(Qeshm Island) Using Ecopath with Ecosim

Modelling trophic structure and energy flows in the coastal ecosystem of the Persian Gulf using Ecopath with Ecosim

Item Type thesis

Authors Hakimelahi, Maryam

Publisher Khorramshahr University of Marine Science and Technology, Faculty of Marine Science and Oceanography, Department of Marine Biology

Download date 30/09/2021 15:54:43

Link to Item http://hdl.handle.net/1834/35825

Khorramshahr University of Marine Science and Technology Faculty of Marine Science and Oceanography Department of Marine Biology

Ph.D. Thesis

Modelling trophic structure and energy flows in the coastal ecosystem of the Persian Gulf (Qeshm Island) using Ecopath with Ecosim

Supervisors: Dr. Ahmad Savari Dr. Babak Doustshenas

Advisors: Dr. Mehdi Ghodrati Shojaei Dr. Kristy A.Lewis

By: Maryam Hakimelahi

June 2018

167

Abstract In the present study, the trophic structure for some species of the coastal ecosystem of south of the Qeshm Island was developed using the mass balance modeling software Ecopath (Version 6.5.1). In this model, 33 functional groups including fish, benthos, phytoplankton, zooplankton, seaweed and detritus were simulated. In total, 3757 samples of stomach contents were analyzed based on the weight and numerical methods. Bony fish and crustacean were found to be the main prey in most of the stomach contents. The mean trophic level in the study area was calculated to be 3.08. The lowest trophic level was belonged to Liza klunzingeri, (2.50) and the highest belong to Trichiurus lepturus (4.45). The range of total mortality varied from 1.11 per year for T. Lepturus to 3.55 per year for Sillago sihama. The lowest and highest Consumption/Biomass value was belonged to Trichiurus lepturus (2.94) and Rastrelliger kanagurta (11.60) respectively. In this study, the role of ecological groups in the food web was determined using Keystone Index. The results showed that benthos and subsequently phytoplankton and zooplankton are among the dominant ecological groups in the southern ecosystem of Qeshm Island. The Connectance Index and the System Omnivory Index of this ecosystem were estimated to be 0.14 and 0.15 respectively. Population dynamics studies especially the exploitation rate (E) showed that Pomadasys kaakan, Rastrelliger kanagurta and Sillago sihama are under fishing pressure. Overall, the indices of ecological relationships with the population dynamics parameters showed that the exploitation of the fisheries resources in the study area is not in a good condition. The results of Ecosim model using time series of mean biomass from 2004-2016, showed a decrease in the biomass of Nemipterus japonicus, Otolithes ruber and Rastrelliger kanagurta. In general, the results of this study showed that the current fish harvest on the population of carnivorous species is very high which in long term could decrease the number of fish recruitment and consequently the capacity and sustainability of Qeshm Island fishing. Moreover, the high exploitation of fishing resources in this area could have negative effect on top-down food web and gradually reduce the top predator species, and eventually resulted in changing the trophic level of the species and reduction of the top of the food web species, which could be resulted in damaging of the ecological balance and endangering long-term sustainable utilization.

Keywords: Population dynamics, Trophic levels, Food web, Keystone Index, Connectance Index, Ecopath with Ecosim, Persian Gulf.

166

( )

1397

. . .

33 . . . 3757

3/08 (Trophic level) . 2/50 (Liza klunzingeri) . 4/45 (Trichiurus lepturus) 1/11 (Q/B) . 3/55 Rastrelliger ) 2/94 (Trichiurus lepturus) . 11/60 (kanagurta 1 (Keystone Index) 1 . 0/89 0/94 System ) (Connectance Index) . . 0/15 0/14 (Omnivory Index Pomadasys ) (E=0/6) (Sillago sihama) (E=0/6) (Rastrelliger kanagurta) (E=0/7) (kaakan . 2004 - 2016 Rastrelliger ) (Otolithes ruber) (Acanthopagrus latus) . (kanagurta -

- : .

1 ...... :

1...... 1 - 1 4...... 1 - 1 - 1 7...... 2 - 1 7...... 1 - 2 - 1 8...... 2 - 2 - 1 9...... 3 - 2 - 1 10 ...... 4 - 2 - 1 11 ...... 5 - 2 - 1

15 ...... :

24 ...... :

23 ...... 1 - 3 25 ...... 2 - 3 26 ...... 3 - 3 27 ...... 4 - 3 27 ...... 5 - 3 27 ...... 1 - 5 - 3 28 ...... ((K) (L ) ) 2 - 5 - 3 29 ...... ( ´ ) 3 - 5 - 3 30 ...... 4 - 5 - 3 31 ...... ( Z ) 1 - 4 - 5 - 3 31 ...... ( M ) 2 - 4 - 5 - 3 32 ...... ( F ) 3 - 4 - 5 - 3 32 ...... ( E ) 4 - 4 - 5 - 3 32 ...... 6 - 3 32 ...... 1 - 6 - 3 32 ...... (VI) 1 - 1 - 6 - 3 33 ...... ( FP) 2 - 1 - 6 - 3

33 ...... (WI) 2 - 6 - 3 34 ...... 7 - 3 34 ...... 8 - 3 35 ...... 9 - 3 35 ...... 1 - 9 - 3 35 ...... 2 - 9 - 3 35 ...... (P/B) / 3 - 9 - 3 36 ...... (Q/B) / 4 - 9 - 3 36 ...... (DC) 5 - 9 - 3 37 ...... 10 - 3 37 ...... 11 - 3 38 ...... 12 - 3 38 ...... 13 - 3 38 ...... (TL) 1 - 13 - 3 39 ...... (EE) 2 - 13 - 3 39 ...... 3 - 13 - 3 40 ...... ( OI) 4 - 13 - 3 40 ...... ( E ) 5 - 13 - 3 40 ...... ( MTI) 6 - 13 - 3 41 ...... (OI) 7 - 13 - 3 41 ...... Aspect Ratio 8 - 13 - 3 42 ...... (CI) 9 - 13 - 3 43 ...... 10 - 13 - 3 43 ...... 14 - 3

45 ...... :

45 ...... 1 - 4 45 ...... (Trichiurus lepturus) 1 - 1 - 4 54 ...... (Sillago sihama ) 2 - 1 - 4 61 ...... (Rastrelliger kanagurta ) 3 - 1 - 4 69 ...... (Liza klunzingeri) 4 - 1 - 4

74 ...... (Acanthopagrus latus) 5 - 1 - 4 80 ...... (Pomadasys kaakan) 6 - 1 - 4 86 ...... 2 - 4 88 ...... 3 - 4 88 ...... 1 - 3 - 4 89 ...... 2 - 3 - 4 91 ...... 3 - 3 - 4 93 ...... 4 - 3 - 4 95 ...... 5 - 3 - 4 97 ...... 6 - 3 - 4 99 ...... 7 - 3 - 4 100 ...... 8 - 3 - 4 101 ...... 9 - 3 - 4 102 ...... 10 - 3 - 4 102 ...... 4 - 4

104...... :

104 ...... 1 - 5 117 ...... 2 - 5 123 ...... 3 - 5 123 ...... 4 - 5

124 ......

126 ......

24 ...... 1-3 -1396) 1-4 46 ...... (1395 -1396) 2-4 48 ...... (1395 (%W) 3-4 51 ...... ( 1395-1396) (%N) (%F) 4-4 53 ...... (1395 - 1396) 54 ...... (1395-1396) 5-4 56 .... (1395 - 1396) 6-4 (1395-1396) 7-4 58 ...... 61 ...... (1395-1396) 8-4 63 ..... (1395-1396) 9-4 65 ...... (1395 - 1396) 10-4 69 ...... ( 1395-1396) 11-4 71 ... (1395-1396) 12-4 73 ...... (1395 - 1396) 13-4 (1395-1396) 14-4 74 ...... -1396) 15-4 76 ...... (1395 (%W) 16-4 79 ...... (1395-1396)

(%N) (%F) 17-4 80 ...... (1395 - 1396) - 1396) 18-4 81 ...... (1395 - 1396) 19-4 82 ...... (1395 (%W) 20-4 85 ...... ( 1395-1396) (%N) (%F) 21-4 86 ...... ( 1395-1396) Q/B EE 22-4 88 ...... 90 ...... 23-4 92 ...... 24-4 100 ..... 25-4 102 ...... 26-4 1-5 106 ...... 2-5 107 ...... 112 . 3-5

12 ...... 1-1 13 ...... 2-1 14 ...... 3-1 25 ...... 1-3 26 ...... 2-3 42 ...... Aspect Ratio 3-3 1-4 46 ...... (1395-1396) 47 . (1395-1396) 2-4 3-4 49 ...... ( 1395-1396) 4-4 49 ...... (1395-1396) 5-4 50 ...... (1395 - 1396) - 1396) 6-4 51 ...... (1395 55 . (1395-1396) 7-4 56 ...... (1395-1396) 8-4 -1396) 9-4 57 ...... (1395 ( 11 - 16/90) 10-4 60 ...... (1395 - 1396) ( 17 - 23/90) 11-4 60 ...... (1395 - 1396)

62 .( 1395-1396) 12-4 63 ...... (1395-1396) 13-4 -1396) 14-4 64 ...... (1395 15-4 65 ...... (1395-1396) 16-4 68 ...... (1395-1396) 17-4 68 ...... (1395-1396) 70 ...... ( 1395-1396) 18-4 71 ...... (1395 - 1396) 19-4 -1396) 20-4 72 ...... (1395 21-4 73 ...... (1395-1396) 75 . (1395-1396) 22-4 76 ...... (1395 - 1396) 23-4 24-4 77 ...... (1395-1396) (1395-1396) 25-4 78 ...... - 1396) 26-4 78 ...... (1395 -1396) 27-4 81 ...... (1395

82 .. (1395-1396) 28-4 29-4 83 ...... (1395-1396) (1395 - 1396) 30-4 84 ...... 31-4 84 ...... (1395-1396) 87 ...... 32-4 33-4 89 ...... 34-4 94 ...... 96 ... 35-4 36-4 98 ...... 99 ...... 37-4 38-4 101 ...... ( ) ( ) 39-4 103 ...... (2004-2016) 118 ...... 1-5

: 1 - 1

.(Lotze et al., 2006; Halpern et al., 2015) (Jackson et al., 2001) .(Worm et al., 2006)

. Pikitch et al., ) .(2004 Browman et al., ) - .(2005a, 2005b

.(Cury and Christensen, 2004)

.(Rombouts et al., 2013)

.(Fulton et al., 2005)

Blanchard et al., 2010; ) - .(Kleisner et al., 2013; Large et al., 2013 - .(Boldt et al., 2014; Large et al., 2015; Levin et al., 2009)

Allesina and Bondavalli, ) .(2004; Valentini and Jordan, 2010

.(Piroddi et al., 2015; Smith et al., 2015) 1 EwE - EwE .(Christensen and Walters, 2004) . Coll et al., ) 400 .(2015 EwE .(Coll and Libralato, 2012; Heymans et al., 2014)

.(Cerco et al., 2010) . 200 10 - 2 .

Ecopath with Ecosim Ecosystem-Based Management

. .(Pikitch et al., 2004) - ( ) . .(Hoese et al., 2006) (1998) Pauly .

0/1 . .

( ) .

) .(1385

. .

. . - 1

. -

... 2 . 5 4 3

7 6

...... 1 - 1 - 1

. . . .

Modelling Trophic Level Omnivory Index Mixed Trophic Impact Electivity Index Overlap Index Connectance Index

. .

Christensen and Pauly, 1993; Nasir, ) ) ( ) .(2000 .(Myers and Worm, 2003) (

.(Pauly et al., 2000)

- .

Whipple et ) .(Magnússon, 1999) . .(al., 2000

. .

.(Scharf and Schlicht, 2000)

. .(Magnússon, 1999) .

- . . : Christensen et al., ) + + + = .(2000

.(Lauria, 2010)

- .(Lauria et al., 2012)

. .

. . .

( )

- . - .

2 - 1 1 - 2 - 1 2 1

. 1491

Marginal Sea Semi-enclosed Neritic

.(1384 )

2 - 2 - 1 . 1

- .(Deus and Petrere-Junior, 2003)

. ... . - . . - ) . (Euphausiids .(Laevastu and Hayes, 1981)

Trophodynamic

. .(Laevastu and Hayes, 1981) (... )

. . . . : - 1 . : - 2 . . .

: -3 . 137 - 32- ( ) .

.(Biswas, 1993)

.(Amundsen et al., 1996)

3 - 2 - 1

. .(Christensen and Pauly, 1998) 2 .

3 . 4 . .(Biswas, 1993)

4 - 2 - 1 -

. . .

.(Pauly and Christensen, 2000)

. 1950 .

- .

Trophic Level Herbivores Carnivores Omnivores

. . . 1

.(Christensen and Pauly, 1998) ( )

2 .

.(Christensen and Pauly, 1998)

. .

5 - 2 - 1 500 164 6000 Ecospace 3EWE ( 120) Ecopath . - ) (Polovin, 1984; Christensen and Pauly, 1992 ) ( .(Walters et al., 1997)

Virtual Population Analysis (MSVPA) Trophic Level Ecopath with Ecosim

EWE . (Walters and Martell, 2004) 1 - 1 . . .

.(Chen et al., 2011) 1-1

- 2-1 . .

2-1

.( 3-1 )

Mass-Balance

3-1

.(Pauly et al., 2000) - 2 - 1 - - 3 - 4 .

.(Pauly et al., 1998, 2000) ( )

. . Polovina (1984) . Ulanowicz (1986)

. Abdurahiman et al., 2010; Pauly et al., ) .(2000; Nascimento et al., 2012 - .

- .

(2014) Taghavimotlagh .1 . . (2016) Razzaghi .2 2011 2002

. . (2017) Behzadi .3 4/32 (Rachycentron canadum) . 6/13 (Q/B) (Himantura gerrardi) (2017) Rastgoo .4 . 4 (2017) Navarro Rastgoo .5

. .

. (1984) Polovina .6 8 . P/B 25 . (1992) Pauly Christensen .7 . -

(1996) Christensen .8

. (1997) Walters .9 . . (1998) Christensen .10

. . (2000) Shannon .11 . . ´Caete (2000) Wolff .12 .

%10 %10 %13 (fiddler) %19 %34 . %14 14 (2001) Arregu´n-Sa´nchez Vega-Cendejas .13 - . . %7 %4 %64 . )

22 (EE ) . ( Q/B P/B . . 4 (2002) Wolff Ortiz .14 . (MTI23) - Chondracanthus chamissoi Argopecten purpuratus - Malacoglanis gelatinosus . Huizache (2003) Zetina-Rejon .15 2 . Caimanero . 3/6 . . (2003) Alias .16 . . (2003) Vivekanandan .17 . . . San Pedro (2003) Campos .18 . . 3/25

Ecotrophic Efficiency Mixed Trophic Impact 26

Sarawak Sabah (2003) Garces .19 3/3 (TL24) . . Kuosheng (2004) Lin .20 . . Kuosheng %96 (2005) Velasco .21 . . (2006) Coll .22 4 . anglerfish . Hake - . . . (2007) Sandberg .23 . . ( ) . Kivu (2008) Villanueva .24 . . . 9/4 4/5 (TL)

Trophic Level 27

- . - (2009) Panikkar Feroz Khan .25 . 4 . 3/57 0/402 0/422 Pearl spots Nile . . (2011) He Chen .26 . Hangzhou . (O) . (2012) Bundy Araújo .27 -1 . Scotian Shelf - 2 . ( ) -3 . . (2013) Tecchio .28 .

. (2014) Heymans .29 - .

(... ) . (2015) Chaalali .30 .

. (2016) Angelini .31 . . (2017) Dutta .32 . (Sundarban ) . 15 . 0/961 0/424 (EE) 2/716 Penaeid prawn .

. .

. (2017) Bornatowski .33 . .

(2017) Mutsert .34 . . - (2017) Piroddi .35 .

. (2017) Sagarese .36

. lionfish (Pterois volitans) (2017) Bumbeer .37 - . lionfish - 2 . - 1 . . . - lionfish -1 . . -2 lionfish (2017) OFarrell .38

(2017) Serpetti .39

. (2017) Shirakihara Sakamoto .40

. .

1 - 3 (N 26° 55 E 56° 10 ) . 1491 . .(1 - 3 ) . (1396 1395 ) . 1 - 3

1-3

N 26° 58 37,76 E 56° 14 37,50 N 26° 46 21,31 E 56° 03 35,03 N 26° 41 03,39 E 55° 42 09,63 N 26° 38 02,24 E 55° 16 32,09

1-3

2 - 3 6 (Sillago sihama) Trichiuridae (Trichiurus lepturus) (Liza klunzingeri) Scombridae (Rastrelliger kanagurta) Sillaginidae Sparidae (Acanthopagrus latus) Mugilidae .(2-3 ) Haemulidae (Pomadasys kaakan) -2 -1 . - 4 - 3

3-2

-3 3 - 01/0 01/0 - %70 %4 - - 6.5.1 EWE - 1.0.0 FiSAT II - 6 Primer -

. Excel 2010 -

4 - 3 6 3757 1396 1395 . . ( ) ) ( ) . ( . 0/01 70 4 .(Berg, 1979) Grethe (1997) Carpenter (2006 1989) Al-Yamani (1380) (2009) Al-kandari (1996) .(Hyslop, 1980)

5 - 3 1 - 5 - 3 . .(King, 1995 ) 1-3 b 1 - 3 = aL W b a L W b . .

.(Cadima, 2003) . .(Bertalanffy, 1938) (L ) x (W ) y 1

3 . (2 ) 2 b . 3/0 [ ] (King, 1996) (King, 1996) (Ln) (Le Cern, 1951) . 2 - 3 Log W = Ln a + bLn L 2 - 3 Ln W = Ln a + b(Ln Fl) Ln a Ln Fl Ln W . b . b Sparre ) 3 b 3 (Pauly, 1984b) t .(and Venema, 1998 .(Biswas, 1993) 2 .(Pauly, 1984b) 3 - 3 2 t = [(s.dx) / (s.dy)] * [(1b-31) / ( (1-r ))] * [ (n-2)] 3 - 3 s.dy s.dx 2 . n r b

((K) (L ) ) 2 - 5 - 3 L L .(Ingles & Pauly, 1984)

Isometric Cube Law 48

.(Gulland, 1991) . ( ) K L (Gulland, 1991) k . k (Pauly, 1984a) (Ingles & .(Pauly, 1980) Pauly, 1984). .(Sparre & Venema, 1998) k .(Cushing, 1981) M k . L

:1 - . K . . .

. . Rn (L K) Rn = 10 ESP/ ASP/ 10 . ESP ASP

( ´) 3 - 5 - 3 - ( ) k .(Munro and Pauly, 1983)

W 2/3 .

. W (k)

Munro and Pauly, ) (L ) .(1983 ´ . . 4 - 3

´= Log K + 2 Log L 4 - 3 ´ (1983) Munro Pauly . K (L ) .(Sparre and Venema, 1998) .(Pauly and Munro, 1984 ) ( L K) .(Pauly and Munro, 1984 )

4 - 5 - 3

. . .

(Z) 1 - 4 - 5 - 3

- FiSAT II .( Gayanilo and Pauly, 1997) . t = t0 1 / K Ln [ 1- ( Lt / L )]

: . Ln (f / dt) = a Z :f :dt :a :Z

( M ) 2 - 4 - 5 - 3

.(Sparre and Venema, 1998 ) . (5-3 ) 1980 . ( ) .(Pauly, 1980 ) 0/246 Log M = 0/0066 0/279 Log L + 0/06543 Log K + 0/4634 Log T 5 - 3 - L K M T . . 26/5

(F) 3 - 4 - 5 - 3 Sparre ) .(and Venema, 1998 51

(E) 4 - 4 - 5 - 3 .(Pauly, 1980) 0/5 E E > 0/5 .F = M .(Pauly, 1980 ) E < 0/5

6 - 3

.(Hyslop, 1980)

1 - 6 - 3 . 1 .(Hyslop, 1980)

(VI) 1 - 1 - 6 - 3 Euzen, 1987, Chrisafi et ) 2 . 6-3 .(al., 2007 VI= (ES / TS) × 100 6 - 3 4 3 TS ES VI 40 20 VI . 20 0 VI . . 60 40 VI .

Numerical Method Vacuity Index Empty Stomach Total Stomach 52

100 80 VI 80 60 VI .(Euzen, 1987)

(FP) 2 - 1 - 6 - 3 1 .(Euzen, 1987; Chrisafi et al., 2007) . 7 - 3 FP = (NSJ / NS) × 100 7 - 3 NS NSJ FP %50 .(Hyslop, 1980) %50 10 . . %10

(WI) 2 - 6 - 3 2 .(8 - 3 ) WI = (Wi / Wi) × 100 8 - 3 .(Hyslop, 1980) i Wi

7 - 3 - Christensen ) .(and Walters, 2004 . 9 - 3 .

Food Preference Index Gravimetric Method 53

B × i × EEi 9 - 3

EE i i,j Bi, Bj i Pi

(Q/B)j i (P/B)i i

j i DCji . j

. i ( ) EXi B, P/B, Q/B, EE . EE 4 3 .

8 - 3

(Lin et al., 2013) (... ) . . . .

9 - 3 1 1 - 9 - 3

( ) . ( ) .

Functional Group 54

- .(Pauly and Christensen, 2000) .

2 - 9 - 3

.(Tang et al., 2006) ( ) - .(Lin et al., 2013) a (1993) Pauly Christensen 2 gC/m .(Chen, 2004)

) .(1395 (P/B) / 3 - 9 - 3 (Allen, 1971) (Z) (P/B) . (Pauly, 1980) (M) Beverton and ) . (Holt, 1957

(Q/B) / 4 - 9 - 3

. .(Palomares and Pauly, 1989) 10 - 3 Log Q/B = 7.964 + 0.204 log W 1.965 T + 0.083 Ar + 0.532 h + 0.398 d 10 - 3

. W

1Ar .(Sparre and Venema, 1992) 2 h h / s Ar) h (. s d . . Q/B P/B .(Lin et al., 2004) Mohamed (2005) .

2 (DC) 5 - 9 - 3

.(Pauly and Christensen, 2000)

Palomares and Pauly, ) .(1989 .

.(Zhang et al., 2010)

Fauchald (2003) Hajisamae (2006) Grandcourt

Aspect Ratio Diet Composition 56

(2000) Pitcher Standford (2005) Mohamed (2006) Sabrah (1979) (1387) (1982) Khalaf Sirajul Islam (2007) Daskalov Mackinson .

10 - 3 EE EE 1 1 .

.(Christensen and Walters, 2004)

...... 11 - 3 .

. 1 .

12 - 3

2004 . ) 2016 .(1395

13 - 3

Trophic level 57

.(Libralato et al., 2006)

(TL) 1 - 13 - 3 . 11 - 3 .(Pauly and Christensen, 1997) 11 - 3

j TROPHj i j DCij . G . 5 3 - 2 . . 5 . 5 / - .(Pauly et al., 1998)

(EE) 2 - 13 - 3 1 . . (P/B) .(Christensen and Pauly, 1993) Z Z = M + F + OM .

Ecotrophic Efficiency 58

. OM F M Z P/B .(Christensen and Pauly, 1993) 12 - 3 EE = 1-OM / (P/B) 12 - 3

3 - 13 - 3

- 5 1 . . . 1 (OI) 4 - 13 - 3

13-3 . .(Pianka, 1973)

13 - 3

. k j i Pki Pji . 1 0 1 .

2 (E) 5 - 13 - 3

.(Chesson, 1983) 14-3 14 - 3

Overlap Index Electivity Index 59

. Pi ri - (1) (-1) - 1 . +1 .(Zavala-Camin, 1996)

1 (MTI) 6 - 13 - 3 ) ( . .(Christensen and Pauly, 1993) 15-3 15 - 3

i j DCi,j

j i FCj,i . .

.(Pauly et al., 1998)

2(OI) 7 - 13 - 3 16 - 3 .(Pauly et al., 1993) 16 - 3 DCij i TLi j TLj OI . j j

Mixed Trophic Impact Omnivory Index 60

. - . .(Pauly et al., 1998) OI

Aspect Ratio 8 - 13 - 3

.(3-3 ) .

( ) . .(Pauly et al., 1998) . 17 - 3 17 - 3 . s h

Aspect Ratio 3-3

1 (CI) 9 - 13 - 3

. 18 - 3 .(Keitt et al., 1997)

18 - 3

i (1= 0= ) k j cijk

- ni 1 0 . . Odum, 1969; ) .(Christensen et al., 2005 .(Christensen and Walters, 2004)

10 - 13 - 3 . .

( i) 2 19-3 .(Libralato et al., 2006) ( KSi) . 19 - 3

j i mij

MTI (KSi) . . 20 - 3 (Power et al., 1996)

KSi = log ( i(1-Pi)) 20 - 3

Connectance Index Keystoneness 62

. i Pi

14 - 3 .(King, 1996) Pauly, ) t PCoA1 .(1984b .(Anderson, 2006)

: (Trichiurus lepturus) 6 3757 (Liza klunzingeri) (Rastrelliger kanagurta) (Sillago sihama) - (Pomadasys kaakan) (Acanthopagrus latus)

Coordinates Analysis Principal 63

... .

1-4

(Trichiurus lepturus) 1 - 1-4

486 - - (1395 - 1396) - 2 .(1 - 4 )

- . 78/09 ±(10/85) ( )± . %89 % 39 76-90 %89 76 %39 . 90

(1395-1396) 1-4 ± ±

( ) ( ) ( ) ( )

78/09 ±(10/85) 369/15 ±(144/04) 486 96/50 50/50 789 120

. 1-4

-1396) 1-4 (1395

. - 2/64 0/003 b a 95 3 (2/64) b t . . (P<0/05) 2-4 . .

(1395-1396) 2-4

FiSAT II . 2-4

(1395-1396) 2-4

Z) M) F) K) L ) (E) ( ) ( ( ( ( (

0/5 1/11 0/49 0/62 3/45 0/25 107/40

- 486 . 15 . . ( 3 - 4 ) 63 22 62/55 (VI1 ) .

Vacuity Index 90

3-4 (1395-1396)

46/45 ) 3/14 6/07 .(4 - 4

-1396) 4-4 (1395

Synodontidae Clupeidae Scombridae Mugilidae Carangidae Mullidae Trichiuridae . Engraulidae Platycephalidae Penaeidae Loliginidae Sepiidae 3 . . Octopodidae - 11/81 ( 7/06) ( 11/02) 0/08 .(1 ) 22/13 . .(5-4 )

5-4 (1395-1396)

6-4 (%90) . .

(1395-1396) 6-4 3 - 4 17/68 50/75 .

(%W) 3-4 (1395-1396) %W 1 Rastrelliger kanagurta 8/64 2 Sardinella longiceps 7/42 3 Saurida tumbil 8/43 Trichiurus lepturus 1/48 4 Upeneus sulphureus 13/28

1 2 3

1 Atule mate 17/68 2 Liza klunzingeri 50/75 Platycephalus sp.3 0/69 4 Thryssa sp. 0/68 19/67 5 Penaeus semisulcatus 0/08 4/73 0/49 Sepia sp. 0/03 Loligo sp. 8/73 Octopus sp. 0/37

- ( 46/45) .(4 - 4 ) ( 57/40) 20/43 . 10/74 . 4/49 4/86

(%N) (%F) 4-4 (1395-1396) %F %N

2 3 4

1 Rastrelliger kanagurta 1/93 1/56 2 Sardinella longiceps 4/49 2/12 3 Saurida tumbil 4/86 2/12 Trichiurus lepturus 0/19 0/08 Upeneus sulphureus4 20/43 11/02 5 Atule mate 57/40 7/06 6 Liza klunzingeri 10/74 11/81 Platycephalus sp.7 1/47 0/68 8 Thryssa sp. 0/49 0/26 22/13 9/74

9 Penaeus semisulcatus 9/80 2/21 9/47 3/69 0/83 0/17

Sepia sp. 0/08 0/09 Loligo sp. 6/07 2/ 71

Octopus sp. 0/35 0/34

(Sillago sihama) 2 - 1-4 795 (1395-1396)

1 2 3 4

6 7 8

.(5 - 4 ) 2 . 17/39 ±(2/43) ( )± . %91 % 63 18-20

(1395-1396) 5-4 ± ±

( ) ( ) ( ) ( )

17/39 ±(2/43) 41/68 ±(17/84) 795 25/60 12/40 131/28 13/28

. 7-4

(1395-1396) 7-4

. . 2/96 0/008 b a 95 3 (2/96) b t

. (P>0/05) 8-4 . .

(1395-1396) 8-4

FiSAT II . 6 - 4

(1395-1396) 6-4

K) L ) Z) M) F) (E) ( ) ( ( ( ( ( 97

0/6 3/55 1/09 2/46 2/53 0/47 26/91

- 795 - . .(9-4 ) 33/79 35/43 30/78 . 30/94 (VI) .

(1395-1396) 9-4

. ( ) Dinophyceae ( ) Bacillariophyceae ( - ) Cyanophyceae

. 7-4 . 35/71 68/40 76/20

(1395-1396) 7-4

( )

NM FO NM FO NM FO NM FO NM FO (%) (%) (%) (%) (%) (%) (%) (%) (%) (%)

Surirella sp. - - - - 0/32 - 6/90 7/89 6/33 3/80 Gyrosigma sp.- - - - 52/29 38/10 - - 3/73 10/13 Amphora sp. 1/85 7/14 - - 5/52 23/81 1/40 2/63 1/70 8/86 Pleurosigma - - - - 29/22 14/29 - - 2/09 3/80 sp. Cymbella sp.------1/19 13/16 1/09 6/33 Navicula sp.------28/07 23/68 25/67 11/ 39 Cerataulina ------1/52 2/63 1/40 1/27 sp. Pinularia sp.------0/38 - 0/35 - Cymatopleura ------0/13 2/63 0/11 1/27 sp. Hemidiscus ------3/05 2/63 2/79 1/27 sp. Coscinodiscus 44/44 28/57 - - - - 15/84 7/89 15/04 8/ 86 sp. Rhizosolenia ------0/05 2/63 0/05 1/27 sp. Peridinium sp.------0/05 2/63 0/46 1/27 46/29 35/71 - - 87/35 76/2 59/03 68/4 60/81 59/ 52 - Oscillatoria 1/85 7/14 ------0/02 1/27 sp.

Alexanderium ------0/02 2/63 0/02 1/27 sp. Polykrikas sp. ------2/53 5/26 2/32 2/53 Dinophysis sp. ------0/05 - 0/05 - Prorocentrum ------1/50 7/89 1/37 3/80 sp. Pyrocyst sp. ------0/50 2/63 0/46 1/27

------4/60 18/41 4/22 8/87 1/85 7/14 ------0/02 1/27 35/20 28/57 - - - -35/79 7/89 33/17 8/86 - -33/33 33/33 - - - - 0/04 2/53 - - 66/67 66/67 - - - - 0/09 5/06 1/85 7/14 - - 0/97 4/76 - - 0/09 2/53 ------0/50 2/63 0/46 1/27

12/96 14/29 - -11/68 19/05 0/05 2/63 1/04 8/86

2(FO) 1(NM)

( 5-10/90) - ( 17-23/90) ( 11-16/90) ( 5-10/90) . 11 - 4 10- 4 . - 16/90) . 90/39 83/30 (11 .

Numerical method Frequency occurrence method 100

- ( 11-16/90) 10-4 (1395-1396)

- ( 17-23/90) 11-4 (1395-1396)

101

(Rastrelliger kanagurta) 3 - 1-4 756 (1395-1396) 2 ( )± .(8 - 4 ) . 20/55 ±(1/33) 21-23 . %99 % 62

(1395-1396) 8-4 ± ±

( ) ( ) ( ) ( )

20/55 ±(1/33) 153/85 ±(28/01) 756 23/20 17/80 228/51 96

. 12-4

(1395-1396) 12-4

102

- b a . . 2/63 0/053 95 3 (2/63) b t . (P<0/05) 13 - 4 . .

(1395-1396) 13-4

FiSAT II . 9 - 4

103

(1395-1396) 9-4

K) L ) Z) M) F) (E) ( ) ( ( ( ( ( 0/6 3/28 1/15 2/13 2/46 0/49 24/42

- . 756 . 15/75 22/01 15/14 36/15 (VI) .(14-4 ) .

(1395-1396) 14-4

28/01 . 104

.(15-4 ) 71/99 (Dinophyceae) (Bacillariophyceae) . 66/67 71/99

15-4 (1395-1396)

Coscinodiscus sp. . 10 - 4 12/88 46/92 .

(1395-1396) 10-4

Tintinnidae 910 2/53 20 2/86

105

2724 7/57 40 5/72

1551 4/31 45 6/44

Alpheus sp. 1300 3/72 12 1/72

Harpacticoida 124 0/34 18 2/58

794 2/21 26 3/72

515 1/43 31 4/43

Amphora sp. 105 0/29 30 4/29

Cerataulina sp. 19 0/05 1 0/29

Nitzschia sp. 3 0/01 2 0/14

Leptocylindrus sp. 3 0/01 1 0/14

Melosira sp. 25 0/07 1 0/14

Proboscia sp. 130 0/36 1 0/14

Helicotheca sp. 30 0/08 1 0/14

Pinnularia sp. 33 0/09 11 1/57

Bellerochea sp. 78 0/21 4 0/57

Bacteriastrum sp. 1 0/01 1 0/14

Coscinodiscus sp. 16890 46/92 90 12/88

Guinardia sp. 3507 9/74 53 7/58

Gyrosigma sp. 361 1/00 34 4/86

Navicula sp. 183 0/51 16 2/29

Pleurosigma sp. 370 1/03 55 7/87

Rhizosolenia sp. 828 2/30 9 1/29

106

Surirella sp. 70 0/19 21 3/00

Ceratium sp. 229 0/64 28 4/01

Dinophysis sp. 1327 3/69 31 4/43

Noctiluca sp. 760 2/11 12 1/72

Polykrikos sp. 90 0/25 3 0/43

Prorocentrum sp. 41 0/11 3 0/43

Peridinium sp. 673 1/87 48 6/87

Pyrocystis sp. 161 0/45 9 1/29

Forams

18 0/05 10 1/43

2057 5/71 24 3/43

49 0/14 7 1/00

16 - 4 48/1 53/3 ) . 40 . ( 5/83 ) ( 36/26 . 17-4 .

107

16-4 (1395-1396)

17-4 (1395-1396)

(Liza klunzingeri) 4 - 1-4

108

776 - (1395 - 1396) 2 .(11 - 4 ) - 13/83 ±(1/30) ( )± 15 . . %84

(1395-1396) 11-4 ± ±

( ) ( ) ( ) ( )

13/83 ±(1/30) 36/48 ±(9/89) 776 18/30 9/20 73/62 10/60

. 18-4

(1395-1396) 18-4

109

- b a . . 2/77 0/024 95 3 (2/77) b t . (P>0/05) 19-4 . .

(1395-1396) 19-4

FiSAT II . 12 - 4

110

(1395-1396) 12-4

K) L ) Z) M) F) (E) ( ) ( ( ( ( ( 0/3 2/12 1/33 0/79 2/31 0/55 19/34

- . 776 . ) 51 32 17 39/82 (VI) .(20 - 4 .

(1395-1396) 20-4 - 12/6 4/47 .(21 - 4 )

111

12/6 (Bacillariophyceae ) . 58/38

21-4 (1395-1396)

- 13 - 4 3/99 Cyclotella sp. . . 13/41

(1395-1396) 13-4

12 0/29 8 4/05 31 0/74 23 11/65 11 0/26 11 5/57 7 0/17 7 3/55 24 0/57 19 9/62 20 0/48 16 8/10 82 1/96 38 19/25

112

Proboscia sp. 14 0/36 2 0/56 Chaetoceros sp. 30 0/76 25 6/98 Nitzschia sp. 80 2/03 32 8/ 94 Melosira sp. 8 0/20 6 1/68 Pinnularia sp. 9 0/23 2 0/56 Cyclotella sp. 157 3/99 48 13/41 Bacteriastrum sp. 11 0/28 2 0/56 Rhizosolenia sp. 6 0/15 2 0/56 Coscinodiscus sp. 21 0/53 17 4/75 Guinardia sp. 4 0/10 2 0/56 Gyrosigma sp. 81 2/06 30 8/38 Navicula sp. 24 0/61 17 4/75 Pleurosigma sp. 24 0/61 11 3/07 27 0/69 13 3/63

(Acanthopagrus latus) 5 - 1-4 528 (1395-1396) 2 .(14-4 ) 27/03 ±(4/92) ( )± 26-34 . . %95 % 45

(1395-1396) 14-4 ± ±

( ) ( ) ( ) ( ) 27/03 ±(4/92) 515/88 ±(257/12) 528

113

36/80 16/80 968 116

. 22 - 4

(1395-1396) 22-4

- b a . 2/83 0/041 95 3 (2/83) b t . . (P<0/05) 23-4 . .

114

(1395-1396) 23-4

FiSAT II . 15-4

(1395-1396) 15-4

K) L ) Z) M) F) (E) ( ( ( ( ( ( )

0/4 1/72 0/89 0/83 2/77 0/40 38/45

- 115

. 528 . ) 39 28 33 36 (VI) .(24 - 4 .

-1396) 24-4 (1395

(%80/54) (%2/57) (%38/63) (%18/02) .(25 - 4 )

116

(1395-1396) 25-4 26-4 (%57) . .

(1395-1396) 26-4

117

16-4 6/73 6/93 .

(%W) 16-4 (1395-1396) %W

Synodontidae 1/92 Engraulidae 1/76 Gerreidae 6/73 Nemipteridae 6/93 22/84

4/99 7/37

8/32 7/49

1/30

17-4 . 17 - 4 80/22 70/54 .

118

- (%N) (%F) 17-4 (1395-1396) %F %N

Synodontidae 5/93 6/95 Engraulidae 3/58 5/65 Gerreidae 1/42 1/40 Nemipteridae 1/62 1/43 5/47 2/59

19/21 21/44 34/17 17/19

54/46 39/52 27/76 31/02

2/82 2/57

(Pomadasys kaakan) 6 - 1-4 416

(1395-1396) 2 ( )± .(18-4 ) - 40-54 . 43/77 ±(7/50) . %94 % 35

119

(1395-1396) 18-4 ± ±

( ) ( ) ( ) ( ) ±(624/31) 43/77 ±(7/50) 416 62/50 28/30 1304/29 2691 322

. 27 - 4

-1396) 27-4 (1395

. 0/021 b a 3 (2/89) b t . 2/89 . (P>0/05) 95

120

28-4 . .

(1395-1396) 28-4

FiSAT II . 19 -4

(1395-1396) 19-4

K) Z) M) F) L ) (E) ( ( ( ( ( ) (

0/7 3/05 0/65 2/4 3/11 0/31 64 /76

121

. 416 . 74 14 12 71/87 (VI) .(29-4 ) .

-1396) 29-4 (1395

(%14/12) (%9/35) (%31/22) (%56/74) .(30-4 )

122

(1395-1396) 30-4

31 - 4 (%40/12) . .

-1396) 31-4 (1395

123

20 - 4 . 17/19

(%W) 20-4 (1395-1396)

0/74 10/67 3/09 0/09 2/07

Sillaginidae 0/92 Engraulidae 1/83

Squilla 17/19 mantis 22/93

20/34 . 21 - 4 . 79/84

124

- (%N) (%F) 21-4 (1395-1396)

%F %N 32/83 9/35 17/82 6/40 15/32 1/49 19/11 7/44 14/17 5/19

Sillaginidae 20/82 5/57 Engraulidae 16/36 5/65

Squilla 14/86 9/32 mantis 47/16 11/02

2-4 . PCoA1 2 ( ) . ( )

32-4 . . 73 .

Principal Coordinates Analysis matrix Similarity 125

- . .

32-4

126

3-4 1 - 3-4 (EE) (Q/B ) (W ) Aspect ratio (Ar) - . 22-4 (2/94) (11/60) .(33 - 4 )

Q/B EE 22-4

W Q/B Ar EE (gr) (/y) 1 150/ 55 2/94 0/88 1/01 86 /45 3/99 0/75 4/44 353/60 11/60 0/ 33 1/59 105/16 6/60 0/09 2/45 1749/84 5/10 0/24 1/31 3372/27 3/90 0/22

127

33-4

2 - 3-4 23-4 Pray/predator S. commerson ) 30 1 30 1 .( . S. commerson 1 . Pray/predator

128

129

23-4

90 3 - 3-4 1 . (TL ) .

. 24-4 ( )

. 4 . . 1 . 2 (TL=2/10) (TL=2) (TL=2/50) 3 . (TL=2/50) (TL=2/98) (TL=2/86) (TL=3/61) (TL=3/95) (TL=3/49) (TL=3/48) (TL=3/09) (TL=3/65) (TL=3/30) (TL=3/19) (TL=3/37) (TL=3/51) 4 . (TL=3/01) (TL=3) (TL=3/20) (TL=4/56) (TL=4/20) (TL=4) (TL=4/16) (TL=4/45) - . (TL=4/41) (TL=4/13) (TL=4/14)

Trophic Level 99

24-4 Biomass P/B Q/B Trophic level EE P/Q (t/km²) (/year) (/year) Scomberomorus commerson Trichiurus lepturus Saurida tumbil Epinephelus coioides Nemipterus japonicus Upeneus vittatus Atule mate Lutjanus johni i Lutjanus malabaricus Pomadasys kaakan Sardinella sindensis Otolithes ruber Lethrinus nebulosus Psettodes erumei Acanthopagrus latus Grammoplites suppositus Parastromateus niger Drepane punctata Sillago sihama Rastrelliger kanagurta Liza klunzingeri Sepia pharaonis Squid Crab Shrimp Benthos (other) Bivalve Gastropoda Polychaete Zooplankton Seaweed Phytoplankton Detritus

100

4 - 3-4 1

1 0 . 1 34-4 .

Overlap Index

101

34- 4 )

102

5 - 3-4 1 35-4 . (1) ( - 1) 1 - . .

. .

Electivity Index

103

) 35-4

104

6 - 3-4 1 - . 36 - 4

- . . . .

Mixed Trophic Impact

105

36-4 - . )

7 - 3-4

106

37-4 . . 4 - . .

) 37-4

8 - 3-4

107

1 . (OI )

25-4 .

( ) .

25-4

OI Scomberomorus commerson Trichiurus lepturus Saurida tumbil Epinephelus coioides Nemipterus japonicus Upeneus vittatus Atule mate Lutjanus johnii Lutjanus malabaricus Pomadasys kaakan Sardinella sindensis Otolithes ruber Lethrinus nebulosus Psettodes erumei Acanthopagrus latus Grammoplites suppositus Parastromateus niger Drepane punctata Sillago sihama Rastrelliger kanagurta Liza klunzingeri Sepia pharaonis Squid Crab

9 - 3-4

Omnivory Index

108

1 . 38 - 4 . .

38-4 ) (.

10 - 3-4

Keystone Index

109

26 - 4 1 0 . 3/08 0/14 1 . 2

- . 0/15 . .

26-4

23099/34 t/km2/yr 14475/56 t/km2/yr 3/080875 / 43/26934 / 0/003563478 /yr ( ) 165/7789 t/km2 0/1490139 0/1517912

4-4

( ) . - 2016 2004 ( ) . 39 - 4 .

Connectance Index System Omnivory Index

110

- ( ) ( ) 39-4 (2004-2016 )

111

: Trichiurus ) (Liza klunzingeri) (Rastrelliger kanagurta) (Sillago sihama) (lepturus (Pomadasys kaakan) (Acanthopagrus latus)

. .

... .

1 - 5 -

78/09 ±(10/85) ( )± (2012) Fofandi .(1-4 ) . 52/1

112

. . (2004) Abdurahiman (2012) Fofandi (2 - 4 ) 107/40 L Pauly Ingles ( 45/5) (1958) Misu . ( 106/83) (1997) Reuben ( 78) (1984) - 131 /25) (2012) Fofandi L Dawson ( 138/1 145/2) (1994) Narasimham ( (1989) Joseph Somavanshi (1967) Thiagrajan ( 129/7) (1990) Chakraborty ( 109) Abdussamad ( 126 129) (1992) . ( 128/2) (2006)

1 - 5 .(Jones, 1981) . (Z) (M) (F) (2-4 ) 0/5 1/11 0/49 0/62 (E) . (1-5 ) .

123

1-5

Z M F K L E

3/16 0/4 2/7 138/1 Narasimham (1994) 0/4 148 Chakraborty (1997) 0/63 2/41 0/89 1/52 0/61 106/83 Reuben et al. 1997 0/71 2/66 0/77 1/89 0/5 128 Mohite and Biradar (2001) 0/75 3/47 3/23 0/67 127/3 Khan (2006) 0/3 0/44 0/31 0/13 0/13 131/25 Fofandi (2012) 0/5 1/11 0/49 0/62 0/25 107/40

62/55

. (2015) Rohit

3/14 6/07 46/45 . (1379) 13/9 16 70/1

. 2 - 5

.(De la Cruz-Torres et al., 2014)

Martins and Haimovici, 2000; )

124

- .(Wojciechowski, 1972

.(Agnaldo et al., 2005)

2-5

Rohit et al. 2015 Yan et al. 2011 Chiou et al. 2006 Bittar et al. 2012 - Bakhoum (2007)

( ) Agnaldo et al. 2005 De la Cruz-Torres et al. 2014 ) Portsev (1980) ( ) Munekiyo and Kuwahara (1985) ( ) ( Martins et al. 2005

1 - 5 . .

(1978) Sin .

Euryphagous 125

. . Bakhoum, ) .(2007

. - 4/45 (Portsev, 1980) 4/45 4/10 Bakhoum (2007) . Akhtar, ) .(2008 ) (r=0/81) 1385 ( . (E = 0/5) .

Pauly et al., ) .(1998

. Rohit . Karnataka (2015)

126

- 131 /28 25/60 (1395) .(5 - 4 ) 106 24/6 (2010) Basavaraja Vinod . . 77/64 21/96

.(Yildirim et al., 2002; Beverton and Holt, 1957; Innal et al., 2015)

. Gowda (1393) (1380) (1978) kaliamurthy Krishnamurthy (1988) (2004) Annappaswamy . (2010) Ansari Shamsan . Dulcic and Karaljevic, ) .(1996 . 26/91 0/47 L K (1984) Pauly Ingles (2015) Hu L 23/5 K 0/7 (1957) Radhakrishnan L 26/1 K 0/6 . L 28/1 K 0/7

127

(2008) Ansari Shamsan . %53/3 .

Ansari Shamsan .(Kariman et al., 2009) (1991) Neelakantaan Reddy (1991) Jayasankar (1988) Gowda (2008)

(Bacillariophyceae) (Cyanophyceae) - . %59/52 (Dinophyceae) Shamsan . (2008) Ansari (1988) Gowda . . . . .

.(Kariman et al., 2009) .

. (1988) Gowda Radhakrishnan (1957) Shamsan (1985) Milward Gunn (2003) Mohammed (2006) Hajisamae (1997) Weerts (1991) Reddy (2008) Ansari .

128

. 83/30 (11 - 16/90) . 90/39 (2008) Ansari Shamsan . (2008) Ansari Shamsan .

. (2008) Ansari Shamsan (1985) Gunn . 3/30 . 3/37 (2004) Hajisamae . . 3/4 Caledonia (2005) Bozec 3/1

Prasert . . (2005) .

. . (2014) Amin (9 - 4 ) 24/42 L Mehanna ( 27/83) (2014) Amin ( 30/70) (2004) Gupta Rohit ( 30/76) (2001) Mehanna Al-Mahdawi ( 33/28) (2010) Abdussamad . ( 32/5) (2010)

129

Ragonese and Bianchini, ) . .(1998

. 3-5 .(Beverton and Holt, 1957) .

3-5

Z M F K L E

0/45 4/44 2/46 1/98 1/50 27/83 Amin et al. (2014) 0/63 30/76 Mehanna (2001) 0/66 7/63 2/60 5 1/80 30/70 Rohit and Gupta (2004) 0/86 6/04-11/85 2/52 3/52 -9/33 1/63 33/28 Abdussamad et al. (2010) Al-Mahdawi and 0/61 1/91 0/75 1/16 0/58 32/5 Mehanna (2010) 0/6 3/28 1/15 2/13 0/49 24/42

(2000) Dadzie .

Bagheri . 46/28 93 (2013) .

130

( 5/83 ) ( 36/26 ) . . (2016) Sree Renjima

. . Sivadas (1973) Luther (1952) George Bhimachar (2013) Bagheri . (2016) Supraba (2009) Bhaskaran (2005) Prasert .

. . (2002) Abou-Seedo 0/55 19/34 K L K L .(12 - 4 ) .(Dadzie et al, 2005) 0/46 24/8 ( )

.(Jones, 1981) .

. .(Sparre and Venema, 1998)

131

King, ) .(1995 . . 1/33 2/12 ) (12-4 ) 0/79 3/56 1/06 4/61 ( (Dadzie et al., 2005) . - 12/6 4/47 . . Uddin . (Bacillariophyceae) . (2012) - 2/50 . (Santini et al., 2015) 2/6

.(Akhtar, 2008)

. (1991) Samuel Matthews (1997) Abdullah Hussain .

132

38/45 0/40 L K L K (1991) Samuel Matthews .(15 - 4 ) (1985) Morgan 44/5 0/2 ( ) (2004) Grandcourt 44/5 43 0/23 0/23 ( ) (1985) Edwards 55 0/23 64/6 0/14 (2009) Al-Mamry 57/8 0/21 (1981) Jones . .

(15 - 4 ) 0/83 0/89 1/72 Matthews 0/31 ( ) . (1991) Samuel .(Gulland, 1991) . 3/51 3/15 (2000) Nasir .

(%2/57) (%38/63) (%18/02) . (%80/54) (1993) Al-Daham

. .(Deshmukh, 1973) . -

133

43/77 ±(7/50) ( )± (1396) .(18 - 4 ) (2011) Fakhri . 42/7 ±(4/95) - (2011) Valinassab 43 ±(3) - (2002) Al-Husaini 61 . 16 - 75

.(Al-Husaini et al., 2002)

. (1396) 64/76 0/31 L K (19-4 ) .(L = 63/5 K=0/21) (1396) 62/55

. (1389)

(%9/35) (%31/22) (%56/74) 79/84 20/34 . (%14/12)

134

(2011) Valinassab . -

Fischer . (1990) Niameimandi (1984) Bianchi Tonozuka Kuiter (1992) Talebzadeh (1389) (2001)

(1389) . III II . .

. . Van 3/65 . (3/46) (1991) Adkin der Elst .

2 - 5

135

(TL) Gascuel et ) .(al., 2005 . .(Pauly et al., 2002)

.(Essington et al., 2006) 4 8 (2017) Dutta . 15 2 2/716 . 4 1 - 5 . .

136

1-5

3/081 (2008) Mohamed (2012) Ullah . 3/04 Arreguin-Sanchez 4/48 (2003) Bradford-Griveetal 2/45 (2003) Vivekanandan 2/82 (2004) . 2/71 (2017) Dutta 3/61

.(Dutta et al., 2017) Hunter ) (2000) Shannon .(and Price, 1992

. .

137

Duarte and ) - 77 .(Garc, 2004 .(Frank et al., 2005)

78 .

.(Frank et al., 2005)

.(Springer et al., 2003) (2007) Okey (2002 1999) Kitchell 79 12 . (3/51 -4/56) . 1

Q/B P/B . Maldeniya, 1996; Olson and Galvan-Magana, ) .(2002; Griffiths et al., 2007

- . (1999) Kitchell

Wasp-Waist Bottom-up Ecological Redundancy 138

. (2002) Schindler

) ( ) . (

- (%23) (%64) 59 . 70 . (%1) (%6) (%6) 16 . (%3) (%6) (%91) (%28) (%64) . (%8)

. .

.(Matthew et al., 2007) %96 %100 %22 %55 %10 %18 (2003) Vivekanandan .

139

. (2010) Abdurahiman . Ross (2010) Bradford-Grieve Pinkerton %14 %61 toothfish . .

. (2007) Matthew .

- . - .(1384 )

80 .(Kurian, 1971) (1961) Slobodkin . .

. (1969) Odum . 10 10 .

Ecological Efficiency 140

. 81 (SOI) . .(Christensen et al., 2000) Arreguin- . 0/15 (2003) Vivekanandan 0/17 (2004) Sanchez (2012) Ullah . 0/1 Mohamed 0/35 (2011) Chen 0/22 (2017) Dutta 0/29 (2008) . 0/35

3 - 5

.(Bentley et al., 2017)

( )

System Omnivory Index 141

. 38-4

- .(Sakamoto et al., 2017)

.(Mutsert et al., 2017)

.(Christensen, 1998) Sagarese . (2017)

4 - 5 ( ) . 83 82

. . .

- .

Connectance Index System Omnivory Index 142

- 1 . - 2 . ( ) - 3 . - 4 . - 5 .

CPUA CPUE .1396 ...... ( ) (Pomadasys kaakan) .1 :16

143

Sillago ) .1380 . . . .37 - 54 :(1)10 . (sihama .1395 ...... 1387 . . . . .68-77 :(3)17 . (Sardinella sindensis) .1395 . . . . . - . (Sillago sihama Forskal, 1775) .29 :8 / .( ) .1380 . . 437 . : .1393 . . . . - 47 :(17) 5 .( ) (Sillago sihama ) .41 . (Trichiurus lepturus) .1379 . .65-72 :(1) 9 .

144

.1389 . . . . .1:4 . (Pomadasys kaakan) .1391 . . . . . 50 . . ¬ .1384 . . 106 . 280 .382 . . .1385 . . .1395 ...... 333 .

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