BIOLOGY, STOCK ASSESSMENT AND FISHERY MANAGEMENT … · There was no dominance of either sex in the...
Transcript of BIOLOGY, STOCK ASSESSMENT AND FISHERY MANAGEMENT … · There was no dominance of either sex in the...
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
I
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
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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.
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� 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 (�) 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.
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� 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.
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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.
� Dissolved O2 in the surface water was high (4–6 ml l-1) in all areas dur ing
al l 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).
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� 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.
VII
� 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
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� 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.
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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.
X
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
XI
3.3. Distribution of C.nufar in relation to oceanography parameters…………………82
4. DISCUSSION……….………………………………………………………………..87
5. REFERENCES……………………………………………………………………….97
XII
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
XIII
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
XIV
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 during2005-06………………………………………………………………………………..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
XV
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 during2005-06 …………………………………………………………………....................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 during2005-2007 (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 t ime 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
XVI
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
1
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
s tu d y w a s co nd u c t ed i n t wo p ha s es . 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
the fisheries sector o f t h e c o u n t r y to manage the resource sustainably.
1.1 Literature review
The fisheries sector has always been important to Oman as a significant source of
a n i m a l p r o t e i n and employment. Prior to the discovery of oil and i t s
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.
2
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,
3
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 di f fers 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 a re ca r r i ed rapidly away from the
reef and its dependent high density plankton feed in g f i s h (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, 1 9 7 8 ; 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
4
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 bot tom 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 t h e Arabian Sea between the lat i tude
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; Kumar et al., 2001a; Weller et al., 2002) and the strongest annual
variability in water circulation (Esenkov et al., 2003).
5
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
6
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
concentration less than 0.1 ml l-1
(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
than 0.1 ml l-1
(Morrison et al.1999), larger organisms, such as fish, are limited by
sl ightly higher levels around 0.3 ml l-1
(Ashjian et al., 2002). However, demersal
fish and crustaceans (motile organisms) were rarely found where oxygen
7
concentration was below 1.5 ml l-1
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.
8
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.
9
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
No.
Voyage period Season demersal
stations
CTD
station
1 17 Sep 2007 - 15 OctPost SW
monsoon101
215
2 1 Nov - 17 Dec 2007Autumn Intermonsoon
104253
3 29 Jan 2008 - 18 Mar NE monsoon 120 348
4 19 Apr - 10 Jun Spring Intermonsoon 123 319
51 Aug - 23 Sep 2008
Post SW monsoon122 256
Total 570 1391
10
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
11
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.
12
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,
13
= ∗ 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
14
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.
15
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
16
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
17
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
used. =∑( )2
Where, O = observed number of males and females in each month
E = expected number of males and females in each month
18
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.
19
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.
20
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 − ) ,
21
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.
22
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
23
Fig. 4. Autumn inter- monsoon 2007: Sea surface temperature
Fig. 5. NE monsoon 2008: Sea surface temperature
24
Fig. 6. Spring Inter- monsoon 2008: Sea surface temperature
Fig. 7. Post- SW monsoon 2008: Sea surface temperature
25
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
26
Fig. 9. Autumn Inter-monsoon 2007: Bottom temperature
Fig. 10. NE monsoon 2008: Bottom temperature
27
Fig. 11. Spring Inter-monsoon 2008: Bottom temperature
Fig. 12. Post- SW monsoon 2008: Bottom temperature
28
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
29
Fig. 14. Autumn monsoon 2007: Surface salinity
Fig. 15. NE monsoon 2008: Surface salinity
30
Fig. 16. Spring Inter-monsoon 2008: Surface salinity
Fig. 17. Post- SW monsoon 2008: Surface salinity
31
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
32
Fig. 19. Autumn monsoon 2007: Bottom salinity
Fig. 20. NE monsoon 2008: Bottom salinity
33
Fig. 21. Spring Inter-monsoon 2008: Bottom salinity
Fig. 22. Post- SW monsoon 2008: Bottom salinity
34
Dissolved oxygen
Generally, the dissolved oxygen was high (4–6 ml l-1
) 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
35
Fig. 24. NE monsoon 2008: Bottom dissolved oxygen
During the spring inter-monsoon period (April–June), the concentration of dissolved
oxygen was low in the bottom waters, 0.1 – 1 ml l-1
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
pockets of shallow areas (40-50 m) on the shelf, oxygen poor waters (up to 0.3 ml l-1
)
were present (Fig. 26).
36
Fig. 25. Spring inter-monsoon 2008: Bottom dissolved oxygen
Fig. 26. Post- SW monsoon 2008: Bottom dissolved oxygen
37
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)
38
Fig. 27. C. nufar distribution from all trawl survey stations
Fig. 28. Biomass of C. nufar in different depth zones in the Arabian Sea survey
0
250
500
750
1000
1250
1500
1750
2000
Voyage 1 Voyage 2 Voyage 3 Voyage 4 Voyage 5
Biom
ass
( t)
20 – 50 m
50 – 100 m
100 – 150 m
150 – 250 m
39
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.
Fig. 29. Length frequency distribution of C. nufar in the commercialcatches during 2005-06
0
2
4
6
8
10
12
14
14 18 22 26 30 34 38 42 46 50 54 58 62
%
TL(cm)
N=1996
40
Fig. 30. Length frequency distribution of C. nufar in the commercial catchesduring 2006-2007
Fig. 31. Pooled length frequency distribution of C. nufar in the commercial catches
during 2005-2007
0
2
4
6
8
10
12
14
14 18 22 26 30 34 38 42 46 50 54 58 62
%
TL(cm)
N=2136
0
2
4
6
8
10
12
14
14 18 22 26 30 34 38 42 46 50 54 58 62
%
TL(cm)
N=4132
41
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)
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
MaleW = 0.018*L 2.907
(R2 = 0.986)
a: -0.033 – 0.071
b: 2.872 – 2.940
0.02650 0.01736
Female W = 0.020*L2.890
(R2 = 0.985)
a: - 0.028 – 0.068
b: 2.859 – 2.921
0.02439 0.01595
PooledW = 0.019*L2.897
(R2 = 0.985)
a: -0.016 – 0.055
b: 2.874- 2.920
0.01795 0.01174
FL = 0.892TL - 0.151R² = 0.991
10
20
30
40
50
60
15 25 35 45 55 65 75
FL (c
m)
TL (cm)
N= 1,300
42
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.
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.
Fig. 34. VBGF curve of C. nufar by SLCA technique
W = 0.019L2.897
R² = 0.985
0
1000
2000
3000
4000
10 20 30 40 50 60 70
Tota
l wei
ght (
g)
Total length (cm)
0 0 . 5 1 . 0 1 . 5 2 . 0 2 . 50
1 0
2 0
3 0
4 0
5 0
6 0
7 0
S a m p l e Ti m i n g
Le
ng
th C
las
s
G r o w th C u r v e = N o n S e a s o n a l . L i n f = 7 0 . 3 5 . K = 0 . 3 1 . Tz e r o = -0 . 2 3 .
43
Fig. 35. VBGF curve of C. nufar by PROJMAT technique
Fig. 36. VBGF curve of C. nufar by ELEFAN 1 technique
0 0.5 1.0 1.5 2.0 2.50
10
20
30
40
50
60
70
Sample Timing
Leng
th C
lass
Growth Curve = Non Seasonal. Linf = 68.26. K = 0.25. Tzero = -0.55.
0 0.5 1.0 1.5 2.0 2.50
10
20
30
40
50
60
70
Sample Timing
Leng
th C
lass
Growth Curve = Non Seasonal. Linf = 70.97. K = 0.30. Tzero = -0.18.
44
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
Method L∞ (cm) K (y-1) 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.
45
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.
0 0.5 1.0 1.5 2.0 2.50
2
4
6
8
Sample Timing
Nom
inal
Age
Age Slice
46
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.
Fig. 39. Length at first capture of C. nufar
15
25
35
45
55
65
75
1 2 3 4 5 6 7 8 9 10 11
Tot
al le
ngth
(cm
)
Age (y)
SLCAPROJMATELEFANAverage
0102030405060708090
100
15 20 25 30 35 40 45 50 55 60 65 70
Prob
ablit
y of
cap
ture
TL(cm)
Lc=31.4 cm
47
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.
48
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).
0
10
20
30
40
50
60
70
80
90
100
J F M A M J J A S O N D
%
Months
Opaque
Hyaline
49
Table 5. Growth parameters of male and female C. nufar using otolith reading.
Sex L∞ K to r2
Male 66.47 0.1939 -0.36055 0.89652
Female 69.77 0.182 -0.24886 0.92528
Fig. 42. VBGF curves for C. nufar, Female (n =159) and Male (n =99).
0
10
20
30
40
50
60
70
0 2 4 6 8 10 12 14
Size
(cm
)
Age(y)
Male - Data
Female - Data
M - VBG
F-VBG
50
Table 6 A. Age - length key for Male of C. nufar - sectioned otolith.
TL(cm)Age (y)
Total1 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
Total 12 12 16 18 13 6 3 5 4 2 2 4 2 99
51
Table 6 B. Age - length key for female of C. nufar - with sectioned otolith.
TL(cm)Age (y)
Total1 2 3 4 5 6 7 8 9 10 11 12 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 2 1 3 1 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
52
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 3 2 4 1 9
60 3 6 2 2 3 3 19
64 1 5 5 4 14
Total 19 102 767 730 189 35 30 20 12 11 15 14 8 1951
53
Table 6 D. Age structure of C. nufar population in the Arabian Sea
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.
TL(cm) Age (y) Female Total
1 2 3 4 5 6 7 8 9 10 11 12 13
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 2 11 3 3 1 3 3 24
56 3 2 4 1 11
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
54
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
55
to attain ripe stage for spawning. Hence, it appears that individual fish would spawn
more than once during the spawning season in Arabian Sea.
Fig. 43 A. Development of ova to maturity stage I in C. nufar
Fig. 43 B. Development of ova to maturity stage II in C. nufar
0
25
50
75
1000.
014-
0.04
20.
056-
0.08
40.
098-
0.12
60.
140-
0.16
80.
182-
0.21
00.
224-
0.25
20.
266-
0.29
40.
308-
0.33
60.
350-
0.37
80.
392-
0.42
00.
434-
0.46
20.
476-
0.50
40.
518-
0.54
60.
560-
0.58
80.
602-
0.63
00.
644-
0.67
20.
686-
0.71
40.
728-
0.75
60.
770-
0.79
8
%
Ova diameter (mm)
A I
0
20
40
60
0.01
4-0.
042
0.05
6-0.
084
0.09
8-0.
126
0.14
0-0.
168
0.18
2-0.
210
0.22
4-0.
252
0.26
6-0.
294
0.30
8-0.
336
0.35
0-0.
378
0.39
2-0.
420
0.43
4-0.
462
0.47
6-0.
504
0.51
8-0.
546
0.56
0-0.
588
0.60
2-0.
630
0.64
4-0.
672
0.68
6-0.
714
0.72
8-0.
756
0.77
0-0.
798
%
Ova diameter (mm)
B II
56
Fig. 43 C. Development of ova to maturity stage III in C. nufar
Fig. 43 D. Development of ova to maturity stage IV in C. nufar
Fig. 43 F. Development of ova to maturity stage V in C. nufar
0
10
20
30
0.01
4-0.
042
0.05
6-0.
084
0.09
8-0.
126
0.14
0-0.
168
0.18
2-0.
210
0.22
4-0.
252
0.26
6-0.
294
0.30
8-0.
336
0.35
0-0.
378
0.39
2-0.
420
0.43
4-0.
462
0.47
6-0.
504
0.51
8-0.
546
0.56
0-0.
588
0.60
2-0.
630
0.64
4-0.
672
0.68
6-0.
714
0.72
8-0.
756
0.77
0-0.
798
%
Ova diamater (mm)
C III
0
5
10
15
20
0.01
4-0.
042
0.05
6-0.
084
0.09
8-0.
126
0.14
0-0.
168
0.18
2-0.
210
0.22
4-0.
252
0.26
6-0.
294
0.30
8-0.
336
0.35
0-0.
378
0.39
2-0.
420
0.43
4-0.
462
0.47
6-0.
504
0.51
8-0.
546
0.56
0-0.
588
0.60
2-0.
630
0.64
4-0.
672
0.68
6-0.
714
0.72
8-0.
756
0.77
0-0.
798
%
Ovadiameter (mm)
D IV
0
5
10
15
0.01
4-0.
042
0.05
6-0.
084
0.09
8-0.
126
0.14
0-0.
168
0.18
2-0.
210
0.22
4-0.
252
0.26
6-0.
294
0.30
8-0.
336
0.35
0-0.
378
0.39
2-0.
420
0.43
4-0.
462
0.47
6-0.
504
0.51
8-0.
546
0.56
0-0.
588
0.60
2-0.
630
0.64
4-0.
672
0.68
6-0.
714
0.72
8-0.
756
0.77
0-0.
798
0.81
2-0.
840
%
Ova diameter (mm)
F V
57
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).
58
Fig. 44. Distribution of different maturity stages of gonads of C. nufar
during 2005-06
0
25
50
75
I II III IV V VI
Perc
enta
ge M (n=19)F (n=24)
Apr 05
0
25
50
I II III IV V VI
Perc
enta
ge
M (n=29)F (n=16)
May
0
25
50
I II III IV V VI
Perc
enta
ge M (n=35)F (n=46)
June
0204060
I II III IV V VI
Perc
enta
ge M (n=35)F (n=48)
July
020406080
I II III IV V VI
Perc
enta
ge M (n=33)F (n=42)
August
0
20
40
I II III IV V VI
Perc
enta
ge
Maturity stages
M (n=38)F (n=34) September
0
20
40
60
I II III IV V VI
Perc
enta
ge M (n=17)F (n=24)
October
020406080
I II III IV V VI
Perc
enta
ge
M (n=32)F (n=47)
November
0
50
100
I II III IV V VI
Perc
enta
ge M (n=31)F (n=40)
December
0
50
100
I II III IV V VI
Perc
enta
ge
M (n=31)F (n=40)
Jan 06
0
20
40
60
I II III IV V VI
Perc
enta
ge
M (n=22)F (n=24)
February
0
50
100
I II III IV V VI
Perc
enta
ge
Maturity stages
M (n=25)F (n=25)
March
59
Fig. 45. Distribution of different stages of gonads of C. nufar during 2006-07
0255075
100
I II III IV V VI
Perc
enta
ge M (n=24)F (n=16)
April 06
0
20
40
60
I II III IV V VI
Perc
enta
ge
M (n=20)F (n=20)
May
0
20
40
60
I II III IV V VI
Perc
enta
ge
M (n=55)F (n=65)
June
0
20
40
60
I II III IV V VI
Perc
enta
ge
M (n=16)F (n=24) July
0
20
40
60
I II III IV V VI
Perc
enta
ge
M (n=20)F (n=33)
August
0
20
40
60
I II III IV V VI
Perc
enta
ge
Maturity stages
M (n=20)F (n=22) September
0
20
40
60
I II III IV V VI
Perc
enta
ge M (n=30)F (n=21)
October
0204060
I II III IV V VIPe
rcen
tage
M (n=19)F (n=22)
November
020406080
I II III IV V VI
Perc
enta
geM (n=19)F (n=21)
December
0
50
100
I II III IV V VI
Perc
enta
ge M (n=12)F (n=28)
Jan 07
0
50
100
I II III IV V VI
Perc
enta
ge
M (n=26)F (n=12)
February
0
50
100
I II III IV V VI
Perc
enta
ge
Maturity stages
M (n=19)F (n=19)
March
60
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
61
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).
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.
22
23
24
25
26
27
28
29
30
0
0.5
1
1.5
2
2.5
A M J J A S O N D J F M
GSI
Months
Female
Male
Temp
62
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.
0.6
0.8
1
1.2
1.4
1.6
1.8
April
May
June July
Augu
st
Sept
embe
r
Oct
ober
Nov
embe
r
Dece
mbe
r
Janu
ary
Febr
uary
Mar
ch
HSI
Female
Male
63
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
0.9
0.95
1
1.05
1.1
1.15
1.2
1.25
1.3
1.35
A M J J A S O N D J F M
Kn
Months
FemaleMale
64
Fig. 51 Juveniles distribution: Autumn monsoon 2007
Fig. 52 Juveniles distribution –NE monsoon 2008
65
Fig. 53 Juveniles distribution Spring Inter-monsoon 2008
Fig. 54 Juveniles distribution -Post SW monsoon 2008
66
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.
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)
0
0.2
0.4
0.6
0.8
1
10 15 20 25 30 35 40 45 50 55 60 65
Prop
ortio
n m
atur
e
Total length (cm)
MaleMaleFemaleFemale
Male: Lm = 33.7 cmFemale: Lm = 31.9 cm
67
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.
Fig. 56 A. Fecundity in relation to total length (cm) of C. nufar.
y = 0.0061x4.3232
R² = 0.8977
0
50
100
150
200
250
300
350
400
450
500
20 30 40 50 60 70
Fecu
dity
('00
0)
Total length(cm)
A
68
Fig. 56 B. Fecundity in relation to body weight(g) of C. nufar .
Fig. 56 C. Fecundity in relation to ovary weight (g) of C. nufar
y = 4.3082x1.3938
R² = 0.9085
0
50
100
150
200
250
300
350
400
450
500
0 500 1000 1500 2000 2500 3000 3500 4000
Fecn
dity
('00
0)
Total Weight(g)
B
y = 3074.1x0.9857
R² = 0.9921
0
100
200
300
400
500
600
0 50 100 150 200
Fecu
ndity
('00
0)
Ovary Weight (g)
C
69
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).
Fig. 57. Sex ratio of C. nufar during the period April 2005 - March 2007
0
10
20
30
40
50
60
70
80
Apr-
05M
ayJu
ne July
Aug
Sep
Oct
Nov De
cJa
n-06 Fe
bM
ar Apr
May
June July
Aug
Sep
Oct
Nov De
cJa
n-07 Fe
bM
ar
%
Months
Male
Female
70
Table. 7A Sex-ratio in C. nufar during 2005-06
M h
Total No.
f fi h
No. of
lNo. of females Male: Female Chi-square value
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
71
Table. 7B Sex-ratio in C. nufar during 2006 -07
*Significant at 5% level
Month Total No.
f fi h
No. of
lNo. of females Male: Female Chi-square value
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
72
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.
Fig. 58. General food composition of C. nufar during 2005-06
50
2.382.781.98
42.86
Fish
Crustaceans
Cephalopods
Polychaete worms
Semidigested matter
73
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).
57.96
1.331.33
39.39
Fish
Crustaceans
Cephalopods
Semidigested matter
74
Fig. 60. Monthly percentage of composition of food items of C. nufar
during 2005-06
Fig. 61. Monthly composition of food items of C. nufar during 2006-07
60
25
42.11
65.9563.01
40
52.94
26.67
55.56
16.67 20
4.26 5.8811.11
6.6710.53 6.38 6.67 5.56
40
75
47.37
23.430.14
53.33
41.18
73.33
27.78
83.33
100
73.33
0
20
40
60
80
100
Apr 0
5
May
June July
Augu
st
Sept
embe
r
Oct
ober
Nov
embe
r
Dece
mbe
r
Jan
06
Febr
uary
Mar
ch
% c
ompo
sitio
n
Months
N=775FishCrustaceansCephalopodsPolychaete wormsSemidigested matter
11.11
36.8445
55.5556.75
73.0883.79
55.55
90
33.33
63.64
33.34
8.11 7.69 9.09
88.89
63.16 55
44.4435.14
19.23 16.22
44.44
10
66.67
27.27
66.67
0
20
40
60
80
100
Apr 0
6
May
June July
Augu
st
Sept
embe
r
Oct
ober
Nov
embe
r
Dece
mbe
r
Jan
07
Febr
uary
Mar
ch
% C
ompo
sitio
n
Months
N=550FishCrustaceansCephalopodsSemidigested matter
75
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.
Fig. 62. Food items found in the gut of various size groups of C. nufar
during 2005-2007 (pooled)
53.8559.46
48.8456.06
60.8765.86
71.43
50
75
50
15.39
2.7 2.3305.41 4.65 0.76 1.45
6.25
30.77 32.43
43.41 41.6736.23
31.71 28.57
43.75
25
50
0
20
40
60
80
100
16-2
0
21-2
5
26-3
0
31-3
5
36-4
0
41-4
5
46-5
0
51-5
5
56-6
0
61-6
5
% C
ompo
sitio
n
Size groups (TL-cm)
N=1325FishCrustaceansCephalopodsPolychaetesSemidigested matter
76
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.
Fig. 63. Feeding intensity in C. nufar relation to months
75
43
80
81
61
48
52
41
40
80
40
53
42
5141
41
41
40
40
0
20
40
60
80
100
Sept
05
Oct
Nov De
cJa
n 06 Fe
bM
arch Ap
rM
ay Jun Jul
Aug
Sept Oct
Nov De
cJa
n 07 Fe
bM
arch
Feed
ing
inte
nsity
(%)
Months
Empty1/4 full1/2 full3/4 fullFull
N=990
77
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.
Fig. 64. Feeding intensity in different maturity stages of C. nufar
7.23 9.3517.65 16.85 22.22 16.91 22.22 17.39
30.564.01 0.93
7.35 2.8114.29
5.155.56
4.3524.24
14.28
2.78
12.04 10.28
11.03 9.55
11.11
10.29
22.2217.39
25
24.2428.57
13.89
0
20
40
60
80
100
MI
F MII
F MIII
F MIV
F MV
F MVI
F
Feed
ing
inte
nsity
(%)
Maturity stages
Empty1/4 full1/2 full3/4 fullFull
M= male(n=477)
F= female(n=513)
78
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%).
Fig. 65. Feeding intensity in different size groups of C. nufar
18.42 12.03 11.62 13.3623.47
13.8925 22.22
12.5
66.67
2.633.01 2.14
1.02 2.27
100
2.63
3.01 5.5 8.91
1.02
2.78
2.27 11.11
12.5
33.33
10.53
11.28 13.46 13.36
10.2
8.33
4.55 11.1112.5
100
65.79
70.68
67.28
64.37
64.29
75
65.91
55.56
62.5
0
20
40
60
80
100
16-2
0
21-2
5
26-3
0
31-3
5
36-4
0
41-4
5
46-5
0
51-5
5
56-6
0
61-6
5
66-7
0
71-7
5
Feed
ing
inte
nsity
(%)
Length class (cm)
Empty1/4 full1/2 full3/4 fullFull
N=990
79
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.
80
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).
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.
0
2000
4000
6000
8000
10000
12000
14000
16000
18000
0
500
1000
1500
2000
2500
0 1 2 3 4
Biomass (t)Yi
eld
(t)
F-factor
Yield (t)Biomass (t)
MSY
81
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.
0
100
200
300
400
500
600
700
800
0 0.4 0.8 1.2 1.6 2 2.4
Yiel
d (g
)
Fishing mortality
YW/R (g)
SSB/R (g)
TB/R (g)
82
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 i t occur at 26–28ºC. During Post southwest monsoon 2008
the fish was found at temperature between 17ºC and 21ºC (Fig. 68).
Fig.68. Catch of C. nufar at di fferent t ime of monsoon.
0
20
40
60
80
100
120
140
15 16 17 18 19 20 21 22 23 24 25 26 27 28 29
Catc
h (k
g)
Temperature ºC
Post SW 2007
Autumn monsoon 2007
Northeast Monsoon 2008
Spring monsoon 2008
Post SW 2008
83
Fig. 69 Distribution of C. nufar: Post SW monsoon 2007
Fig. 70 Distribution of C. nufar : Autumn monsoon 2007
84
Fig. 71 Distribution of C. nufar: NE monsoon 2008
Fig. 72 Distribution of C. nufar: Spring Inter-monsoon 2008
85
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).
86
Fig.74 Catch rate of C. nufar by stratum during monsoon periods
Fig.75. Occurrence of C. nufar at different level of bottom dissolved oxygen.
0200400600800
1000120014001600180020002200
Post
SW
200
7
Autu
mm
onso
on20
07
NE
mon
soon
2008 Sp
rinm
onso
on20
08
Post
SW
200
8
Kg. k
g-2
20-50 m
50 - 100 m
100 - 150 m
150 - 250 m
0
20
40
60
80
100
120
140
0 1 2 3 4 5
Catc
h(Kg
)
Dissolved O2 ml l-1
Post SW 2007
Autumn monsoon 2007
Northeast Monsoon2008Spring monsoon 2008
87
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
88
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 S e a 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
the region where oxygen is less than 0.1 ml l-1
. .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.,
89
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 l imi tat ions 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 f i sh 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.
90
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
Country L∞ (TL; cm) K y-1 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
91
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
92
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
93
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
94
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).
95
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
96
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.
97
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110
Ann
ex.1
. Cla
ssifi
catio
n of
mat
urity
stag
es in
C. n
ufar
Stag
e of
mat
urity
Mal
eFe
mal
ePo
sitio
n of
last
mod
e an
dm
axim
um d
iam
eter
of
ova
(in p
aren
thes
es) i
nm
.d. (
1 m
.d. =
0.0
14m
m)
Nat
ure
and
exte
nt o
f tes
tis in
body
cav
ityN
atur
e an
d ex
tent
of o
vary
inbo
dy c
avity
App
eara
nce
of o
va u
nder
mic
rosc
ope
I-Im
mat
ure
Smal
l, tra
nspa
rent
, pal
e,oc
cupy
ing
a ve
ry sm
all
porti
onto
1/3
of b
ody
cavi
ty
Smal
l, tra
nspa
rent
, pal
e,oc
cupy
ing
a ve
ry sm
all p
ortio
nto
1/3
of b
ody
cavi
ty, o
va n
otvi
sibl
e to
nak
ed e
ye
Irre
gula
r, sm
all,
yolk
less
/yol
kde
posi
t jus
t sta
rted,
tran
spar
ent
with
cle
arly
vis
ible
/par
tially
visi
ble
nucl
eus
1-3
(9)
II-M
atur
ing
1W
hitis
h, tr
ansl
ucen
t,oc
cupy
ing
abou
t 1/2
of b
ody
cavi
ty
Pale
yel
low
, gra
nula
r ova
vis
ible
to n
aked
eye
, occ
upyi
ng a
bout
1/2
of b
ody
cavi
ty
Med
ium
size
d, a
ssum
e ro
und
shap
e, o
paqu
e, w
ith fa
iram
ount
of y
olk
16-1
8 (2
4)
III-
Mat
urin
g 2
Cre
amy
whi
te, o
ccup
y ab
out
3/4
of b
ody
cavi
tyPa
le y
ello
wis
h, b
lood
ves
sels
visi
ble
on d
orsa
l sid
e, o
va c
lear
lyvi
sibl
e, o
ccup
ying
abo
ut 3
/4 o
fbo
dy c
avity
Med
ium
size
d, o
paqu
e, fu
llyyo
lked
25-2
7 (3
3)
IV-M
atur
eC
ream
y w
hite
, sof
t,oc
cupy
ing
abou
t ful
l len
gth
of b
ody
cavi
ty
Pink
ish
yello
w, b
lood
ves
sels
prom
inen
t, la
rge
ova
prom
inen
tly v
isib
le,
occu
pyin
gab
out f
ull l
engt
h of
bod
y ca
vity
Larg
e si
zed,
mat
ure,
trans
pare
nt a
t per
iphe
ry31
-33
(39)
V-R
ipe/
Run
ing
Cre
amy
whi
te, o
ccup
ying
full
leng
th o
f bod
y ca
vity
, milt
exud
ed o
n sl
ight
pre
ssur
e
Pink
ish
yello
w, o
ccup
ying
full
leng
th o
f bod
y ca
vity
, ova
rian
wal
l thi
n, la
rge
ova
sepa
rate
dfr
om e
ach
othe
r and
ooz
ed o
utun
der s
light
pre
ssur
e
Eggs
sphe
rical
, mea
sure
d 0.
77m
m-0
.84
mm
in d
iam
eter
, asi
ngle
oil-
glob
ule
and
narr
owpe
rivite
lline
spac
e pr
esen
t
55-5
7 (6
0)
VI-
Spen
tFl
abby
, litt
le re
ddis
h, o
ccup
ying
abou
t 1/2
of b
ody
cavi
ty.
Flac
cid,
redd
ish,
occ
upyi
ngab
out 1
/2 o
f bod
y ca
vity
Med
ium
size
d ov
a pr
esen
t with
disi
nteg
ratin
g rip
e ov
a19
-21
(-)
111
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)
RipeOvary 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.
112
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