From the Arabian Sea Off Oman

BIOLOGY, STOCK ASSESSMENT AND FISHERY MANAGEMENT OF THE SANTER SEABREAM CHEIMERIUS NUFAR (VAL. 1830) FROM THE ARABIAN SEA OFF OMAN

Dissertation Submitted By

Abdulaziz Said Mohamed Al-Marzuqi

In partial fulfilment of the requirements for the degree of Doctor of Natural Sciences (Dr. rer. nat.)

Leibniz Center for Tropical Marine Ecology

Faculty of Biology / Chemistry University of Bremen

February 2011 Zusammenfassung Im Mittelpunkt der Dissertation steht die Biologie und die fischereibezogene Einschätzung von Santer seabream, Cheimerius nufar (Valenciennes, 1830), entlang der Küste des Arabischen Meeres des Sultanates Oman in den Jahren 2005-2008. Zusätzlich wurden die saisonbedingte Verteilung bestimmter Umweltparameter im Arabischen Meer und ihr Einfluss auf den Bestand von C. nufar untersucht. Diese Untersuchungen basieren auf Einschätzungen der Grundfischerei durch das Forschungsschiff AL Mustaqila 1 in dem Zeitraum September 2007 – September 2008. Vier Untersuchungsgebiete wurden erfasst. Wichtige Ergebnisse der Studie bezüglich C. nufar werden weiter unten zusammengefasst. Die Oberflächentemperaturen lagen zwischen 25°C und 28°C in der Herbst- Zwischenmonsunperiode (Sep – Dez), fielen dann nach der NE-Monsunperiode (Feb) auf 22°C – 24°C und stiegen dann erneut in der Frühjahrs- Zwischenmonsunperiode (Apr. – Jun.) auf 25°C – 28°C an. Die Oberflächentemperatur war generell gering (18°C – 24°C) unmittelbar nach dem SW-Monsun. In Küstennähe (20-50 m Schicht) war die Bodentemperatur mit 25°C am höchsten während der Herbst-Zwischenmonsunperiode (Sep-Okt) im Jahre 2007 und am geringsten (17°C) unmittelbar nach dem SW-Monsun im Jahre 2008.

Der Seeoberflächensalzgehalt schwankte wenig während der Untersuchungen und lag im Bereich zwischen 36 und 37 ppt. Einen geringeren Salzgehalt (34ppt) wiesen im Jahre 2008 lediglich die Gewässer südlich der Insel Masirah unmittelbar nach der SW-Monsunperiode auf. Kleine Taschen mit einem höheren Salzgehalt wurden im Gebiet 3 des Schelfs beobachtet. Der Bodensalzgehalt zeigte ein Bild ähnlich dem Oberflächensalzgehalt.

Der Gehalt an O2 im Oberflächenwasser war in allen Gebieten während der Untersuchungen hoch (4-6 ml l-1). Die Sauerstoffminimumzonen (OMZs) waren die pelagischen Zonen mit geringem Sauerstoffgehalt die sich nach unten erstreckten direkt von der Mischschicht bis zu einer Tiefe von mehr als 1000 m in

I einigen Gebieten. Der Bodensauerstoffgehalt war gering in den meisten der Untersuchungsgebiete während der Frühjahrs-Zwischenmonsunperiode (Apr-Jun).

Es gab Unterschiede im Vorkommen von C. nufar während der Untersuchungen die in der Periode nach dem Südwestmonsun 2007 und 2008 durchgeführt wurden. Der Fisch trat auf bei Temperaturen zwischen 20°C und 24°C während des NE- Monsuns und bei Temperaturen zwischen 26°C-28°C während des Frühjahrs- Zwischenmonsuns. Während des Post-Südwest-Monsuns wurde der Fisch bei Temperaturen zwischen 17°C und 21°C angetroffen. Der Fisch C. nufar scheint an ein geringes Sauerstoffniveau während der verschiedenen Untersuchungsperioden angepasst zu sein.

Der geschätzte Fang an C. nufar während der Jahre 2005-2006 bzw. 2006-2007 betrug 2075t bzw. 2018t.

Die Größe der kommerziellen Anlandungen an C. nufar erstreckt sich von 16 cm TL bis 64 cm TL.

Etwa 67.5% des Fanges bestand aus Fisch mit einer Länge unter 32 cm TL.

Da die Längen-Gewichts-Beziehung von männlichen und weiblichen Exemplaren sich nicht signifikant unterscheidet, kann die übliche Gleichung W=0.019L2.897 für diese Fischart benutzt werden.

Diese Fischart ist ein Fischfresser, der sich gelegentlich von cephalopoden, crustacaens und polychaetes ernährt.

Leere Mägen waren üblich in all den Monaten und Reifestadien des Fisches.

II Es wurden Inkonsistenzen bezüglich der Intensität der Nahrungsaufnahme des Fisches in den verschiedenen Stadien der Reife bei beiden Geschlechtern festgestellt.

Die mittlere Erstlaichlänge der weiblichen Fische (31.9 cm TL) war geringer als die der männlichen (33.7 cm TL).

Die Laichreife tritt bei beiden Geschlechtern zu Beginn des dritten Jahres ein.

Die Population von C. nufar laicht während der Monate Mai – Oktober. Einzelne Fische scheinen zweimal im Jahr zu laichen.

Die Fertilität erstreckt sich von 11406 – 473277 Eier.

Es wurde keine Dominanz einer der Geschlechter in der Population beobachtet. Ein Test des monatlichen und jährlichen Geschlechterverhältnisses mittels Chi- Square-Test ergab keinen signifikanten Unterschied.

Folgende VBG-Parameter wurden für C. nufar ermittelt: basierend auf Längenfrequenzanalysen L∞ = 69,9 cm, K = 0.29 y-1 und t0 = -0,32 y und basierend auf Otholithenanalysen für männliche Fische L∞ =66,47 cm, K = 0,1937 y-1 und t0 = -0,36055 y und für weibliche Fische L∞ = 69,77 cm K = 0,182 y-1 und t0 = -0,24886 y. Das maximale Alter basierend auf Längenfrequenzen wurde mit etwa 11 Jahren geschätzt; der Otholith gibt jedoch eine Lebensdauer von etwa 13 Jahren an. Der Wachtumsindex (Ø) von C. nufar erreichte in diesen Untersuchungen basierend auf Längendaten einen Wert von 3,14 und basierend auf Othlithenanalysen einen Wert von 2,95. Hieraus folgt, dass diese Fischart vergleichsweise in den Gewässern Omans schnell wächst.

Die ermittelte Fanglänge (Lc) betrug 31,4 cm. 67,4% der kommerziell angelandeten Fische wies eine Länge <34 cm auf.

III Die ermittelten Koeffizienten totale Sterblichkeit (Z), natürliche Sterblichkeit (M) und Fischereisterblichkeit (F) wiesen die Werte 1,3, 0,62 und 0,68 auf.

Das ermittelte M/K – Verhältnis weist den Wert 2,13 auf.

Die Ausbeutungsrate (E) für die Untersuchungsperiode ist gleich 0,52.

Der geschätzte MSY für den Bestand an C. nufar betrug bei Anwendung der Formel von Cadima 1953 t und bei Anwendung des Thompson und Bell Models 1946 t.

Das durchschnittliche beständige Aufkommen wurde auf 3010 t geschätzt.

Der mittlere totale Bestand (Y/U) wird auf 5387 t geschätzt.

Die vorhergesagte totale Biomasse von 16052 t verringert sich auf 12713 t bei einem Wert F=0,1, auf 6167 t bei einem Wert von F=0,5 und auf 5235 t bei einem Wert von F=0,6. Bei einem Wert von F=2,0 betragen der Ertrag und die Biomasse 18 t bzw. 57 t.

Bei dem gegenwärtigen Wert von F (=0,68) erreichen die Beziehungen Yw/R, TB/R, SSB/R und FB/R die Werte 111 g, 260 g, 191 g und 163 g.

Die vorliege Studie zeigt das der Bestand an C. nufar an der Küste Omans des Arabischen Meeres geringfügig überausgebeutet ist. Die mittlere Größe des zu fangenden Fisches sollte auf 36 cm TL begrenzt werden. Weiterhin ist es zur Erreichung eines nachhaltigen dauerhaften Ertrages zu empfehlen das derzeitige Aufwandsniveau um 17% zu senken.

IV Summary

This thesis focuses on the biology and fishery assessment of the santer seabream, Cheimerius nufar (Valenciennes, 1830) along the Arabian Sea coast of the Sultanate of Oman during 2005–2008. In addition the seasonal distribution of certain environmental parameters in the Arabian Sea and their influence on the stock of C. nufar were studied based on the demersal fishery resources survey conducted by the R.V. Al Mustaqila 1 during Sept 2007 – Sept 2008 covering four survey areas.

The Important findings of the study on C. nufar are summarized below.

Sea surface temperature tended to be warmest that ranged between 25 ºC and 28 ºC during the autumn intermonsoon period (Sept–Dec), cooled after the NE monsoon period (Feb) to 22 ºC – 24 ºC, warmed again to 25 ºC – 28 ºC during the spring intermonsoon period (Apr–Jun), and was generally very cool (18 ºC - 24 ºC) immediately after the SW monsoon. Nearshore (20–50 m strata) bottom temperatures w ere highest 25 ºC during the autumn intermonsoon period (Sept- Oct) in 2007 and were coolest (17 ºC) immediately after the SW monsoon in 2008. Sea surface salinity varied little throughout the surveys that ranged between 36 and 37 ppt except for immediately after the SW monsoon period in 2008 when waters south of Masirah Island were less saline (34 ppt), although small pockets of higher saline waters were seen in area 3 on the shelf. Bottom salinity showed a pattern similar to surface salinity. -1 Dissolved O2 in the surface water was high (4–6 ml l ) in all areas during all thesurveys. The OMZs were the midwater zones of low oxygen that extend from just below the mixed layer to more than 1000 m in some areas. Bottom oxygen was low in most of the survey areas during the spring intermonsoon period (April–June).

V There was difference in the distribution pattern of C. nufar between the surveys done in the post southwest monsoon periods of 2007 and 2008. The fish occurred at temperatures between 20ºC and 24ºC during NE monsoon and spring inter-monsoon and at 26ºC-28ºC during autumn inter-monsoon. During post-southwest monsoon of 2008 the fish was found at temperatures between 17ºC and 21ºC. The fish C. nufar appeared to adapt to low oxygen level during different periods of survey. The catches of C. nufar were estimated at 2,075 t and 2018 t during 2005-2006 and 2006-2007 respectively.

The size of C. nufar in the commercial landings ranged from 16 cm TL to 64 cm TL. About 67.5% of the catch composed of fish measuring less than 32 cm TL.

As the length-weight relationships of males and females did not differ significantly, the common equation W = 0.019L2.897 can be used for the species.

The species is a piscivore feeding occasionally on cephalopods, crustaceans and polychaetes. Empty stomachs were common in all the months and maturity stages of fish.

There was inconsistency in feeding intensity of fish in different stages of maturity in both the sexes

The average length at first maturity in female (31.9 cm TL) was lower than in male (33.7 cm TL).

The maturity in both males and females appeared to attain at the beginning of third year.

The population of C. nufar spawned during May – October and individual fish appeared to spawn twice a year.

VI Fecundity of ranged from 11,406 - 473,277 of eggs.

There was no dominance of either sex in the population and the monthly and annual sex-ratios tested by chi-square test did not differ significantly.

The calculated VBG parameters of C. nufar based on length frequency analysis were: L∞ = 69.9 cm, K = 0.29 y-1 and t0 = -0.32 y. and based on otolith study were : for the male, L∞ = 66.47 cm, K = 0.1939 y-1 and t0 = -0.36055 y and for female, L∞ = 69.77 cm, K = 0.182 y-1 and t0 = -0.24886 y. The estimated maximum age based on length frequency would be around 11 years; however, otolith indicated the longevity of about 13 years. The growth performance index (Ø’) of C. nufar in the present study was estimated at 3.14 based on length data and 2.95 based on otolith analyses suggested the species is comparatively growing fast in the Omani waters. The calculated length at capture (Lc) was 31.4 cm and about 67.4% of the fish in the commercial catches measured < 34 cm.

The estimated total mortality (Z), natural mortality (M) and fishing mortality (F) coefficients were 1.3, 0.62 and 0.68 respectively.

The M/K ratio estimated for the species was 2.13.

The exploitation rate (E) for the period of study stood at 0.52.

The estimated MSY for the stock of C. nufar by Cadima’s formula was 1,953 t and by Thompson and Bell model was 1,946 t.

The average standing stock was estimated as 3,010 t.

The estimated average total stock (Y/U) stood at 5,387 t.

The predicted total biomass of 16,052 t decreased to 12,713 t at F=0.1, to 6,167 t at F= 0.5 and to 5,235 t at F= 0.6. At F= 2.0, the yield and biomass stood at 18 t and 57 t respectively

VII At the current F (= 0.68), the values of Yw/R, TB/R, SSB/R and FB/R were found to be 111 g, 260 g, 191 g and 163 g respectively.

The present study indicates the stock of C. nufar in the Arabian Sea off Oman is marginally overexploited. The average size of the fish to be caught may be restricted to 36 cm TL. Further, a reduction of 17% of effort from the present level of effort is recommended for obtaining sustainable yield.

VIII Acknowledgements

First of all my thanks to the God, who created me, helps me and has given me the health and ability to reach this stage. I would like to express my thorough respect and appreciation to my supervisor, Prof. Dr. Ulrich Saint-Paul, who freely discussed my ideas for this study, for his helpful guidance, advice and valuable comments. I extend my thanks to my co-supervisor Dr.Werner Ekau for his incessant support throughout my study period. Special thanks go to my local supervisor Prof. Dr. Nachiappan Jayabalan for his valuable advice and comments. I appreciated his quality and friendly supervision.

I would like to thanks my Government, Sultanate of Oman, represented by Ministry of

Fisheries Wealth for offering me this scholarship.

I would like to thanks my colleagues in the Marine Science and Fisheries centre for their support and the invaluable assistance they provide in getting the needed information for this study.

Finally, I would like to record my sincere appreciation of the love, sacrifices and unlimited help and support of my family who encouraged me to study and overcome all obstacles and who always pray and worship God for my success.

IX Table of Contents

Summary………………………………………………………………………………….I

Acknowledgement……………………………………………………………………....IX

Table of Contents………………………………………………………………………..X

List of Figures………………………………………………………………………....XII

List of Tables…………………………………………………………………………XVI

1. INTRODUCTION…………………………………………………………………....1 1.1. Literature review…………………………………………………………………..1 1.2. The Study Area……………………………………………………………………4 1.3. Aims of the study………………………………………………………………….7 2. MATERIALS AND METHODS...... …...8 2.1. Sampling procedure of the Arabian Sea Survey ………………………………….8 2.1.1. Resource Assessment Survey of the Arabian Sea Coast of Oman…………..8 2.1.2. Survey area, design, stratification and station allocation…………………….8 2.1.3. Survey procedure and data collection………………………………………10 2.2. Collection and analysis of C. nufar samples from landing sites………………...12 2.2.1. Age and growth estimation…………………………………………………12 2.2.2. Maturation and spawning…………………………………………………...15 2.2.3. Food composition and feeding intensity……………………………………18 2.2.4. Population dynamics and stock assessment………………………………...19

3. RESULTS……………………………………………………………………………22 3.1. Seasonal distribution of physico-chemical characteristics of the water of the Arabian Sea………………………………………………………………………22 3.2. Study of C. nufar………………………………………………………………...37 3.2.1 Age and growth estimation……………………………………………………..39 3.2.2. Maturation and spawning………………………………………………………54 3.2.3. Food composition and feeding intensity……………………………………….72 3.2.4. Population dynamics and stock assessment……………………………………79

X 3.3. Distribution of C.nufar in relation to oceanography parameters…………………82

4. DISCUSSION……….………………………………………………………………..87 5. REFERENCES……………………………………………………………………….97

XI List of Figures

Fig. 1. Sultanate of Oman geographical location……………………………………5

Fig. 2. Survey area and stations in the Arabian Sea…………………………………10

Fig. 3. Post- SW monsoon 2007: Sea surface temperature…………………………..22

Fig. 4. Autumn inter- monsoon 2007: Sea surface temperature……………………..23

Fig. 5. NE monsoon 2008: Sea surface temperature…………………………………23

Fig. 6. Spring Inter- monsoon 2008: Sea surface temperature……………………….24

Fig. 7. Post- SW monsoon 2008: Sea surface temperature…………………………..24

Fig. 8. Post- SW monsoon 2007: Bottom temperature………………………………25

Fig. 9. Autumn Inter-monsoon 2007: Bottom temperature………………………….26

Fig. 10. NE monsoon 2008: Bottom temperature……………………………………26

Fig. 11. Spring Inter-monsoon 2008: Bottom temperature…………………………..27

Fig. 12. Post- SW monsoon 2008: Bottom temperature……………………………..27

Fig. 13. Post- SW monsoon 2007: Surface salinity………………………………….28

Fig. 14. Autumn monsoon 2007: Surface salinity…………………………………...29

Fig. 15. NE monsoon 2008: Surface salinity………………………………………...29

Fig. 16. Spring Inter-monsoon 2008: Surface salinity…………………………….…30

Fig. 17. Post- SW monsoon 2008: Surface salinity…………………………………30

Fig. 18. Post- SW monsoon 2007: Bottom salinity………………………………….31

Fig. 19. Autumn monsoon 2007: Bottom salinity…………………………………...32

Fig. 20. NE monsoon 2008: Bottom salinity………………………………………...32

Fig. 21. Spring Inter-monsoon 2008: Bottom salinity……………………………….33

XII Fig. 22. Post- SW monsoon 2008: Bottom salinity………………………………..….33

Fig. 23. Post- SW monsoon 2007: Bottom dissolved oxygen………………..……….34

Fig. 24. NE monsoon 2008: Bottom dissolved oxygen…………………………....….35

Fig. 25. Spring inter-monsoon 2008: Bottom dissolved oxygen……………………...36

Fig. 26. Post- SW monsoon 2008: Bottom dissolved oxygen………………………...36

Fig. 27. C. nufar distribution from all trawl survey stations……………………….…38

Fig. 28. Biomass of C. nufar in different depth zones in the Arabian Sea survey ………...…38

Fig. 29. Length frequency distribution of C. nufar in the commercial catches during

2005-06………………………………………………………………………39

Fig. 30. Length frequency distribution of C. nufar in the commercial catches during

2006-2007……………………………………………………………………40

Fig. 31. Pooled length frequency distribution of C. nufar in the commercial catches

during 2005-2007...... 40

Fig. 32. Total length-fork length relationship in C. nufar …………………………....41

Fig. 33. Length-weight relationship in C. nufar……………………………………....42

Fig. 34. VBGF curve of C. nufar by SLCA technique………………………………..42

Fig. 35. VBGF curve of C. nufar by PROJMAT technique………………………..…43

Fig. 36. VBGF curve of C. nufar by ELEFAN 1 technique…………………………..43

Fig. 37. Age distribution of C. nufar in the commercial catches during 2005-2007….45

Fig. 38. Growth curves derived from length frequency analyses in C. nufar…………46

Fig. 39. Length at first capture of C. nufar…………………………………………...46

Fig. 40. Photomicrograph of a transverse section through the sagittal otolith………..47

XIII Fig. 41. Monthly percentage of otoliths with hyaline and opaque edge………………48

Fig. 42. VBGF curves for C. nufar, Female (n =159) and Male (n =99)……………..49

Fig. 43 A. Development of ova to maturity stage I in C. nufar……………………….55

Fig. 43 B. Development of ova to maturity stage II in C. nufar………………………55

Fig. 43 C. Development of ova to maturity stage III in C. nufar……………………..56

Fig. 43 D. Development of ova to maturity stage IV in C. nufar……………………..56

Fig. 43 F. Development of ova to maturity stage V in C. nufar………………………56

Fig. 44. Distribution of different maturity stages of gonads of C. nufar during 2005-06………………………………………………………………………………..58

Fig. 45. Distribution of different stages of gonads of C. nufar during 2006-07………59

Fig. 46 - A. Fish with running ripe maturity stage during post SW monsoon 2007…..60

Fig. 46 - B. Fish with running ripe maturity stage during post SW monsoon 2008…..60

Fig. 47. Monthly gonado-somatic indices in C. nufar 2005 – 2007…………………..61

Fig. 48. Monthly hepato-somatic indices in C. nufar 2005 -2007………………….....62

Fig. 49. Monthly Kn in C. nufar 2005-2007…………………………………………..63

Fig. 50.Juveniles distribution -Post SW monsoon 2007……………………………....63

Fig. 51 Juveniles distribution: Autumn monsoon 2007……………………………….64

Fig. 52 Juveniles distribution –NE monsoon 2008…………………………………....64

Fig. 53 Juveniles distribution Spring Inter-monsoon 2008……………………………65

Fig. 54 Juveniles distribution -Post SW monsoon 2008……………………………....65

Fig. 55 Length at first maturity in C. nufar…………………………………………...66

Fig. 56 A. Fecundity in relation to total length (cm)………………………………….67

Fig. 56 B. Fecundity in relation to body weight(g)…………………………………...68

XIV Fig. 56 C. Fecundity in relation to ovary weight(g) of C. nufar……………………..68

Fig. 57. Sex ratio of C. nufar during the period April 2005 - March 2007………...... 69

Fig. 58. General food composition of C. nufar during 2005-06………………………72

Fig. 59. General food composition of C. nufar during 2006-07………………………73

Fig. 60. Monthly percentage of composition of food items of C. nufar during 2005-06 …………………………………………………………………...... 74

Fig. 61. Monthly composition of food items of C. nufar during 2006-07…………….74

Fig. 62. Food items found in the gut of various size groups of C. nufar during 2005-2007 (pooled)………………………………………………………....75

Fig. 63. Feeding intensity in C. nufar relation to months……………………………..76

Fig. 64. Feeding intensity in different maturity stages of C. nufar……………………77

Fig. 65. Feeding intensity in different size groups of C. nufar………………………..78

Fig. 66. Predicted yield for C. nufar for different F…………………………………..80

Fig. 67. Equilibrium yield-per-recruit and biomasses-per-recruit in C. nufar

against F…………………………………………………………………...... 81

Fig.68. Catch of C. nufar at different time of monsoon……………….82

Fig. 69 Distribution of C. nufar: Post SW monsoon 2007…………………………....83

Fig. 70 Distribution of C. nufar : Autumn monsoon 2007……………………………83

Fig. 71 Distribution of C. nufar: NE monsoon 2008………………………………….84

Fig. 72 Distribution of C. nufar: Spring Inter-monsoon 2008………………………...84

Fig. 73 Distribution of C. nufar: Post SW monsoon 2008…………………………...85

Fig.74 Catch rate of C. nufar by stratum during monsoon periods…………………...86

Fig.75. Occurrence of C. nufar at different level of bottom dissolved oxygen……...... 86

XV List of Tables

Table 1. Details of survey period, demersal fish survey and CTD stations …………..9

Table 2 Estimated biomass (t) and coefficients of variation for C. nufar……………....37

Table 3. Length – weight relationship parameters ……………………………………..41

Table 4. VBGF parameters of C. nufar estimated by different methods ………………44

Table 5. Growth parameters of male and female C. nufar using otolith reading……….49

Table 6 A. Age - length key for Male of C. nufar - sectioned otolith. …………………50

Table 6 B. Age - length key for female of C. nufar - with sectioned otolith. ………….51

Table 6 C. Age structure of C. nufar population in the Arabian Sea…………………...52

Table 6 D. Age structure of C. nufar population in the Arabian Sea…………………...53

Table 7A Sex-ratio in C. nufar during 2005-06………….………………………………70

Table 7B Sex-ratio in C. nufar during 2006 -07…………………………………………71

Table 8. Comparison of growth parameters of C. nufar from different studies………....90

XVI 1. Introduction

This thesis focuses on the fishery assessment of the santer seabream, Cheimerius nufar along the Arabian Sea coast of the Sultanate of Oman during 2005–2008 taking into consideration of the influence of the environmental parameters. T h e study was conducted in two phases. While the phase-1 study was related to the biology and stock assessment of C. nufar based on samples collected from the commercial fish catches, phase-2 investigation was concerned with the distribution of certain environmental parameters in the Arabian Sea and their influence on the distribution of C. nufar based on the fishery resource survey conducted by the R.V. Al Mustaqila 1. The work is situated in the context of an ongoing requirement for thefisheries sector of the country to manage the resource sustainably.

1.1 Literature review

The fisheries sector has always been important to Oman as a significant source of animal protein and employment. Prior to the discovery of oil and its subsequent exploitation in the late 1960s, the fishing industry supported about 80% of the population, and even today, around 50% of Omanis rely upon fishing as the source of income retaining its prominence within the limits of renewable economic resources (MoI, 2 0 0 8 ).

Though, there is general increase in the total annual catch of fish in recent years, the landings of quality fish have declined due to overexploitation (MAF, 2006). As in most developing countries, Oman’s coastal fisheries are supported by multispecies, more than 150 species of fishes and crustaceans (Al-Abdessalaam, 1995) and harvested by multigear. The number of fishermen and the gears employed increase from year to year and at present about 34,757 fishermen are engaged in fishing using 13,862 boats (GOSO, 2009). As the consequence, there is concomitant reduction in the prosperity of coastal fisherfolk who depend upon fishing for their livelihood. The reason for carrying out this study was due to the serious concerns repeatedly raised o n the sustainability of demersal stocks and the welfare of the traditional fishers despite long coastline and frequent reports of large number of foreign vessels trawling in the closed areas of the Omani waters.

1 Prior to 1990s, a decline in total catch and reduction in the a v e r a g e size of the individuals caught were recorded from the Omani waters; however, no specific reason for the decline could be attributed (Siddeek, 1999). Besides, there has been a shift from pelagic to demersal fishing in Oman as some of the higher value pelagic species like the kingfish Scomberomorus commerson were depleted (Siddeek et al., 1998).

The sparid C. nufar is locally known as ‘kofwar’ or ‘frenka’ inhabits inshore waters up to the depth of 60-100 m and forms an important component of the demersal fishery of Oman targeted by both the traditional fishers using handlines and gillnets and by industrial trawlers (Abdessalaam, 1995). Due to its high demand in the market, increased fishing pressure has lead to overexploitation of the stock (Abdessalaam, 1995). The estimated biomass and potential yield from the stocks of all the sparids of Oman were reported to be around 40,000 t and 6,000 t respectively (Abdessalaam, 1995). The estimated catches of C. nufar from the Arabian Sea coast of Oman for the years 2005-06 and 2006-07 were 2,075 t and 2,018 t respectively (GOSO, 2005; 2006; 2007). While the contribution of all sparids to the annual total catch of Oman was about 4.2% with a value of about US$ 11.95 million during 2007; the catches of C. nufar represented about 45% of the sparid catches from Arabian Sea with an estimated value of about US$ 5.38 million (GOSO, 2007).

Fishes of the family Sparidae are represented by nineteen species in the Omani waters (Fouda et al., 1997) and many of them contribute to the traditional and industrial fisheries of the country (GOSO, 2007). The santer seabream, C. nufar is a moderate sized fish and is distributed in the western Indian Ocean including the Red Sea, coasts of Oman, Arabian Gulf, eastern African coast, Madagascar and the Mascarene Islands (Bauchot and Smith, 1984; Smith and Smith, 1986; Randall, 1995; Connell et al., 1999). Individuals of C. nufar may attain a total length of 75 cm and weigh about 6 kg; however, most of the fish caught are in the weight range of 1 to 3kg (Connell et al., 1999). Detailed studies on the biology and population structure on C. nufar are available from the Gulf of Aden (Druzhinin, 1975; Edwards et al., 1985.), South African coasts (Coetzee and Baird, 1981; Coetzee, 1983,

2 Garratt, 1985, 1985a, 1991; Smale, 1986; Buxton and Garratt, 1990) and Mozambique (Timochin, 1992).

Though, C. nufar is a priority species and intensively fished in Oman, except the report on its spawning and reproductive pattern (McIlwain et al., 2006), no other study on fishery biological characteristics of the species is available. Presently, an attempt has been made to study the biological and population parameters of C. nufar from the Arabian Sea harvested by artisanal gears for two years from April 2005 to March 2007. Subsequently, the distribution pattern of the environmental parameters in the Arabian Sea exclusive economic zone (EEZ) of Oman and their influence on the distribution and abundance of C. nufar as well as spawning time of the species were investigated to determine if there were consistent patterns that could be related to the physico-chemical parameter variability of water through the survey cruises conducted by the R.V. Mustaqila 1 during September 2007 – September 2008.

Many events take place in the reproductive life of fishes with clear periodicity, including within-year, within-spawning season, within-lunar cycle, and within-day patterns. The timing of events at each of these scales differs from species to species and from location to location. For e x a m p l e , many species of reef fishes congregate at reef passes to spawn during high tide (Shibuno, et al., 1993; Hensley, et al., 1994; Domeier and Colin, 1997). It is a strategy of the fish to direct spawning to a time when eggs and larvae are carried rapidly away from the reef and its dependent high density plankton feeding fish (Shibuno, et al., 1993; Hensley, et al., 1994; Danilowicz, 1995). Ecological factors such as predation, competition or food availability can influence the timing of spawning (Johannes, 1978; Norcross and Shaw, 1984). Alternatively, spawning time may be dictated by physiology for synchronization in the population (Qasim, 1973).

During particular time of the year, t h e best conditions l i k e temperature and salinity trigger the adults to build up body reserves that can be used subsequently to elaborate mature gonads.Spawning in fish coincides with environmental conditions suitable for survival and growth of eggs and larvae or juveniles (Bye, 1984; Norcross and Shaw, 1984). Hence, even minor increase in the

3 water temperature due to global warming might impose constraints on the fish spawning time. It is hoped that all the present study will help to understand the status of the stock of C. nufar along the Omani coast to plan for sustainable harvest.

1.2 The study Area: Arabian Sea and Oman coast

The Sultanate of Oman is located in the North West Indian Ocean and is surrounded by three seas, the Arabian Gulf and the Oman Sea in the north, and the Arabian Sea in the south (Fig.1). Fisheries productivity is heavily influenced by the oceanographic dynamics in the area. The region is dominated by the seasonal monsoon winds, particularly the southwest monsoon, which generates upwelling of nutrient rich bottom waters to the surface that enhance productivity, particularly in the northern part of the area. The other major feature likely to strongly influence fish productivity and distribution is the seasonal development of oxygen deficient layers in depths between 50 and 1000 m in which most fish species cannot live(Wishner et al., 1998).

The area for this study was located in the Arabian Sea between the latitude 16º 33' N and 22º 21' N and longitude between 53º 09' and 59º 55' E. This area is unique tropical basin in the world which has a semi-annual cycle (Bohm et al.,1999;Kumaret al.,2001a;Welleret al., 2002) and the strongest annual variability in water circulation (Esenkov et al., 2003).

4 Fig. 1. Sultanate of Oman geographical location

The Arabian Sea is one of the most complex marine regions of the world (Richardson et al., 2006) and is influenced by seasonal variations in conditions (Savidge et al., 1990).There are two different water masses in the study area, which are characterized by strong upwelling during the SW monsoon (May to September) accompanied by cloud cover. These conditions have been confirmed by various investigations (Schott, 1983; Quraishee, 1984; Kindle, 2001; Prasad, 2004). During upwelling w h i c h is more intense in July, sea water temperature is reduced (Rao et al., 2005). The SW monsoon later reverses to the

5 north-easterly direction with moderate winds and clear skies from November to February (Flagg and Kim, 1998). During the SW monsoon strong currents and upwelling bring colder, less saline and nutrient rich bottom water from about 500 m depth to the surface mixed layer to compensate the loss of water in the upper 500 m layer (Shi et al., 1999). These features are also capable of transporting nutrient rich cold water to hundreds of kilometers offshore (Brink et. al., 1998). The effect of the monsoon on the water temperature is well documented (Wishner et al. 1998; Lee et al., 2000, Weller et al., 2002). Warm sea surface temperatures are typically seen offshore during the SW monsoon period, with cooler, more recently upwelled water present near-to-shore.

The oxygen minimum zone (OMZ) is a midwater zone of low oxygen that extends from just below the mixed layer to more than 1000 m, in some areas with oxygen -1 concentration less than 0.1 ml l (Morrison et al., 1999). Species survival might be significantly influenced by the fluctuations of dissolved oxygen. That impacts the species composition in the ecosystem followed by trophic pathways and productivity changes (Ekau et al., 2010).

Occurrence of the OMZs in the study area is common (Morrison et al., 1995). The reasons for the existence of the OMZs may be due to large scale local ox ygen consumption rates from high productivity in surface waters, very slow movement of decaying organic matter within the layer consuming most oxygen present, low oxygen concentration in the waters entering the layer from the southern ocean (Olson et al., 1993), or the lack of an opening to seas to the north (Morrison et al., 1999).

While, small organisms are physiologically limited by dissolved oxygen levels less -1 than 0.1 ml l (Morrison et al.1999), larger organisms, such as fish, are limited by -1 slightly higher levels around 0.3 ml l (Ashjian et al., 2002). However, demersal fish and crustaceans (motile organisms) were rarely found where oxygen

6 -1 concentration was below 1.5 ml l in “The Dead Zone” in the Gulf of Mexico (Rabalais et al., 2002).

Fluctuations in the extent of the OMZs can lead to redistribution of populations of species (John, 2004), reduced distribution (less habitat available) (Arntz et al., 1988), reduced catch (Banse, 1984), shifting of demersal fishes into the pelagic zone, direct mortality, forced migration, increased susceptibility to predation, changes in food resources and disruption of life cycles (Rabalais et al., 2002). The OMZs also serve as biogeographic barriers by limiting cross- slope movements of populations (Rogers, 2000).

1.3.Aims of the study o To understand the seasonal variation of the marine environmental parameters in the Arabian Sea and their influence on the distribution of C. nufar. o To study the biological and population characteristics of C. nufar in the Arabian Sea o To find out whether there is any consistent spatial difference in the timing of spawning of C. nufar in relation to environmental factors o To support the assessment and management of the stock(s) of C. nufar targeted commercially in the Arabian Sea.

7 2. Material and Methods

2.1 Sampling procedures for the Arabian Sea Survey

2.1.1. Resource Assessment Survey of the Arabian Sea Coast of Oman

During 2007 – 2008 the Ministry of Fisheries Wealth (MFW) Sultanate of Oman conducted a survey to provide estimates of the fishable biomass of principal demersal, small pelagic and mesopelagic fish resources off the Arabian Sea coast of Oman for ongoing stock assessment of Omani fisheries, to guide development and investment decisions. Five seasonal surveys with an average duration of 47 days were completed in the project period using the RV Al Mustaqila 1. The surveys were timed to ensure coverage of the main seasons, with an overlap of one season, between August 2007 and September 2008 (Table 1). The specific timing of the surveys was designed to achieve a reasonable seasonal coverage but to avoid the limitations on survey activity that were likely, based on normal weather patterns, during the June-July part of the SW monsoon period. As the period of one cruise overlapped with two subsequent season, for presentation of oceanographic data, the seasons broadly clasiified as shown in Table -1 were followed.

2.1.2. Survey area, design, stratification and station allocation

The defined area for the survey was the continental shelf in the 20–250 m depth range from Ra’s al Hadd to the Yemen border southwest of Salalah (Fig.2). The region of survey was divided into 4 major areas, the boundaries of which were based on features where the shelf edge is at its narrowest. A two phase stratified random survey design was adopted in order to achieve greatest precision in surveys for mixed demersal species; fisheries resources (Francis 1981,1984). The major 4 survey areas were further divided by depth to make 15 strata. As the surveys progressed, the survey area was recalculated including areas of strata. Initial stratum boundaries were defined from electronic copies of British Admiralty charts.

8 The depth contours considered from these charts were 20–50, 50–100, and, in some areas, the 100–150 m depth ranges.

Phase 1 of each seasonal survey consisted of 100 random trawl stations allocated to strata based on an average station density of about 1 station per 190 km2 of trawlable area, or 1 station per 317 square km of the total area. A minimum of three stations per stratum was sampled in phase 1 of each survey. Up to 20 phase 2 stations were then directed to the strata with the most variable catch rates. Station positions were selected randomly before each voyage with a minimum distance between stations of 2 nautical miles (Fig. 2).

Trawl survey tows were carried out during daylight hours only. Care was taken to ensure that trawls started after dawn and were completed by dusk to minimize variations in catchability which might result from day/night variation in fish reaction to the trawl. The target numbers of demersal trawl stations per survey are given (Table 1).

Table 1. Details of survey period, demersal fish survey and CTD stations

Voyage Voyage period Season demersal CTD No. stations station Post SW 215 1 17 Sep 2007 - 15 Oct 101 monsoon Autumn Intermonsoon 253 2 1 Nov - 17 Dec 2007 104

3 29 Jan 2008 - 18 Mar NE monsoon 120 348 4 19 Apr - 10 Jun Spring Intermonsoon 123 319 5 Post SW monsoon 1 Aug - 23 Sep 2008 122 256

Total 570 1391

9 Fig. 2. Survey area and stations in the Arabian Sea

2.1.3. Survey procedures and data collection

The R.V. Al Mustaqila 1 is 46 m length overall, has a beam of 12.5 m, horsepower of 3602 and a displacement of 1745 tonnes. The trawl gear was rigged and deployed in the same manner for all the demersal tows across the 5 surveys. The 40 m cod-end selected for the demersal trawl survey was constructed of 4 mm knotted nylon braid. The nominal inside mesh measurement was 40 mm. A series of 3 x 20 meshes were measured using callipers at the beginning of the first survey, when the cod-end was dry (i.e., had not been used). The cod-end was marked and repeat measurements were made during the second survey to evaluate any variation in mesh sizes. At each station, the trawl target distance was 2 nautical

10 miles at a speed over the ground of 3.5 knots. Towing speed and gear configuration were maintained as constant as possible during all seasonal surveys. The trawl doors were of Thyboron Type 7 capable of being used in midwater or on the seafloor. The door weight and size were 1968 kg and 9.37 m2 respectively. Measurements of door spread, headline height, and vessel speed were recorded every 5 min during each tow.

Beside the fishery resources, data on physio-chemical parameters of the water in the Arabian Sea were collected during the survey. Temperature, salinity, dissolved oxygen and depth were recorded by CTD deployment after every trawl operation. The maximum amount of line deployed on the CTD casts was 500 m. To measure 100% dissolved oxygen (DO) saturation, the CTD was deployed just below the water surface on each cast for at least 1 minute or held in a holding tank with constant surface water flow for at least 5 minutes at each station. Two different modules of CTDs were used during the survey, XR-620 and XR-420. The first module replaced the X R - 4 2 0 on t h e f i r s t cruise, when it was found t h a t the XR-420 titanium casing caused temperature lag. The XR-420 was used after station 142 on t h e t h i r d c r u i s e because the XR-620 failed.

A biological data collection program on C. nufar was undertaken during the survey based on special request for the purpose of this study. These data include fish weight recorded to the nearest 0.1 g, sex, fish total length measured to the nearest 0.1 cm. The stages of gonadal maturity identified are given in Annex 1.

11 2.2. Collection and analysis of C. nufar samples from landing sites

Random samples of C. nufar were collected from commercial catches landed by different types of artisanal gears such as gillnets, handlines, traps and commercial trawl nets on monthly basis between April 2005 and March 2007.

2.2.1. Age and growth

A total of 4132 fish were measured for their total length (TL) and fork-length (FL) to the nearest 1 mm using a fish measuring board and rounded off to the nearest cm to calculate the monthly size frequency distribution in the catches. The relationship between the TL and FL was estimated by regression analysis.

For the estimation of length-weight relationships of C. nufar, the TL and total weight (TW) of the fresh fish were used. While, the TL was measured to the nearest 1 mm, the TW was recorded to the nearest 1 g using a top pan balance. The general equation,

= where, TW and TL are the total weight and total length respectively, and 'a' and 'b' are the constants to be determined were fitted separately for males and females.

Analysis of covariance (ANCOVA) (Snedecor and Cochran, 1967) technique was employed to test the significant difference if any, in the relationships between males and females at 5% level.

2.2.1.1. Estimation of growth parameters using length – frequency data

A total of 4132 fish measured for their TL and the lengths were grouped into 2 cm class intervals. The size range of the fish varied from 16 cm to 64 cm TL. To estimate the von Bertalanffy growth function (VBGF) parameters (L∞, K and t0) the equation,

12 =∗1−∗( °) was fitted using the LFDA version 5.0 of FMSP- Fish Stock Assessment Software (Hoggarth et al., 2006). Three alternative fitting techniques such as Shepherd's Length Composition Analysis (SLCA), Projection Matrix (PROJMAT) and ELEFAN 1 with non-seasonal version were used to calculate VBGF parameters. Also, the routine Powell-Wetherall technique available with the FMSP-software was used to estimate L∞ and Z/K. Since the suitability of ELEFAN 1 technique is suggested for tropical fish (Pitcher, 2000), the growth parameters estimated by ELEFAN 1 technique were considered for further analysis of stock assessment. Length at age was estimated from the plots of SLCA, PROJMAT and ELEFAN 1, and by the empirical method of Froese and Binohlan (2000). Length at first capture was estimated by plotting cumulative percentages of length against length classes.

2.2.1.2. Estimation of growth parameters using otolith

A total of 258 fish (159 females and 99 males) ranging in size from 16 to 66cm (TL) was used for otolith based ageing. To estimate the mass of the otoliths, one of the sagittae from each fish was weighed to the nearest 0.01g. To determine the fish age, the internal structure of the otolith, including the primordium (core) was examined in resin-embedded sections.

One otolith from each fish was embedded in small moulds (< 2cm) containing epoxy resin (Brothers et al., 1983). Thin sections (300μm) of otoliths were made through the nucleus along a transverse, dorsoventral plane with a Buehler Isomet low-speed saw containing a diamond wafering blade. In order to have a polished face for viewing, excess resin on the sectioned face of the otolith was removed using a grinding wheel fitted with silicon carbide paper of different grit sizes (400 to 1200 grit) and flushed with water.

Each section of the otolith was mounted on a microscopic slide and observed at 10x X 4x magnification using a compound microscope with transmitted light. All the sectioned otoliths (n=258) displayed clear increments and rings were used to determine the age of the fish. The term “opaque zone” refers to the area that

13 appeared milky under reflected light or dark under transmitted light in unstained samples. In our study, terminology was based on that recommended by Morales-Nin (1991).

The age of each fish was determined by counting the number of rings; i.e. pairs of alternating opaque and translucent rings from the sectioned otoliths. Rings were counted, without reference to fish length or date of capture, in the ventral lobe of the sagittal otolith from the primordium to the otolith margin close to the ventral margin of the Sulcus acusticus. All counts were made in the same way and repeated at least three times, with a two-week interval between readings, to ensure consistency. For each fish the third count was used to estimate age, since by this time considerable experience had been gained in the interpretation of the rings and otolith structure. The relationship between size and age was determined using the von Bertalanffy Growth Function (VBGF) (von Bertalanffy, 1934). The outer edge of the otoliths was examined on a monthly basis for the presence of either an opaque or hyaline zone to establish whether these zones were deposited annually. The growth parameters estimated were compared with earlier studies. Since, growth is not linear, comparison of growth in two fish populations using L∞ and K may be misleading. Hence, the overall growth performance index (Munro’s phi prime index, Ø’) for C. nufar was calculated empirically (Munro and Pauly, 1983) using the formula,

(∅′) = log + 2 log ∞ where, K is expressed on annual basis and L∞ in cm.

14 2.2.2. Maturation and Spawning

For studying the maturation and spawning in C. nufar, a total of 1,329 specimens collected during 2005-07 were used.

State of maturity of gonads was recognized by modifying the stages defined by the International Council for Exploration of the Sea (Lovern and Wood, 1937) to six stages of maturity key (I-Immature; II-Maturing 1; III-Maturing 2; IV-Mature; V- Ripe/Running and VI-Spent) to suite the present study. The stages were recognized both in females and males by examining the ovaries macroscopically and also by the microscopic structure of the intraovarian eggs; whereas, in males macroscopic structure of testes alone was considered (Annex 1). The fixed ovarian stages were counter checked with the histological sections (Annex 1B).

To study the development of ova from immature to mature stage, 36 ovaries in maturity stages I to VI were used. Samples of ova from anterior, middle and posterior regions of both the lobes were examined. As there was no difference in the size of ova from different parts of the ovary, ova were sampled from the middle region of the ovary for ova diameter measurements. About 200 ova from each ovary were measured for the diameters. The method of measurement of ova diameters was similar to that of Clark (1934); Prabhu (1956) and Jayabalan (1986). Ova diameters were measured using an ocular micrometer fitted to the eye piece of the microscope and each ocular micrometer division (o.m.d) was equal to 14μ (1000 μ=1mm). As the ova measuring up to 6 o.m.d. were present in large numbers in all the stages of ovary indicating the general stock of eggs, they were not measured from stage II onwards. Ova diameter data from ovaries of same stage of maturity were pooled and were grouped into 3 o.m.d for plotting percentage frequency curves.

The spawning season was determined based on commercial catch and samples collected during the R.V. Al Mustaqila 1 demersal trawl survey. For spawning season based on commercial catch, monthly percentage occurrence of different maturity stages of gonads, gonado-somatic index (GSI), hepato-somatic index (HSI) and relative condition factor (Kn) were calculated and plotted. A comparison of

15 spawning of C. nufar was also made using the data on maturity stages of gonads during various surveys.

The monthly gonado-somatic index (GSI) was calculated separately for males and females adopting the formula,

= × 100

where, GW denotes the gonad weight (g) and GuW denotes gutted weight (g).

For the nutritional state of the fish, the monthly hepato-somatic index was calculated as suggested by Busacker et al., (1990) using the formula,

= / × 100

where HW = the liver weight (g) and TW = the total weight (g) of fish.

The monthly relative condition factor, Kn was estimated to understand the condition of the fish following Le Cren (1951) as,

=/ where TW = observed weight, aLb = calculated weight obtained from the length- weight relationship.

For estimating length at first maturity (Lm), fish were grouped into 1 cm class interval and gonads from stage-III onwards were considered as mature for calculating the fraction of mature fish at different lengths. A logistic function was fitted to the fraction of mature fish against length interval as suggested by King (1995).

The logistic equation used was:

=1/ 1+ − ( − ) where P = proportion mature fish in length class L,

r = the width of the maturity curve and

16 Lm = length at 50% maturity

Based on the batches of intraovarian eggs that formed modes in the advanced stages of ovaries (mature, ripe and spent ovaries), the spawning frequency in individual fish was determined (Zacharia and Jayabalan, 2007). Fecundity of females were estimated gravimetrically from 27 ovaries of stage IV. The minimum and maximum sizes of the fish used for estimation of fecundity ranged between 28 cm TL and 63.4 cm TL and the body weight ranged from 280 g to 3,730 g. The weight of the ovaries used varied from 3.5 g to 185.91 g respectively. From the known weight of the ovary preserved in 5% neutral formalin, a small portion of the ovary was removed and weighed to the nearest 0.001g in an electronic microbalance and then kept in 'modified Gilson's fluid' (Simpson, 1951) for 4 days. The piece of the ovary was taken to a counting chamber and all the mature eggs in the piece of the ovary were counted under a binocular microscope. From the number of mature ova in the piece of the ovary, the total number of mature ova was estimated using the formula,

Fecundity =Total weight of the ovary / Weight of the sample * Number of mature ova in the sample

The relationship between the fecundity and variables like total length, body weight and ovary weight was found out by the least square method,

= where, F=fecundity, a = constant, X = variable (fish length, fish weight or ovary weight) and b = correlation coefficient.

The sex-ratio was estimated as male to female ratio. To test the monthly and annual sex-ratios for the expected ratio of 1:1, the technique of chi-square (X2) analysis was

∑()2 used. =

Where, O = observed number of males and females in each month

E = expected number of males and females in each month

17 2.2.3. Food composition and feeding intensity

For analysis of quantitative and qualitative estimation of food of fish, samples were collected at random from the commercial catch. The gut content from 775 specimen during 2005-06 and 550 fish during 2006-07 were analysed. However, some bias might have been introduced while sampling the fish for food and feeding habits as the fish are targeted by handlines and trap fishery using baits, besides gillnets and trawl nets.

For the general food items found in the gut, they were identified up to group level. When the food item was in advanced state of digestion, it was recorded as ‘semidigested matter’. For the seasonal variation of the quantity of food items, points (volumetric) method (Hynes, 1950) was used.

Feeding intensity in C. nufar was studied from September 2005 to March 2007. A total of 990 fish were analyzed for the fullness of the stomachs and were grouped into actively fed (full and 3/4 full stomachs), moderately fed (1/2 full stomachs), poorly fed (1/4 full stomachs) and empty. The feeding intensity in relation to months, maturity stages and size of the fish was estimated.

18 2.2.4. Population dynamics and stock assessment

The instantaneous total mortality rate (Z) of C. nufar was estimated by the length- converted catch curve method (Pauly, 1983) and Beverton-Holt method (Beverton and Holt, 1956) using the routines provided in the LFDA version 5.0 of FMSP software. Adopting the Powell-Wetherall technique, L∞ and Z/K of C. nufar were also calculated.

Natural mortality coefficient (M) of C. nufar was estimated by three different empirical methods (Rikhter and Efanov 1976; Pauly, 1980; Alagaraja 1984) and the average M was taken for further analyses.

In the empirical method of Pauly (1980), the following equation was adopted.

ln = −0.0152 − 0.279 ∗ ln ∞ + 0.6543 log∗ + 0.4634 ∗ ln where ‘T’ indicates the annual mean temperature (oC) of the surrounding water in which the fish lives. In the present study, the T value was taken as 24°C as this value represented the mean bottom water temperature in the Arabian Sea off Oman (Thangaraja, 1995).

In Rikhter and Efanov's method (1976), the following formula was adopted

= 1.521/ 0.72 − 0.155 where, tm = the age at which 50% of the population matures.

Alagaraja's (1984) method assumes that 99% of a cohort had died, if it had been exposed to natural mortality only. The formula used was:

1% = − ln(0.01)/

Where Tm indicates longevity and M1% indicates natural mortality corresponding to 1% survival.

19 Fishing mortality coefficient (F) was obtained by subtracting natural mortality (M) from total mortality (Z) as,

=−

The term standing stock refers to the concentration of fish population for a given area at a given time. This can be estimated in terms of numbers or weight. In this study, the standing stock was estimated by weight using the formula,

=/ where Y is the yield and F, the fishing mortality

The total stock or biomass in weight was estimated using the relation between yield and exploitation ratio as

=/ where Y is the annual yield and U is the exploitation ratio.

The maximum sustainable yield (MSY) was calculated by the equation suggested by Cadima (in Traodec, 1977). Cadima's estimator used was,

=0.5( + ) where Y is the total catch in a year, is the average biomass in the same year and M the natural mortality. The MSY was also estimated using the ‘Yield’ software (Hoggarth et al., 2006).

The exploitation rate (E) of C. nufar was estimated as suggested by Sparre and Venema (1992) using the formula,

=/, where, E= the exploitation rate; F= the fishing mortality and Z= total mortality. The exploitation ratio was calculated by the equation given by Ricker (1975) as,

=/(1− ) ,

20 where, U= the exploitation ratio; F= the fishing mortality and Z= total mortality

Per-recruit analyses were made using routine available with the 'Yield' package incorporated in the FMSP Software (Hoggarth et al., 2006). Estimations of equilibrium yield-per-recruit (Yw/R), total biomass-per-recruit (TB/R) and stock spawning biomass-per-recruit (SSB/R) were made for a range of F-values of C. nufar. All the input parameters used for “Yield” software were obtained from the present study.

The optimum length of exploitation was estimated empirically from the equation (Froese and Binohlan, 2000) as,

=3∗ ∞/3 + /

Where Lopt denotes the optimum length of exploitation, L, the asymptotic length, M, the natural mortality and K, the growth coefficient.

21 3. Result

3.1. Seasonal distribution of physico-chemical characteristics of waters of the Arabian Sea

Temperature Generally the sea surface water temperature was found to be warm that ranged between 25 and 28 ºC during the autumn inter-monsoon period (Sept– Dec) that cooled after the NE monsoon period (Feb) to 22 ºC – 24 ºC and warmed again to 25 ºC – 28 ºC during the spring inter-monsoon period (Apr–Jun). The water became very cool (18-24 ºC) immediately after the SW monsoon (Figs. 3-7). In the Gulf of Masirah area, the surface sea water was comparatively cooler by several degrees (up to 5˚C) than in the waters in the southwest after the SW monsoon period (Sept) in 2007 (Fig. 3). However, during the autumn inter-monsoon period (Nov–Dec), the sea surface water temperature did not fluctuate very much in the entire Arabian Sea area (Fig. 4). Sea surface water temperatures within Salalah area were comparatively warmer than in other areas during spring inter-monsoon 2008 (Fig. 6).

Fig. 3. Post- SW monsoon 2007: Sea surface temperature

22 Fig. 4. Autumn inter- monsoon 2007: Sea surface temperature

Fig. 5. NE monsoon 2008: Sea surface temperature

23 Fig. 6. Spring Inter- monsoon 2008: Sea surface temperature

Fig. 7. Post- SW monsoon 2008: Sea surface temperature

24 The nearshore (20–50 m strata) bottom temperatures were highest during the autumn inter-monsoon period 25 ºC that subsequently reduced gradually as the surveys progressed (Figs. 8–14). Bottom water temperatures in the entire area surveyed in the Arabian Sea area were at their lowest 17 ºC immediately after the SW monsoon (Fig. 12). During 2008, the survey in the post-SW monsoon period was conducted nearly two months earlier than in 2007. This may be the reason for the higher water temperatures recorded during the post-SW monsoon period in 2007 (Fig. 8). Along the shelf edge in the entire survey area, the bottom water temperatures were consistently low.

Fig. 8. Post- SW monsoon 2007: Bottom temperature

25 Fig. 9. Autumn Inter-monsoon 2007: Bottom temperature

Fig. 10. NE monsoon 2008: Bottom temperature

26 Fig. 11. Spring Inter-monsoon 2008: Bottom temperature

Fig. 12. Post- SW monsoon 2008: Bottom temperature

27 Salinity

The sea surface water salinity did not vary much throughout the surveys (Figs. 13–16) that ranged between 36 and 37 ppt. However, immediately after the SW monsoon period in 2008 drastic changes occurred, 34 ppt, (Fig. 17). The salinity values of the waters south of Masirah Island immediately after the SW monsoon in 2008 were low (34-35 ppt); although, in certain areas on the shelf more saline waters occurred. The reason for variability in sea surface salinity after the SW monsoon in 2008 might be the effect of upwelling that brought colder, less saline waters upon to the shelf. The sea surface water salinity in the area south of Masirah Bay had reverted back to the average salinity of approximately 36 ppt during post SW monsoon 2007 (Fig. 13).

Fig. 13. Post- SW monsoon 2007: Surface salinity

28 Fig. 14. Autumn monsoon 2007: Surface salinity

Fig. 15. NE monsoon 2008: Surface salinity

29 Fig. 16. Spring Inter-monsoon 2008: Surface salinity

Fig. 17. Post- SW monsoon 2008: Surface salinity

30 The bottom water salinity showed similar pattern to surface salinity (Figs. 18–22). The occurrence of less saline water, (33–35 ppt), was evident immediately after the SW monsoon period in entire shelf south of Masirah Island (Fig. 22). During SW monsoon, the bottom waters in the southern stratum (Areas 3 and 4) became more saline, 36 ppt, (Fig. 18). The bottom water salinity was almost uniform throughout the survey area by autumn monsoon 2007 (Fig.19).

Fig. 18. Post- SW monsoon 2007: Bottom salinity

31 Fig. 19. Autumn monsoon 2007: Bottom salinity

Fig. 20. NE monsoon 2008: Bottom salinity

32 Fig. 21. Spring Inter-monsoon 2008: Bottom salinity

Fig. 22. Post- SW monsoon 2008: Bottom salinity

33 Dissolved oxygen -1 Generally, the dissolved oxygen was high (4–6 ml l ) throughout the survey area. Though, the oxygen minimum zone (OMZ) was recorded in the nearshore waters in much of the survey area during the post SW monsoon period 2007; layers of well- oxygenated waters were present in certain pockets located in Sawqirah Bay at depths of 20–100m (Fig. 23). During this period as the oxygen sensor failed, measurements of dissolved oxygen could not be carried out for the southern half of the survey area.

After the NE monsoon period (Jan–Mar), the extent of the OMZ got altered (Fig. 24). The OMZ was not present in shallower waters, but was shifted to near the shelf edge and close to the Oman– Yemen border. The OMZ was present much deeper, at depths of 150-200 m.

Fig. 23. Post SW monsoon 2007: Bottom dissolved oxygen

34 Fig. 24. NE monsoon 2008: Bottom dissolved oxygen

During the spring inter-monsoon period (April–June), the concentration of dissolved -1 oxygen was low in the bottom waters, 0.1 – 1 ml l almost in the entire survey area (Fig. 25). In certain areas close to the shore and in south of Masirah Island, high oxygenated bottom waters existed. Though, the well oxygenated nearshore bottom waters occurred south of Masirah Island soon after the SW monsoon period, in certain -1 pockets of shallow areas (40-50 m) on the shelf, oxygen poor waters (up to 0.3 ml l ) were present (Fig. 26).

35 Fig. 25. Spring inter-monsoon 2008: Bottom dissolved oxygen

Fig. 26. Post- SW monsoon 2008: Bottom dissolved oxygen

36 3.2. Study of C. nufar

Biomass and distribution of C. nufar in the Arabian Sea

Cheimerius nufar occurred throughout the survey area, but more commonly to the south of Masirah Island, and predominantly in waters shallower than 100 m depth (Fig. 27). Total survey biomass (tonnes) of C. nufar and coefficients of variation (CVs) are given Table 2. Estimates range from 1321 t in voyage 5 to 4598 t in voyage 3. Lakbi bay had the highest estimates of biomass during voyages 1, 2, 3 and 4; while, Salalah area estimates were highest on voyage 2 and 3. The 20–50 m depth range held the highest biomass for all voyages; and the 100 - 150 m depth range produced lowest biomass during voyages 1, 2 and 5 (Fig. 28).

Table. 2 Estimated biomass (t) and coefficients of variation (%, below in parentheses) for C .nufar.

Voyage 1 Voyage 2 Voyage 3 Voyage 4 Voyage 5 Average

1513 3022 4598 2251 1321 2,541 t

(33) (31) (35) (17) (27)

37 Fig. 27. C. nufar distribution from all trawl survey stations

2000

1750

1500

1250 20 – 50 m

1000 50 – 100 m 100 – 150 m Biomass ( t) 750 150 – 250 m 500

250

0 Voyage 1 Voyage 2 Voyage 3 Voyage 4 Voyage 5

Fig. 28. Biomass of C. nufar in different depth zones in the Arabian Sea survey

38 3.2.1. Age and growth estimation

In the present study, the estimations of age and growth of C. nufar were made using two different techniques, the length frequency analysis and the interpretation of growth marks on otoliths

Length frequency distribution

The size frequency distribution of C. nufar in the commercial catches ranged from 16 cm to 64 cm of TL during 2005-06 and during 2006-07, the size was ranged between 18 cm and 64 cm of TL. During 2005-06, fish measuring up to 26 cm formed about 32% of the catches. However, individuals in the size range of 28 cm - 30 cm formed about 33% of the catches and the rest of the higher sized fishes collectively formed about 35% of the catches (Fig. 29). This trend changed to 26%, 23% and 51% respectively during 2006-07 (Fig. 30). The size frequency of the pooled data for both the years is given in Fig. 31.

N=1996 14 12 10 8

% 6 4 2 0 14 18 22 26 30 34 38 42 46 50 54 58 62 TL(cm)

Fig. 29. Length frequency distribution of C. nufar in the commercial catches during 2005-06

39 N=2136 14 12 10 8

% 6 4 2 0 14 18 22 26 30 34 38 42 46 50 54 58 62 TL(cm)

Fig. 30. Length frequency distribution of C. nufar in the commercial catches during 2006-2007

N=4132 14 12 10 8

% 6 4 2 0 14 18 22 26 30 34 38 42 46 50 54 58 62 TL(cm)

Fig. 31. Pooled length frequency distribution of C. nufar in the commercial catches during 2005-2007

40 Total length-Fork length relationship

The relationship established between the total length and the fork length of the fish can be expressed as, FL = 0.892TL-0.151 (R2=0.991) (Fig. 32)

60 FL = 0.892TL - 0.151 50 R² = 0.991 N= 1,300 40

FL (cm) 30

10 15 25 35 45 55 65 75 TL (cm)

Fig. 32. Total length-fork length relationship in C. nufar

Length-weight relationship

The estimated length-weight relationships of males, females and sexes pooled data of C. nufar are shown in Table 3.

Table 3. Length – weight relationship parameters

Sex LW Confidence interval Sa Sb W = 0.018*L 2.907 a: -0.033 – 0.071 0.02650 0.01736 Male (R2 = 0.986) b: 2.872 – 2.940 W = 0.020*L2.890 a: - 0.028 – 0.068 0.02439 0.01595 Female

(R2 = 0.985) b: 2.859 – 2.921 W = 0.019*L2.897 a: -0.016 – 0.055 0.01795 0.01174 Pooled (R2 = 0.985) b: 2.874- 2.920

41 The ANCOVA test for length-weight relationships of males and females showed both the slopes and means were not significantly different (P>0.05) and hence, the common equation for the species would fit well (Fig. 33). The coefficient of determination (R2) in males, females and sexes pooled could be highly correlated.

4000

W = 0.019L2.897 3000 R² = 0.985

2000

Total weight (g) 1000

0 10 20 30 40 50 60 70 Total length (cm)

Fig. 33. Length-weight relationship in C. nufar.

Estimation of growth parameters using length frequency data

Figures 34 – 36 show the VBGF parameters of C. nufar estimated using the monthly length frequency data for the period 2005-2007 by SLCA (Fig. 34), PROJMAT (Fig. 35) and ELEFAN (Fig. 36) techniques.

Growth Curve = Non Seasonal. Linf = 70.35. K = 0.31. Tzero = -0.23.

Length Class Length 20

0 0 0.5 1.0 1.5 2.0 2.5 Sample Timing

Fig. 34. VBGF curve of C. nufar by SLCA technique

42 Growth Curve = Non Seasonal. Linf = 68.26. K = 0.25. Tzero = -0.55.

Length Class Length 20

0 0 0.5 1.0 1.5 2.0 2.5 Sample Timing

Fig. 35. VBGF curve of C. nufar by PROJMAT technique

Growth Curve = Non Seasonal. Linf = 70.97. K = 0.30. Tzero = -0.18.

Length Class Length

0 0 0.5 1.0 1.5 2.0 2.5 Sample Timing

Fig. 36. VBGF curve of C. nufar by ELEFAN 1 technique

43 The L∞ of C. nufar estimated by Powell-Wetherall technique was 65.27 cm. Though there were variations in VBGF parameter values between different techniques (Table 4), the values obtained by ELEFAN 1 technique were used for subsequent stock assessment.

Table.4. VBGF parameters of C. nufar estimated by different methods

-1 Method L∞ (cm) K (y ) t0 (y)

SLCA 70.35 0.31 -0.229

PROJMAT 68.26 0.25 -0.55

ELEFAN 1 70.97 0.298 -0.180

Powell-Wetherall 65.27 - -

Age distribution in commercial catches

The age distribution of C. nufar in the commercial catches (Fig. 37) indicated that individuals belonging to the age groups 0-4 years supported the fishery.

44 Age Slice

Nominal Age Nominal

0 0 0.5 1.0 1.5 2.0 2.5 Sample Timing

Fig. 37. Age distribution of C. nufar in the commercial catches during 2005- 2007

Length at age

The lengths of C. nufar were converted into age using growth estimations by the SLCA, PROJMAT and ELEFAN 1 techniques (Fig. 38). The SLCA indicated higher lengths for all the ages followed by the ELEFAN 1 and PROJMAT techniques. The lengths converted into age showed that the life span of C. nufar would be about 11 years in the Omani waters. The life span estimated by the equation 3/K (Pauly, 1983a) indicated that the fish may live up to 10.5 years.

45 75

SLCA 45 PROJMAT ELEFAN

Total(cm) length 35 Average

15 1234567891011 Age (y)

Fig. 38. Growth curves derived from length frequency analyses in C. nufar

Length at capture and recruitment

In the commercial catches, the smallest fish encountered measured 16 cm of TL and the largest fish measured 64 cm TL. The length at 50% of fish caught (Lc) was calculated to be 31.4 cm (Fig. 39). The maturity in both males and females appeared to attain at the beginning of third year.

100 90 80 70 60 Lc=31.4 cm 50 40 30

Probablity of capture 20 10 0

15 20 25 30 35 40 45 50 55 60 65 70 TL(cm)

Fig. 39. Length at first capture of C. nufar

46 Estimation of growth parameters using otoliths

Periodicity of ring formation

A section through the otolith of a 13 year old C nufar is shown in Fig. 40. The distance between the growth rings can be seen to narrow with increasing age.

Fig. 40. Photomicrograph of a transverse section through the sagittal otolith.

A total number of 258 sectioned otoliths were read. The rings were clearly visible and the age of the fish could be determined in most of them. A definite seasonal variation in the formation of opaque and hyaline zones is evident on the otolith edge (Fig.41). Hyaline zone formation takes place from March – June and the opaque zones are laid down from July – January. It can therefore be concluded that one opaque zone and one hyaline zone are formed annually.

47 100

% 50 Opaque 40 Hyaline 30

0 JFMAMJJASOND Months

Fig. 41. Monthly percentage of otoliths with hyaline and opaque edge.

Size at age and growth

The VBGF curves were fitted to lengths-at-age for C. nufar (Table 5, Fig. 42). The age estimation indicates the life span is 13 years. The VBGF suggests that the male fish had a higher growth rate when younger and attained a smaller theoretical maximum size than females. In the age – length key, a wide range of lengths (24 – 64 cm) within most age groups was found (Table 6 A, B, C & D).

48 Table 5. Growth parameters of male and female C. nufar using otolith reading.

2 Sex L∞ Kto r

Male 66.47 0.1939 -0.36055 0.89652

Female 69.77 0.182 -0.24886 0.92528

Male - Data Size (cm) 30 Female - Data 20 M - VBG

10 F-VBG

0 02468101214 Age(y)

Fig. 42. VBGF curves for C. nufar, Female (n =159) and Male (n =99).

49 Table 6 A. Age - length key for Male of C. nufar - sectioned otolith.

Age (y) TL(cm) Total 1 2 3 4 5 6 7 8 9 10 11 12 13 16 2 2 18 6 6 20 4 4 8 22 7 7 24 1 2 3 26 3 3 28 5 5 30 4 4 32 2 1 3 34 1 1 36 4 4 38 3 3 40 5 5 42 3 5 1 9 44 1 3 4 46 1 3 1 5 48 1 1 1 3 50 1 1 1 1 1 5 52 2 1 1 4 54 1 1 1 3 56 1 1 58 1 1 60 1 1 62 2 3 1 6 64 1 1 2 66 1 1 Total12121618136354224299

50 Table 6 B. Age - length key for female of C. nufar - with sectioned otolith.

Age (y) TL(cm) Total 1 2 3 4 5 6 78 9 101112 13 18 5 5 20 2 10 12 22 6 6 24 1 1 2 26 3 3 28 2 1 1 4 30 3 1 4 32 1 1 2 34 4 4 36 7 3 10 38 13 13 40 5 5 42 1 1 2 44 1 1 2 46 1 1 2 48 1 1 1 1 4 50 1 1 1 1 2 6 52 9 1 2 2 14 54 2131 7 56 3 2 5 58 3 3 60 4 3 4 4 15 62 3 3 4 10 64 5 5 8 18 66 1 1 Total 7 17 10 33 8 13 6 8 11 8 11 14 13 159

51 Table 6 C. Age structure of C. nufar population in the Arabian Sea.

TL(cm) Age (y) Male Total 1 2 3 4 5 6 7 8 9 10 11 12 13 16 9 9 20 10 43 53 24 59 267 325 28 362 52 26 440 32 139 277 46 462 36 283 32 315 40 105 45 8 158 44 12 31 18 12 6 80 48 8 8 8 8 2 5 8 45 52 1 9 2 2 1 2 2 21 56 32 41 9 60 362233 19 64 1 5 5 4 14 Total 19 102 767 730 189 35 30 20 12 11 15 14 8 1951

52 Table 6 D. Age structure of C. nufar population in the Arabian Sea

TL(cm) Age (y) Female Total

123456 7 8910111213

16 11 11

20 11 49 60

24 66 298 365

28 405 58 29 492

32 155 310 52 517

36 318 35 353

40 118 50 8 176

44 14 35 21 14 7 90

48 8 8 8 8 3 5 8 50

52 2113313324

56 32 4111

60 3 7 2 3 3 3 21

64 1 5 5 4 16

Total 22 115 859 817 211 40 33 22 14 12 16 15 8 2185

Growth performance index (phi-prime index)

While the growth performance index (Ø’) based on length frequency data was estimated at 3.14, the otolith based estimates of K and L∞ gave a value of 2.945.

53 3.2.2. Maturation and Spawning

The maturity key for the identification of different stage of gonads in males and females of C. nufar is given in Annex 1A. The results of the histological assessment of the stages of ovarian development in female C. nufar are shown in Annex 1B, which show the various stages present during the spawning season, (i.e. immature, maturing, mature and ripe stages).

Development of ova to maturity

Diameters of intraovarian eggs from stages I to VI ovaries studied to understand the process of development of ova from immature to ripe for spawning are given in (Fig. 43 A-F & Annex 1). In stage-I, ova measuring 0.014-0.042 mm formed the mode and several ova measured up to 0.098-0.126 mm. These ova may be considered as the general egg stock. In stage-II, a batch of ova forming mode at 0.224-0.252 mm got separated from the general egg stock for further development. Few larger ova measured up to 0.308-0.336 mm. In stage-III, ova with two modes close to each other (0.226-0.229 mm and 0.350-0.378 mm) were present and the maximum size of ova measured up to 0.462 mm. Both the batches of eggs were in maturing condition. In stage- IV, the mode formed by ova of 0.266-0.294 mm in stage-III was stationary and an additional advanced mode at 0.434-0.462 mm was evident and this mode represented the mature group of eggs. However, in this stage few ova were larger and measured up to 0.546 mm. In stage V, the mature group of ova forming mode at 0.434-0.462 mm. In stage IV further developed and formed the mode at 0.770-0.798 mm. This mode represented the ripe eggs. In stage-VI, ripe ova (mode at 0.770-0.798 mm in stage V) were absent indicating that the ripe ova were shed during spawning. The maturing group of ova forming mode at 0.266-0.294 mm was present and few larger disintegrating eggs were also seen. The batch of ripe ova (0.770-0.798 mm) present in stage V ovary was absent in spent ovary and few larger ova were in the state of resorption (Fig. 43). This shows that the ripe eggs had released during spawning. The intermediate stage of maturing group of ova (mode at 0.266-0.294 mm) present in spent ovary would take probably half the time than the immature ova

54 to attain ripe stage for spawning. Hence, it appears that individual fish would spawn more than once during the spawning season in Arabian Sea.

A 100 I

% 50

0.014-0.042 0.056-0.084 0.098-0.126 0.140-0.168 0.182-0.210 0.224-0.252 0.266-0.294 0.308-0.336 0.350-0.378 0.392-0.420 0.434-0.462 0.476-0.504 0.518-0.546 0.560-0.588 0.602-0.630 0.644-0.672 0.686-0.714 0.728-0.756 0.770-0.798 Ova diameter (mm)

Fig. 43 A. Development of ova to maturity stage I in C. nufar

B 60 II

Fig. 43 B. Development of ova to maturity stage II in C. nufar

55 30 C III 20

% 10

Fig. 43 C. Development of ova to maturity stage III in C. nufar

20 D IV 15

% 10

Fig. 43 D. Development of ova to maturity stage IV in C. nufar

F 15 V 10

% 5

Fig. 43 F. Development of ova to maturity stage V in C. nufar

56 Spawning Season

Spawning Season based on commercial catch Data

The monthly percentage occurrence of different stages of maturity of ovaries and testes for 2005-06 and 2006-07 (Figs. 44, 45) showed the ripe gonads (stage V) to occur from April to September during 2005-06 and 2006-07. However, the spent gonads were recorded only for three months (June, September and October) during 2005-06 and four months (June, August-October) during 2006-07. Occurrence of spent individuals in June may contain few fish that spawned during the later part of May and occurred during June in the catches. Similarly, spent fish recorded in October 2005 and 2006 might be due to the spawning in some fish during later part of September and recorded in October. The presence of spent fish in the commercial catches during June to October shows that C. nufar may spawn for about 6 months (May-October) and peak spawning may be between June and September in Omani waters.

Spawning Season based on Survey Data

Spawning observation

The period of spawning of C. nufar is likely to occur during the SW monsoon as the gonads in running ripe stage occurred almost exclusively in the post sw monsoon 2007/2008 in Area2, Area 3 and Area 4, which coincided with lower value of water temperatures, 18 Cº – 23 Cº, in the entire area of the Arabian Sea that took place immediately during the SW monsoon (Fig. 3, Fig. 7, Fig. 46 A & B).

57 75 60 October M (n=19) Apr 05 M (n=17) 50 40 F (n=24) F (n=24) 25 20

Percentage Percentage 0 0 I II III IV V VI I II III IV V VI

M (n=29) May 80 M (n=32) November 50 F (n=16) 60 F (n=47) 40 25 20

Percentage

Percentage 0 0 I II III IV V VI I II III IV V VI

50 June 100 M (n=31) December M (n=35) F (n=40) F (n=46) 25 50

Percentage 0 Percentage 0 I II III IV V VI I II III IV V VI

60 100 M (n=31) Jan 06 M (n=35) July 40 F (n=48) F (n=40) 20 50

Percentage

0 Percentage 0 I II III IV V VI I II III IV V VI

80 August 60 M (n=22) February 60 M (n=33) F (n=24) F (n=42) 40 40 20 20

Percentage

Percentage 0 0 I II III IV V VI I II III IV V VI

M (n=38) 100 M (n=25) March 40 F (n=34) September F (n=25)

20 50

Percentage

Percentage 0 0 I II III IV V VI I II III IV V VI Maturity stages Maturity stages

Fig. 44. Distribution of different maturity stages of gonads of C. nufar during 2005-06

58 100 60 M (n=30) October April 06 75 M (n=24) F (n=21) F (n=16) 40 50 20 25

Percentage Percentage 0 0 I II III IV V VI I II III IV V VI

M (n=20) May M (n=19) 60 F (n=20) November 60 F (n=22) 40 40 20 20 Percentage 0 Percentage 0 I II III IV V VI I II III IV V VI

M (n=55) 60 June 80 M (n=19) December F (n=65) F (n=21) 40 60 40 20 20

Percentage 0 Percentage 0 I II III IV V VI I II III IV V VI

M (n=16) Jan 07 60 F (n=24) July 100 M (n=12) 40 F (n=28) 50 20

Percentage

Percentage 0 0 I II III IV V VI I II III IV V VI

60 M (n=20) August 100 M (n=26) February F (n=33) F (n=12) 40 50 20

Percentage

Percentage 0 0 I II III IV V VI I II III IV V VI

M (n=20) 60 September 100 M (n=19) March F (n=22) F (n=19) 40 50 20

Percentage 0 Percentage 0 I II III IV V VI I II III IV V VI Maturity stages Maturity stages

Fig. 45. Distribution of different stages of gonads of C. nufar during 2006-07

59 Fig. 46 - A. Fish with running ripe maturity stage during post SW monsoon 2007

Fig. 46 - B. Fish with running ripe maturity stage during post SW monsoon 2008

60 The sex-wise monthly gonado-somatic indices of C. nufar for 2005-2007 (Figs. 47) clearly indicated that females always recorded higher GSI than males. The GSI values were higher in females than the weighted average (0.65) during July and August. For males, the GSI values were higher during May, July, August, September and October than the weighted annual average (0.281). The higher GSI values during June-September coincided with sw monsoon season which was the peak spawning season of C. nufar. The peak spawning activity of fish occurred when the water temperature declined to 23 -25 ºC during the above months (Fig. 47).

2.5 30

29 2 28

1.5 27 Female

GSI 26 Male 1 25 Temp 24 0.5 23

0 22 AM J J A S OND J FM Months

Fig. 47. Monthly gonado-somatic indices in C. nufar 2005 – 2007

The calculated monthly hepato-somatic indices of female fish registered generally higher values than the males except February and March during 2005-2007 (Fig.48). The peak value for both males and females were during September which started declining progressively up to February. The declining trend of HSI values from September to February was almost identical to the declining trend of GSI values.

61 1.8

1.6

1.4

HSI 1.2 Female 1 Male 0.8

0.6

July

May

June

April

March

August

January

October

February

December

November

September

Fig. 48. Monthly hepato-somatic indices in C. nufar 2005 -2007

During 2005-07, while the monthly relative condition factor (Kn) values (Fig. 49) for males ranged from 0.98 (August) and 1.27 (February), the values for females fluctuated between 0.96 (June) and 1.19 (February). The weighted average for males was 1.079 and for females was 1.023. In males higher Kn were recorded than the weighted average during September, October, December and February. While for females the weighted average was 1.023 and values were higher than the average during September, October, December and February.

In general, males registered better condition values than the females during various months. The average monthly Kn for 2005-2007 indicated rise and fall in the values and hence, could not be related to spawning.

62 1.35 1.3 Female Male 1.25 1.2 1.15 Kn 1.1 1.05 1 0.95 0.9 AM J J A S OND J FM

Months

Fig. 49. Monthly Kn in C. nufar 2005-2007

Occurrence of juveniles in survey areas

Juveniles of the species were found throughout the survey area, although most commonly in the three southern Zones (Fig.50-54).

Fig. 50.Juveniles distribution -Post SW monsoon 2007

63 Fig. 51 Juveniles distribution: Autumn monsoon 2007

Fig. 52 Juveniles distribution –NE monsoon 2008

64 Fig. 53 Juveniles distribution Spring Inter-monsoon 2008

Fig. 54 Juveniles distribution -Post SW monsoon 2008

65 Length at first maturity

Males measuring less than 21 cm TL and females measuring less than 23 cm were all immature. The percentage of mature individuals of both the sexes increased with increase in length of the fish. The length at 50% of maturity in males and females were estimated at 33.7 cm and 31.9 cm of TL respectively (Fig.55). This indicates that the females mature at an earlier length than males.

Male: Lm = 33.7 cm 0.8 Female: Lm = 31.9 cm

0.6 Male Male 0.4 Female

Proportion mature Proportion Female

0.2

0 10 15 20 25 30 35 40 45 50 55 60 65 Total length (cm)

Fig. 55 Length at first maturity in C. nufar

Fecundity

In C. nufar, the batch fecundity varied between 11,406 and 473,277 (Annex 2). There was general increase in fecundity as the length, body weight and gonad weight of the fish increased. The relationship between the fecundity (F) and total length of fish (TL) (Fig. 56 A) was found to be,

F = 4.3232+0.0061 TL (R2 = 0.898)

66 The relationship between fecundity (F) and body weight (W) (Fig. 56 B) could be expressed as,

F = 1.3938 + 4.3082 w (R2 = 0.9085)

The relationship between fecundity (F) and ovary weight (OW) (Fig. 56 C) was estimated as,

F =0.9857 + 3074.1 OW (R2 = 0.9921)

Fecundity in C. nufar could be well correlated to the weight of the ovary than the body weight and length of the fish as R2 =0.992.

A 500 450 y = 0.0061x4.3232 R² = 0.8977 400 350

000) 300 250 200

Fecudity (' 150 100 50 0 20 30 40 50 60 70 Total length(cm)

Fig. 56 A. Fecundity in relation to total length (cm) of C. nufar.

67 500 450 B y = 4.3082x1.3938 400 R² = 0.9085 350

000) 300 250 200 Fecndity (' 150 100 50 0 0 500 1000 1500 2000 2500 3000 3500 4000 Total Weight(g)

Fig. 56 B. Fecundity in relation to body weight(g) of C. nufar .

C 0.9857 600 y = 3074.1x R² = 0.9921 500

400

000)

300

200

Fecundity (' 100

0 0 50 100 150 200 Ovary Weight (g)

Fig. 56 C. Fecundity in relation to ovary weight (g) of C. nufar

68 Sex- ratio

The percentage composition of male and female of C. nufar for different months is shown in Fig. 57. As the sexes of the fish could not be identified externally, the fish were dissected to recognize the sex. The monthly sex-ratio for the period April 2005–March 2006 (Table. 7A) and April 2006-February 2007 (Table. 7B) indicated the overall dominance of females over males. However, the sex-ratios did not differ significantly during both the years. The monthly sex-ratios during 2005-06 were not significant at 5% level. During 2006-’07, among the monthly sex ratios, significant difference was found only during January (x2=6.4) and February 2007 (x2=5.1578).

% 40 Male 30 Female 20

Oct

Apr

Sep Feb

July

Dec

Aug

Nov

Mar

May

June

Jan-07

Jan-06

Apr-05 Months

Fig. 57. Sex ratio of C. nufar during the period April 2005 - March 2007

69 Table. 7A Sex-ratio in C. nufar during 2005-06

Total No. No. of No. of females Male: Female Chi-square value Mh ffih l Apr 05 43 19 24 1:1.3 0.5814

May 45 29 16 1:0.6 3.7556

June 81 35 46 1:1.3 1.4938

July 83 35 48 1:1.4 2.0362

Aug 75 33 42 1:1.3 1.08

Sept 72 38 34 1:0.9 0.2222

Oct 41 17 24 1:1.4 1.1952

Nov 79 32 47 1:1.5 2.8482

Dec 71 31 40 1:1.3 1.1408

Jan 06 60 36 24 1:0.7 2.4

Feb 46 22 24 1:1.1 0.087

March 50 25 25 1:1 0

Total 746 352 394 1:1.1 2.3646

*Significant at 5% level

70 Table. 7B Sex-ratio in C. nufar during 2006 -07

Month Total No. No. of No. of females Male: Female Chi-square value ffih l Apr 06 40 24 16 1:0.7 1.6

May 40 20 20 1:1 0

June 120 55 65 1:1.2 0.8334

July 40 16 24 1:1.6 1.6

Aug 53 20 33 1:1.7 3.1886

Sept 42 20 22 1:1.1 0.0952

Oct 51 30 21 1:0.7 1.5882

Nov 41 19 22 1:1.2 0.2196

Dec 40 19 21 1:1.1 0.1

Jan 07 40 12 28 1:2.3 6.4*

Feb 38 26 12 1:0.5 5.1578*

March 38 19 19 1:1 0

Total 583 280 303 1:1.1 0.9074

*Significant at 5% level

71 3.2.3. Food composition and feeding intensity

Of the 775 specimens analyzed during 2005-06 and 550 fish studied during 2006-07 for food and feeding habits of C. nufar, 523 and 324 fish had empty stomachs during the first year and second year of observation respectively.

The common food items found in the gut of C. nufar largely belonged to fishes (sardine, anchovy, Therapon), crustaceans (crab, shrimp) cephalopods (cuttlefish, squid, octopus) and polychaetes. Fish as food items of C. nufar contribute 50% and 57.96% during 2005-06 and 2006-07, followed by semidigested matter 42.9% and 39.4% respectively (Figs. 58 & 59). Rest of the food items were of minor importance and polychaetes were recorded only during 2005-06.

Fish 42.86 Crustaceans

Cephalopods 50 Polychaete worms

Semidigested matter

1.98 2.78 2.38

Fig. 58. General food composition of C. nufar during 2005-06

72 Fish

Crustaceans

39.39 Cephalopods

Semidigested matter

57.96

1.33 1.33

Fig. 59. General food composition of C. nufar during 2006-07

Food Composition during different months

During 2005-06, the volume of fish as food items of C. nufar varied from 0% (February) to 65.95% July (Fig. 60). While, the indices of semidigested matter ranged between 23.4% (July) and 100% (January), the other food items such as crustaceans (0% - 11.1%) and cephalopods (0% -10.5%) occurred for few months. Polychaetes (6.9%) were recorded only during August.

During 2006-2007, semidigested matter was the dominant item in the gut of fish during most of the months and the composition ranged between 10% (December) and 88.9% (April) (Fig. 61). The contribution of fish as food item of C. nufar varied from 11.1% (July) to 83.79% (October).

73 Fish 100 100 N=775 Crustaceans Cephalopods 83.33 80 75 Polychaete worms 73.33 73.33 65.95 Semidigested matter 60 63.01 60 53.33 52.94 55.56 47.37 40 42.11 40 41.18 40 30.14 27.78 % composition 25 23.4 26.67 16.67 20 20 10.53 11.11 4.26 6.38 6.67 5.88 5.56 6.67 0

July

May

June

Jan 06 Jan

March

Apr 05

August

October

February

December

November

September Months

Fig. 60. Monthly percentage of composition of food items of C. nufar during 2005-06

100 Fish 88.89 Crustaceans 90 N=550 83.79 Cephalopods 80 Semidigested matter 73.08 66.67 66.67 63.16 63.64 55 56.75 60 55.55 55.55 45 44.44 44.44 36.84 40 35.14 33.33 33.34 27.27

% Composition 19.23 16.22 20 11.11 8.11 7.69 10 9.09

July

May

June

Jan 07 Jan

March

Apr 06

August

October

February

December

November

September Months

Fig. 61. Monthly composition of food items of C. nufar during 2006-07

74 Food composition in different size of fish

For the food items found in the fish of different sizes, the data were pooled for 2005- 06 and 2006-07. The contributions of fish items and semidigested matter were dominant in all the size groups (Fig. 62). However, the crustaceans were preferred in minor quantities by size groups of 16-30 cm. Cephalopods and polychaetes were preferred by small and medium sized fish.

Fish 100 Crustaceans N=1325 Cephalopods Polychaetes 75 80 71.43 Semidigested matter 65.86 59.46 60.87 60 53.85 56.06 48.84 50 50 50 43.41 41.67 43.75 36.23 40 32.43 31.71 30.77 28.57 % Composition 25 20 15.39 5.41 4.65 6.25 0 2.7 2.33 0.76 1.45 0

26-30 16-20 31-35 21-25 36-40 51-55 41-45 56-60 46-50 61-65 Size groups (TL-cm)

Fig. 62. Food items found in the gut of various size groups of C. nufar during 2005-2007 (pooled)

75 Feeding intensity in relation to month, maturity stage and size of fish

Of the 990 fish analyzed 668 had empty stomachs. The empty stomachs were dominant during all the months excepting July, August and October 2006 (Fig. 63). Active feeding (full and 3/4full stomachs) was found to range from 0% (February 2006) to 58.8% (October 2006) of individuals. Moderate feeding (1/2full stomach) was observed from 0% (December 2005, January-February 2006) to 14.6% (December 2006) of fish. Poor feeding intensity (1/4full stomach) was to vary from 0% to 24.4% and the highest percentage was recorded during January 2007.

100 75 81 40 52 41 41 N=990 53 51 80 40 61 40 43 42 Empty 60 41 80 41 80 40 1/4 full 48 1/2 full 40 3/4 full

Feeding intensity (%) Full 20

Jul

Jun

Oct Oct

Apr

Feb Feb

Dec Dec

Aug

Nov Nov

May

Sept

Jan 07 Jan Jan 06 Jan

March March

Sept 05

Months

Fig. 63. Feeding intensity in C. nufar relation to months

76 There was no consistency in the occurrence of active, moderate and poor feeding in various stages of maturity of males and females (Fig. 64). However, empty stomachs were dominant in all the maturity stages that ranged from 50% (stage IV) to 76% (stage I) in males and 47.2% (stage VI) to 77.57% (stage I) in females.

100

M= male (n=477) 80 F= female (n=513) 60

13.89 40 11.11 22.22 24.24 2.78 Empty 11.03 17.39 1/4 full Feeding intensity (%) 9.55 14.29 10.29 28.57 5.56 1/2 full 20 7.35 2.81 5.15 4.35 12.04 10.28 30.56 3/4 full 0.93 22.22 22.22 25 24.24 4.01 17.65 16.85 16.91 17.39 14.28 Full 7.23 9.35 0 M FMFMFMFMFMF I II III IV V VI Maturity stages

Fig. 64. Feeding intensity in different maturity stages of C. nufar

77 Feeding intensity in relation to different sizes of C. nufar showed inconsistency of feeding among fish (Fig. 65). The percentage of empty stomachs ranged from 55.6% (51-55 mm size group) to 75% (41-45 mm size group). Active feeding was observed in fish of sizes 61-65 mm (66.7%) and 71-75 mm (100%).

100 64.29 N=990 65.79 67.28 65.91 62.5 33.33 80 70.68 64.37 75 55.56

60 Empty 100 100 1/4 full 1/2 full 40 10.2 4.55 11.11 3/4 full Feeding intensity (%) 10.53 66.67 2.27 12.5 Full 13.36 1.02 11.11 2.63 11.28 13.46 1.02 2.27 20 2.63 8.33 12.5 3.01 5.5 8.91 2.78 3.01 2.14 25 18.42 23.47 22.22 12.03 11.62 13.36 13.89 12.5 0

16-20 21-25 26-30 31-35 36-40 41-45 46-50 51-55 56-60 61-65 66-70 71-75 Length class (cm)

Fig. 65. Feeding intensity in different size groups of C. nufar

78 3.2.4. Population dynamics and stock assessment

Mortality

The total mortality (Z) of C. nufar estimated by the length converted catch curve method was found to be 1.24 (SE=0.04). The Beverton-Holt estimation provided the Z value as 1.36 (SE=0.18) which was comparable to the value estimated by the length converted catch curve technique. The Powell-Wetherall method gave the mean Z/K as 3.56 (SE=0.15). The average Z calculated by both length converted catch curve and Beverton and Holt methods which stood at 1.30.

For C. nufar, the natural mortality coefficient (M) calculated by the empirical methods of Pauly (1980), Alagaraja (1984) and Rikhter and Efanov (1976) stood at 0.61, 0.44 and 0.80 respectively. The average of the above three values (0.62) was close to the estimate obtained by Pauly’s techniques (M=0.61). Hence, for further analysis, the M value was taken as 0.61. The calculated fishing mortality (F) of C. nufar was: total mortality minus natural mortality (F = Z – M) was 0.68.

Exploitation rate (E) and Exploitation Ratio (U)

The exploitation rate (E) for the pooled data of 2005-’06 and 2006-’07 was 0.52. The value of U estimated for the combined data of both the years was 0.38.

Yield, Standing stock, Total stock or Biomass and MSY

The catches of C. nufar for the year 2005-2006 estimated 2,075 t and for 2006-2007 2,018 t were considered as the yield of respective years. From the average annual yield for both the years (2,047 t), the estimated annual average standing stock was 3,010 t and the annual average total stock was 5,387 t. The MSY estimated by Cadima’s formula was 1,953 t. The Thompson and Bell analysis also showed a close MSY of 1,946 t (Fig.66). The current yield and MSY estimates indicate that the stock of C. nufar is marginally overfished.

79 The predicted yield and biomass decreased with the increase of F. The predicted total biomass of 16,052 t decreased to 12,713 t at F=0.1, to 6,167 t at F= 0.5 and to 5,235 t at F= 0.6 and at F= 2.0, the yield and biomass stood at 18 t and 57 t respectively (Fig. 66).

2500 18000 16000 2000 MSY 14000 Yield (t)

Biomass (t) 12000 Biomass (t) 1500 10000

Yield (t) 8000 1000 6000

500 4000 2000 0 0 01234 F-factor

Fig. 66. Predicted yield for C. nufar for different F

Yield-per-recruit and Biomass-per-recruit

In the stock of C. nufar, the yield-per-recruit (Yw/R), total biomass-per-recruit (TB/R) and stock spawning biomass-per-recruit (SSB/R) were estimated for the F- values ranging from 0-2 (Fig.67). It was evident that the MSY/R was 115.4 g at the F of 1.48. However, the current estimated F was 0.68. When F was zero, the estimated TB/R and SSB/R were 759 g and 553 g respectively (Fig. 67). These values decreased with the increase in F. At the current F equal to 0.68, the values of Yw/R, TB/R and SSB/R were 111 g, 260 g and 191 g respectively.

80 800

700

600 YW/R (g) 500 SSB/R (g) 400 TB/R (g)

Yield (g) 300

200

100

0 0 0.4 0.8 1.2 1.6 2 2.4 Fishing mortality

Fig. 67. Equilibrium yield-per-recruit and biomasses-per-recruit in C. nufar

against F (Yw/R, yield-per-recruit; SSB/R, stock spawning biomass-per-

recruit; TB/R, total biomass-per-recruit)

Optimum length of capture (Lopt)

From the empirical equation (Froese and Binohlan, 2000), the optimum length of exploitation was calculated at 40.6 cm. The Lopt/L∞ for C. nufar was calculated at 0.58.

81 3.3. Distribution of C. nufar in relation to oceanography parameters

The fish C. nufar occurred throughout the survey area but more commonly in the south to Masirah Island in waters shallower than 100 m depth (Fig. 27). Abundance of fish was highest from January to June. During this period, the depth range of occurrence of the species appeared greatest. There was difference in the distribution pattern of C. nufar between the surveys done in post southwest monsoon of 2007 and 2008. While the fish occurred in areas 1, 2 and 3 during post southwest monsoon period of 2007 period, during southwest monsoon period of 2008 the fish occurred in areas 2,3 and 4.(Fig. 69,73). It has been noted that the C. nufar prefer to occur at temperature between 20 and 24ºC during NE monsoon and Spring Inter monsoon while in autumn inter monsoon it occur at 26–28ºC. During Post southwest monsoon 2008 the fish was found at temperature between 17ºC and 21ºC (Fig. 68).

140

Post SW 2007 120 Autumn monsoon 2007 100 Northeast Monsoon 2008

80 Spring monsoon 2008 Post SW 2008 60

Catch (kg)

0 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 Temperature ºC

Fig.68. Catch of C. nufar at different time of monsoon.

82 Fig. 69 Distribution of C. nufar: Post SW monsoon 2007

Fig. 70 Distribution of C. nufar : Autumn monsoon 2007

83 Fig. 71 Distribution of C. nufar: NE monsoon 2008

Fig. 72 Distribution of C. nufar: Spring Inter-monsoon 2008

84 Fig. 73 Distribution of C. nufar: Post SW monsoon 2008

Response C. nufar to OMZ

When the OMZs were widely prevalent, C. nufar was found closer to the coast mainly within Masirah and Sawqirah bays during the post south west monsoon in 2007 (Sept-Oct) and post south west monsoon in 2008 (Aug-Sept) (Figs. 26,69,73 ). By NE monsoon in 2008 (Jan-Mar), when most of the shelf waters were well oxygenated, C. nufar was widely distributed throughout the survey area (Fig. 24, 71). During the spring inter-monsoon period (Apr-Jun), waters were largely deoxygenated and C. nufar were found in a narrow band nearer to shore (Fig. 25, 72). It appears that C. nufar is adapted to low oxygen level (Fig. 75). These conditions were clearly reflected on the catch rates during different voyages. The highest catch rate was recorded in stratum between 20 and 50 m during post SW monsoon 2007 and 2008 while it was widely distributed during NE monsoon 2008 between 20 – 150 m depth (Fig.74).

85 2200 2000 1800 1600 1400

-2 1200 20-50 m 1000

Kg. kg 50 - 100 m 800 100 - 150 m 600 400 150 - 250 m 200 0

2008 2007

Sprin

Autum

2008

monsoon monsoon

NE monsoon

Post SW 2008

Post SW 2007

Fig.74 Catch rate of C. nufar by stratum during monsoon periods

140 Post SW 2007 120 Autumn monsoon 2007 100 Northeast Monsoon 2008 80 Spring monsoon 2008

Catch(Kg) 60

0 012345 -1 Dissolved O2 ml l

Fig.75. Occurrence of C. nufar at different level of bottom dissolved oxygen.

86 4. Discussion

Climatic variability and increasing pressure from exploitation of marine resources and other human activities would influence and potentially modify the state and functioning of marine ecosystems. The interaction of both climate and exploitation are probably substantially involved in most cases for decline in fish populations and the associated ecosystem changes as climate may cause failure in a fishery management system, fishery exploitation may disrupt the ability of the population to resist or adjust to climate changes

The two unique features of the Arabian Sea that significantly influence productivity in the region are the seasonal monsoons and the occurrence of extensive oxygen minimum zones (Wishner et al., 1998). The NE monsoon occurs from November to mid February and the SW monsoon, with sustained strong winds, typically occurs from June to mid September (Weller et al., 1998). However, upwelling favorable winds along the Oman coast can begin as early as April (Kindle and Arnone 2001).

The productivity in the region has a strong seasonal signal associated with the SW Monsoon (Wishner et al., 1998). Filaments and jets are created by the SW monsoon, which are capable of exporting cool, nutrient rich waters hundred of kilometres offshore (Brink et al., 1998). Mid-open ocean upwelling occurs offshore during the SW monsoon (Smith and Bottero1977). Features along the Oman coast produced by the SW monsoon tend to persist into the autumn intermonsoon period (Kindle and Arnone 2001).

The effect of the monsoon periods on water temperatures is well documented (Wishner et al., 1998; Lee et al., 2000, Weller et al., 2002). Warm sea surface temperatures are typically seen offshore during the SW monsoon period, with cooler, more recently upwelled water present near-to-shore. Sea surface temperatures are coolest after the NE monsoon (Wishner et al., 1998). CTD

87 casts during 2007–08 showed a similar pattern; nearshore temperatures were warmest immediately before the SW monsoon, upwelling was evident immediately after the SW monsoon, and sea surface temperatures were coolest immediately after the NE monsoon period. T h e c ooler waters after the NE monsoon are the result of cross-basin variations in windstress and convective overturning bringing cool waters to the surface (Lee et al., 2000).

Cold, less saline water was noted at mid-depths, particularly to the NE of Masirah Island and Salalah region after the NE monsoon period. Salinity minima in midwater have been noted in other studies (Shetye et al., 1992, Weller et al., 2002) and are thought to be the result of fresh coastal water sinking offshore or seasonal effects of evaporation and capping of the thermocline. Although low salinity intrusions were noted at this time, salinity in surface and deep waters was typically high. Elevated surface salinities during and after the NE monsoon may be the result of southward advection of highly saline surface waters from the Sea of Oman (Warren et al., 1966) or strong evapotranspiration association with the monsoon (Lee et al., 2000)

The OMZs is a midwater zone of low oxygen that extends from just below the mixed layer to more than 1000 m in some areas. The core of this layer is defined as -1 the region where oxygen is less than 0.1 ml l . .There is some debate as to how this layer forms – large local consumption rates from high productivity in surface waters; very slow movement within the layer, which allows decaying organic matter to consume all oxygen present; low oxygen concentration in the waters entering the layer from the southern ocean (Olson et al., 1993); or the lack of an opening to seas to the north (Morrison et al., 1999).

In 2007–08, the OMZ was found at depths as shallow as 40–50 m immediately after the SW monsoon. After the NE monsoon, waters were well oxygenated and the OMZ was typically found at depths of 150–200 m, if observed at all. This depth pattern was noted in several other studies (Weller et al., 1998, Wishner et al.,

88 1998), although Morrison et al. (1999) found that the OMZ had a maximum depth of approximately 150 m. Morrison et al. (1999) found dissolved oxygen was typically higher to the south, similar to observations described here. Naqvi et al. (1992) found dissolved oxygen was at minimum after the NE monsoon, increased throughout the spring inter monsoon, and reached a maximum during the SW monsoon period. Dissolved oxygen levels less than 0.1 ml l-1 have physiological limitations on small organisms (Morrison et al., 1999). However, larger organisms, such as fish, are limited by slightly higher levels of dissolved oxygen of about 0.3 ml l-1 (Ashjian et al., 2002). The present study indicates that C. nufar can thrive even in areas where oxygen levels were low.

The distribution of C. nufar was not similar for the surveys conducted in post southwest monsoon periods during 2007 and 2008.The fish occurred in areas 1, 2 and 3 during post southwest monsoon period of 2007; however, it was captured in areas 2, 3 and 4 during post southwest monsoon period of 2008. The differences in biomass estimates between post southwest monsoon surveys may be due to the after-effects of the cyclone (Gonu) that hit Omani coast during June 2007 and/or the slight difference in timing between the two surveys (the 2007 survey started 1.5 months later than in 2008).

The commercial catches of C. nufar consisted of fish with crucial lengths. The estimated length at first capture was 31.4 cm, and the females matured at the average length of 31.9 cm, and males matured at the average length of 33.7 cm. During July 1999 - July 2000, the catches of C. nufar from the trawl landings of Oman consisted of about 40% of immature fish (McIlwain et al., 2006). However, in the present study, the fish measuring up to 29 cm of TL contributed to 49% during 2005-06, 36 % during 2006-07 and 42 % for the pooled data. This clearly indicates that besides immature individuals, sizeable portions of mature fish are also caught prior to spawning causing recruitment overfishing. The current situation is a matter of management concern and warrants for suggesting the legal size for capture.

89 The value of exponent b in length-weight relationship of C. nufar from South Africa was found to be 2.7831 (Coetzee and Baird, 1981) which fell within in the range generally found for fish (Ricker, 1971). It is interesting to note that while there was significant difference between the slopes of the fork length-weight regressions of males and females in an earlier study from the Arabian Sea off Oman (McIlwain et al., 2006), in the present study, no such significant difference between the total length-weight relationships could be established. However, the b values of males and females in the length-weight relationships in both the studies are comparable i.e. the males and females registered the b values of 2.872 and 2.849 respectively in the former study, and 2.906 and 2.890 respectively in the present study.

The VBGF parameters estimated from different regions of the Indian Ocean indicated variations in the parameter values (Table 8). However, the maximum size of the fish in the commercial catch was reported to range between 60cm and 75 cm (Van der Elst, 1981; Bauchot and Smith, 1984; Garratt, 1985).

Table 8. Comparison of growth parameters of C. nufar from different studies

-1 Country L∞ (TL; cm) K y t0 (y) Reference

Gulf of Aden 100 0.067 -2.96 Pauly (1978), based on Druzhinin (1975)

Gulf of Aden 65 0.18 - Edwards et al., (1985)

Mozambique 70 0.17 - Timochin (1992)

South Africa 95.4 0.654 -2.62 Coetzee and Baird (1981)

Oman 69.9 0.286 -0.180 Present study

90 In the present study, for the largest sized C. nufar recorded in the commercial catch

(Lmax= 64 cm), the estimated age based on length frequency would be around 11 years; however, otolith indicated an age of about 13 years (Table 6 A&B). The Pauly’s equation (3/K) also indicated the longevity of the fish would be around 10.5 years (Pauly, 1983a). From the South African waters, based on otolith, the life span of the fish measuring 76.3 cm was estimated to be around 22 years (Coetzee and Baird, 1981). As per the otolith based length at age estimates provided (Coetzee and Baird, 1981), the age of the fish measuring 64 cm in the South African waters would be between 14 and 15 years. The difference in the age may be due to the differences in the environmental parameters.

The growth performance index (Ø’) of C. nufar in the present study was estimated at 3.14 based on length data and 2.95 based on otolith analysis. It suggests that the species is comparatively growing faster in the Omani waters than in Gulf of Aden where the Ø’ was equal to 2.83 (Druzhinin, 1975) and 2.88 (Edwards et al., 1985) and to 2.92 in Mozambique (Timochin, 1992). The sagittal otoliths in C. nufar showed the formation of one opaque and one translucent ring each year. The high percentage of otoliths with opaque edge was observed during July – January and hyaline edge observed during March - June could be related to low and high values of sea temperature respectively. The species C. nufar has a long life span in the Arabian Sea off Oman; the maximum age estimated was 13 years for the fish 64 cm of total length.

During the SW monsoon season due to the drop in temperature in the surface water and higher primary productivity, few species of sparids including C. nufar spawn along the Arabian Sea coast of Oman (McIlwain et al., 2006). However, the period of spawning of C. nufar reported from May to August (4 months) (McIlwain et al., 2006) was found to extend from May to October (6 months) in the present study (Figs. 44 & 45). It is interesting to note that C. nufar spawned during winter (June - October) along the coast of South Africa (Garratt, 1985; 1991) which may be due to

91 the optimum seawater temperature. The optimum temperature for spawning in C. nufar lies between 18oC and 20.3oC (Garratt, 1985).

Until recently, community level spawning patterns in tropical demersal fish have often been correlated with gross seasonal patterns in environmental condition that might affect food supply, growth and dispersal of larvae (Robertson, 1991). One of the main predictions of this gross environmental seasonality hypothesis is that because the mechanisms are supposedly simple, they should affect communities in the same way, such that the two variables, spawning and environmental conditions, match each other (Robertson, 1991). In this study, C. nufar spawning pattern appeared to match with one gross environmental condition, i.e. the SW monsoon, when larval survivorship is probably maximized due to an increase in primary productivity. During the SW monsoon, regular upwelling events occur when seawater temperatures can decline to 16–18 ºC, but generally average to 20ºC (Sheppard et al., 2000; Wilson et al., 2002). The pelagic productivity during the summer upwelling months may be two orders of magnitude than that during winter (Sheppard et al., 2000), possibly optimizing larval survival and growth. For many species of fish, water temperature can have a pronounced effect on spawning, particularly the rate of gonad development, with sudden increases in temperature triggering maturation and ovulation (Bye, 1990). The exception was noticed in this study where our results suggest that C. nufar spawned mostly during the monsoon period when the water temperature declined to 22–25 ºC. This concurs with the findings of Sadovy (1996), who suggested that many species of sparids spawn during colder periods, compared with co-occurring reef fishes.

Sea water temperature can influence spawning of fish (de Vlaming, 1972; Mischke and Morris, 1997) and in some sparids there was relationship between SST and the timing of spawning (Dubovitsky, 1977; Buxton, 1990).

Species that could not withstand global warming-induced increase in water temperature would be left with two alternatives, either they would shift their geographic ranges to remain in waters of suitable temperature or the adults would

92 tolerate increased water temperatures; but, could adopt the tactics of spawning in cooler months as in some sparids (Gucinski, et al., 1990)

Spawning and subsequent larval production often appear to be timed to match with periods when appropriate food is likely to be abundant (Robertson et al., 1988). Shifting of spawning season to cooler months in tropical species would possibly reduce the duration of spawning and hence, there could be far reaching consequences in the structure of fish faunas and fisheries. The change in timing of spawning and subsequent recruitment in reefs or estuaries depended on seasonal hydrographic patterns for larval supply would harm recruitment of desired species and may lead to recruitment of different species (Gucinski, et al., 1990). Further, the eggs and larvae depending on seasonal hydrographic patterns for transport to nursery and feeding grounds could be transported to inappropriate areas (Norcross and Shaw, 1984). Such alterations in recruitment patterns will lead to major species shifts in fish community structure and biodiversity in an area.

Depending on the spawning habits of teleostean fishes (Hickling and Ruternberg, 1936; James, 1967), C. nufar falls under the group III fishes that spawn more than once a year similar to several other fish species in the Indian Ocean region (Prabhu, 1956; Jayabalan, 1986; Reuben et al., 1992). Estimating size at sexual maturation in relation to capture size is essential to minimize the catch of immature individuals and to determine the size at which individuals enter the reproductive population (Sadovy,1996). In the present study, females matured at an earlier length than males as observed earlier (McIlwain et al., 2006). In some demersal fish species of Oman, female attains maturity earlier than male (McIlwain et al., 2006). In the South African waters, 50% of maturity in female C. nufar was obtained at 25 cm FL (Garratt, 1985) which was lower than the observed Lm in female in the present study (31.9 cm TL or 28.3 cm FL) and in the earlier study (29.2 cm FL) from Oman (McIlwain et al., 2006). While, the male to female ratios of C. nufar tested by chi- square test did not differ significantly in the Omani waters (McIlwain et al., 2006) as observed in the present study, in South Africa, the male to female ratio was found to be 1: 2.5 (Garratt, 1985a) and 1: 2.0 (Buxton and Garratt, 1990). Sex change, where

93 individuals function as both sexes either sequentially or simultaneously, is a common reproductive strategy in tropical reef fishes, occurring in nearly 50% of families (Hourigan and Kelly, 1985; Sadovy,1996). Protogyny and protandry have been found in 10 families and the family sparidae is one of them. Establishing whether a species undergoes sex change is critical to fisheries management. Protogynous species, for example, are particularly vulnerable to overfishing as gear selectivity reduces the reproductive capacity of these species by removing the female fish in the older year classes, thereby decreasing the probability that eggs are successfully fertilized (Huntsman and Schaaf, 1994; Sadovy,1996). Determining the mode of reproduction for a hermaphroditic species is essential as managing such a fishery can be considerably complicated if there is a bias in the sex-ratio that goes undetected (Ferreira, 1993). To establish sexual functionality for C. nufar definitive classification must come from a study of gonadal histology or through experimental manipulation (Sadovy and Shapiro, 1987). If the former is to be performed, it should involve a large sample size that is representative of all size classes and taken monthly from one location (Sadovy and Shapiro, 1987).

In the present study, the fecundity of C. nufar generally increased with the size, body weight and ovary weight of fish and the minimum and maximum fecundity ranged from 11,406 to 473,277 eggs in fish measuring 29 cm and 63.4 cm respectively (Annex 2). However, slightly higher values of fecundity that ranged from 13,050 to 560,566 eggs were reported in the fish of the sizes 21.1 cm to 62 cm from the South African waters (Garratt, 1985).

The food analyses of the gut clearly indicated the fish as the most preferred food of C. nufar that also on cephalopods, crustaceans and polychaete worms (Figs. 58, 59). While, the crustaceans were preferred in minor quantities by size groups of 16-30 cm, cephalopods and polychaetes were preferred by small and medium sized fish. Feeding on nektonic and benthic forms by C. nufar especially on fish as primary food item and occasional feeding on cephalopods crustaceans and polychaetes has been observed from the Gulf of Aden (Druzhinin, 1975) and South African waters (Coetzee and Baird, 1981; Smale, 1986).

94 Occurrence of empty stomachs in several species of fishes from the Arabian Sea is common (Sreenivasan, 1974; Naik et al., 1990; Kalita and Jayabalan, 2000). About 54.9% of C. nufar occurred with empty stomachs from South Africa (Coetzee and Baird, 1981). However, in the present study from Oman, the empty stomachs in C. nufar accounted for about 67.5% (Figs.63, 64, 65). Though, the spent fish feed actively (James, 1967; Jayabalan and Ramamoorthi, 1985; Jayabalan, 1988) to balance the lost energy during spawning, active, moderate and poor feeding in C. nufar occurred in all the maturity stages of males and females indicating inconsistency in feeding intensity (Fig. 64). The incidence of empty stomachs in fish may be related to the regurgitation (Job, 1940); high calorific value of the diet consumed (Longhurst, 1957; Sreenivasan, 1974) and faster rate of digestion (Qasim, 1972).

No information is available on the mortality parameters of C. nufar from previous studies. In the present study, the total mortality (Z), natural mortality (M) and fishing mortality (F) were estimated at 1.3, 0.62 and 0.68 respectively. The Z/K was found to be 3.56 (SE=0.15). In fishes, the growth coefficient K is closely related to natural mortality and high K indicates high natural mortality (Vivekanandan, 2005). The M/K ratio is almost constant among closely related species and similar taxonomic groups (Beverton and Holt, 1959; Banerji, 1973) that varies from 1 to 2.5 (Beverton and Holt, 1959). In the present study, the M/K ratio estimated for C. nufar was 2.13.

The optimum exploitation rate (E) is assumed to be close to close to 0.5 and the sustainable yield is optimized when F≈M (Gulland, 1971). In the present study, the estimated E was equal to 0.52 which is marginally higher than the optimal rate of exploitation. As the MSY estimated by the Cadima’s estimator (1,953 t) and Thomson and Bell technique (1,946 t) was slightly higher than the yield (2,047 t) obtained during the period of study. The current yield and MSY estimates indicate that the stock of C. nufar is marginally overfished. To harvest the fish at the estimated MSY level, a reduction of 17% of effort from the present level (F= 1) is

95 needed. The MSY per recruit estimated as 115.4 g at 1.48 F appears not realistic as at the current F = 0.68, the stock is subjected to overexploitation.

The effects of fishing of larger individuals, besides, having substantial direct consequence on the population, can influence the capacity of population to withstand climate variability through other pathways such as effects on migration and parental care. Hence, in an ecosystem, elimination of species that might occur due to fishing may be destabilizing with reduced resilience to perturbations. Indiscriminate exploitation of marine resources might promote increased turnover rates in marine ecosystems, which would worsen the effects of environmental changes. Overall reduction in marine diversity at the individual, population and ecosystem levels would lead to a reduction in the resilience and an increase in the response of populations and ecosystems to future climate variability and change. Hence, future management plans need to consider the structure and functioning of populations and ecosystems in a broader sense to help maximise the ability of marine fauna to adapt to future climates.

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Arntz, W.E., E. Valdivia and J. Zeballos, 1988. Impact of El Niño 1982–83 on the commercially exploited invertebrates (mariscos) of the Peruvian shore. Meeresforsch, 32: 3–22.

Ashjian, C. J., S. L. Smith, C. N. Flagg and N. Idrisi, 2002. Distribution, annual cycle, and vertical migration of acoustically derived biomass in the Arabian Sea during 1994-1995. Deep-Sea Res. II 49: 2377-2402.

Banerji, S.K., 1973. An assessment of the exploited pelagic fisheries of the Indian seas. Proc. Symp. Living Resources of the Seas around India, CMFRI Special Publication, 176-183 p.

Banse, K., 1984. Overview of the hydrography and associated biological phenomena in the Arabian Sea off Pakistan. In: B. U. Hag and J. D. Milliman, Editors, 1984. Marine Geology and Oceanography of Arabian Sea and Coastal Pakistan, Van Nostrand Rheinhold, New York, pp. 271–304.

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109 Annex.1. Classification of maturity stages in C. nufar

Stage of Male Female Position of last mode and maturity maximum diameter of Nature and extent of testis in Nature and extent of ovary in Appearance of ova under ova (in parentheses) in body cavity body cavity microscope m.d. (1 m.d. = 0.014mm)

I- Immature Small, transparent, pale, Small, transparent, pale, Irregular, small, yolkless/yolk 1-3 (9) occupying a very small occupying a very small portion deposit just started, transparent portion to 1/3 of body cavity to 1/3 of body cavity, ova not with clearly visible/partially visible to naked eye visible nucleus

II- Maturing 1 Whitish, translucent, Pale yellow, granular ova visible Medium sized, assume round 16-18 (24) occupying about 1/2 of body to naked eye, occupying about shape, opaque, with fair cavity 1/2 of body cavity amount of yolk

III- Maturing 2 Creamy white, occupy about Pale yellowish, blood vessels Medium sized, opaque, fully 25-27 (33) 3/4 of body cavity visible on dorsal side, ova clearly yolked visible, occupying about 3/4 of body cavity

IV- Mature Creamy white, soft, Pinkish yellow, blood vessels Large sized, mature, 31-33 (39) occupying about full length prominent, large ova transparent at periphery of body cavity prominently visible, occupying about full length of body cavity

V- Ripe/Runing Creamy white, occupying full Pinkish yellow, occupying full Eggs spherical, measured 0.77 55-57 (60) length of body cavity, milt length of body cavity, ovarian mm - 0.84 mm in diameter, a exuded on slight pressure wall thin, large ova separated single oil-globule and narrow from each other and oozed out perivitelline space present under slight pressure

VI- Spent Flabby, little reddish, occupying Flaccid, reddish, occupying Medium sized ova present with 19-21 (-) about 1/2 of body cavity. about 1/2 of body cavity disintegrating ripe ova

110 Annex 1B. Developmental stages of C. nufar ovary based on histological examination

Immature Ovary is small, tightly packed and dominated by previtelogenic, primary growth stage oocytes. The gonad wall is thin with no evidence of prior spawning in the form of post ovulatory follicles, atretic cortical alveolus (yolk vesicle) or vitelogenic (yolk gobule and migratory nucleolus stage) oocytes, brown bodies or scaring (stromal tissue) in the gonad

Maturing, Mature Yolk vesicle stage is the most advanced oocyte present. The ovary is dominated by previtellogenic, primary growth ooctyes (chromatin nucleolar and perinucleolar stages)

Ripe Ovary is in active vitellogenesis. Oocytes in all stages of development are present: yolk vesicle, yolk globule and mingrstory stages oocytes dominate. Yolk globule or migratory-nucleolus stages are the most advanced oocyte(s) present.

111 Annex.2. Fecundity in C. nufar

S. No. Total length (cm) Total wt (g) Gonad wt. (g) Fecundity

1 28 280 9.79 30,574 2 28.8 310 3.77 12,897 3 29 312 3.5 11,406 4 30.6 382 4.72 14,759 5 31.7 420 4.22 12,592 6 32.2 420 5.91 13,132 7 32.6 448 9.19 27,202 8 32.7 470 7.32 18,922 9 34 486 8.17 23,423 10 34.5 516 5.99 17,718 11 34.5 546 11.42 32,843 12 35 552 10.29 31,919 13 36.2 692 12.16 36,188 14 36.3 636 6.94 20,105 15 38 680 10.51 35,996 16 38.3 800 25.65 77,232 17 38.8 546 10.19 31,772 18 43.7 1032 21 65,583 19 47 1314 30.45 79,809 20 47.2 1326 53.91 168,522 21 50.2 1788 63.69 204,699 22 51.6 1986 63.05 200,940 23 51.8 1805 39.71 109,281 24 52.5 2063 46.82 139,008 25 56.4 2414 77.91 195,632 26 61.5 3178 103.08 326,248 27 63.4 3730 185.91 473,277

112