarkwa Bay…….………………………………. ..43

Plate-15 Station 4 at the East mole of Tarkwa Bay………………………………………43

Plate-16 Study species (A= Thais callifera, B = Patella safiana,

C = Nerita senegalensis and D = Buccinum undatum……………………..….. 45

LIST OF APPENDICES

CONTENTS PAGES

Appendix 1: Physico-chemical parameters investigated at the West mole of Tarkwa

Bay (April, 2008-March, 2010)…………………………………………….246

Appendix 2: Physico-chemical parameters investigated at the Training mole of Tarkwa Bay

(April, 2008-March, 2010)………..……………………………………….247

Appendix 3: Physico-chemical parameters investigated at the East mole XXVIII

of Tarkwa Bay (April, 2008 – March, 2010………………………………..248

Appendix 4: Descriptive statistics for number of individuals collected in dry seasons

at the three moles of Tarkwa Bay (April, 2008 – March, 2010)…………249

Appendix 5: Temporal variations in euryhaline molluscan species sampled at

the three moles of Tarkwa Bay (April,2008-March, 2010)………………251

Appendix 6: Temporal variations in dry season molluscan species sampled at

the three moles of Tarkwa Bay(April, 2008-March, 2010)...... 252

Appendix 7: Temporal variations in rainy season molluscan species sampled at

the three moles of Tarkwa Bay (April, 2008- March, 2010)………..…..253

Appendix 8: Checklist of common molluscan species of the three moles

of Tarkwa Bay (April,2008–March,2010)……………………………..254

Appendix 9: Checklist of rare species of molluscan communities of Tarkwa Bay

(April, 2008 – March, 2010)……………………………………………..255

Appendix 10: Checklist of molluscan ornamental species of the three moles

of Tarkwa Bay ( April 2008- March, 2010)…………………………..256

Appendix 11: Checklist of species of aquaculture potentials sampled at the three

moles of Tarkwa Bay (April, 2008 – March, 2010)…………………….257

Appendix 12: Checklist of bio-fouling species of the molluscan communities of

XXIX

Tarkwa Bay (April, 2008-March, 2010 )………………………………258

Appendix 13: Descriptive statistics for the number of individuals collected at the

three moles of Tarkwa Bay during the dry season

(2008-March, 2010) ……………………………………………………...259

Appendix 14: Descriptive statistics for the number of individuals collected at the

three moles of Tarkwa Bay during the rainy season

(April, 2008- March, 2010)………………………………………………261

Appendix 15: Temporal variations in total abundance, compositions, distributions of

molluscan faunal species the three tidal zone among the three

moles of Tarkwa Bay (April, 2008 – March, 2010)……………………263

Appendix 16: Mean values of male and female monthly gonad dry weight (mg) of

three species of herbivorous gastropods investigated from( May,2008

-December, 2009)………………………………………………………….264

Appendix 17: Temporal variations in Species Richness (d) computed for the communities

of molluscan fauna sampled at the three moles of Tarkwa Bay

(April, 2008 – March, 2010)………………………………………………265

Appendix 18: Temporal variations in Shannon – Weaver Indices computed for the

communities of the molluscan fauna investigated at the three

XXX

moles of Tarkwa Bay ( April, 2008 – March, 2010……………………..266

Appendix 19: Temporal variations in Simpson’s Dominance Index (C) computed for

the communities of the molluscan fauna at the three moles

of Tarkwa Bay (April, 2008 – March, 2010 )…………………………….267

Appendix 20: Temporal variations in values of Evenness Indices computed for the

communities of the molluscan fauna at the three moles

of Tarkwa Bay (April,2008- March ,2010 )……………………………268

Appendix 21: Temporal variations in density ( Total number of Individuals/m2) of

different species of molluscan fauna collected at the 3 moles

of Tarkwa Bay ( April,2008- March, 2010)……………………………..269

Appendix 22: Temporal variations in number of individuals’ molluscan fauna

sampled at the three tidal levels at the three moles of Tarkwa Bay

(April, 2008 – March, 2010)……………………………………………..270

Appendix 23: Temporal variations in the number of species ( S ) computed for the

communities of molluscan fauna sampled at the three moles

of Tarkwa Bay ( April,2008 – March, 2010 )………………………………272

Appendix 24: Studies on the growth of Patella safiana in the natural and the laboratory

environments investigated (May, 2008- July, 2009) on the East mole of

XXXI

Tarkwa Bay………………………………………………………………..273

Appendix 25: Temporal variations of heavy metal (µg/g/dry weight) concentrations

- in different body parts of selected molluscan faunal species at

the west mole, investigated in dry season (April, 2008-March, 2010)….274

Appendix-26 Temporal variations of heavy metal concentration (µg/g dry weight)

in different body parts of selected molluscan species in the West

mole investigated in rainy season (April,2008- March, 2009)………275

Appendix-27 Mean concentrations of heavy metals in seawaters, seaweeds and selected

molluscan species investigated at the three mole of

Tarkwa Bay in Dry season ( April, 2008- March, 2010),………………276

Appendix-28 Mean concentrations of heavy metals in seawaters, seaweeds and selected

molluscan species investigated at the three moles of

Tarkwa Bay in dry season (April, 2008- March, 2010)………………..277.

Appendix-29 Mean concentration of heavy metals in seawater, seaweeds and

selected molluscan species investigated in rainy season at the three moles

of Tarkwa Bay…………………………………………………………278.

XXXII

1.6 Operational definition of terms:

• Anthropogenic - Human induced environmental degradation conditions of a community

or ecosystem.

• Artificial breakwaters - are heterogeneous rocky assemblages built to abate

erosion along the coastline.

• Baseline - Historical data taken to assess the current conditions of a community or

an ecosystem.

• Bay: Part of the sea within a wide curve of the shore.

• Dominant species- Organism that exerts a controlling influence on or define the character of a community.

• Fauna: community found in one or more regions.

• Foraging- Act of acquiring food for optima energy expenditure.

• Intertidal -Alternately flooded and exposed by tides.

• Quadrat- A rectangular or squared-shaped instruments used to estimate density cover and biomass of both plant and .

• Reference site -A minimal impaired site that is representative of the expected ecological conditions and integrity of other of the same type and region.

• Regime – A regular pattern of occurrence or action.

XXXIII

1.7 Lists of Abbreviations

WM West Mole (West artificial breakwaters).

TM Training Mole

EM East Mole

ITCZ Inter-Tropical Convergence Zone

GPS Global Positioning System

S. R Sex Ratio

GSI Gonad Somatic Index

R E Reproductive Effort

GW Somatic Weight

SST Sea Surface Temperature

SSS Sea Surface Salinity

AAS Atomic Absorption Spectrophotometer

MFS Millipore Filtering System

XXXIV

Abstract

The molluscan community living in the three moles of Tarkwa Bay, Lagos was studied monthly between April 2008 and March 2010. These moles are made of rock known as the west, training and east. They provide attachment surfaces for the settlement of littoral communities. The study investigated the physico-chemical parameters, nutrients, heavy metals concentration levels in sea surface water, algae and resident molluscs of the three moles. Molluscan diversity, feeding, guts content analysis; growth and reproduction of selected species were assessed. Water samples were collected monthly at low tides from twelve geo-referenced network of sampling stations established on the littoral shorelines of the study sites. Sea surface water was collected just 0.5 metre below the surface using 2.01 non-metallic Hydrobios water sampler, kept in pre-labelled polyethylene 500ml bottles, and later transported to the laboratory for storage in deep freezer at 40C for further analyses. Algal samples were scraped from rock surfaces at low tides, and washed 3 times in seawaters and kept in pre- labelled 1000ml plastic containers for storage in the laboratory. Molluscs were collected at both subtidal and intertidal using quadrats of inert aluminium frame of size 1 m2 were used at low tides along transects fixed landward-seaward directions at 12 study sites. A total of 8486 individuals belonging to 69 species, 34 families and 4 classes were recorded.

The four classes of Phylum encountered were: 89.92%; Bivalvia

9.55%; Cephalopoda 0.3%; and Polyplacophora 0.2%. A total of 5794 individuals constituting 68.3% of the total population, were collected during the dry seasons, compared to 2692 individuals (31.70%) collected during the rainy season. The One - way ANOVA results showed that growth in Patella safiana and Nerita senegalensis were significantly difference among the three tidal zones on the shore (P < 0.05). The annual mean percentage of Chlorophyta in the gut of Patella safiana was 34.3 ± 7.51% with a range of 1.8% (July) to

XXXV

44% (May). Rhodophyta had a minimum percentage gut content of 4% (April), reaching a maximum of 31% (July) with a mean percentage gut content of 14.58± 8.43%. Detritus accounted for a minimum of 17% (March) and a maximum of 32% (November) with annual mean value of 23.7 ± 4.31%. The mean value for the level of metal concentrations decreased in the order: Mn > Fe > Zn > Cu > Cr > Pb; in seawater, algae and molluscs. The level of heavy metal concentrations in the body parts of molluscs decreased in the order: Body tissues > foot > mantle > shell. The highest level of metal concentration was recorded in

Buccinum undatum (266.6µg/g dry wt.) for Mn and lowest concentration was found in Nerita

Senegalensis (0.01µg/g dry wt. of Pb).The assessment of Gonadosomatic Index (GSI) in male and female showed similar temporal patterns (t 0.05, 38= 2.02), however, but there was significance different among months (ANOVA; P < 0.05) for reproductive cycles of Patella safiana and Nerita senegalensis. The annual reproductive efforts for patella safiana (0.7048) male and female (0.5360), while Nerita senegalensis male (3.2436) and female (3.0190). The study generated a comprehensive checklist of molluscan communities of Tarkwa Bay which will be useful for future assessments and ecotourism development. Information on water quality, dietary preference and heavy metal concentration will be handy for shellfish industries. The established seasonal influence and reproductive patterns in selected molluscs, which will serve as valuable tools for breeding and conservation of intertidal molluscs in maritime communities in Nigeria.

XXXVI