Species, sex, size and male maturity composition of ... - Seaturtle.org

Journal of Fish Biology (2012) 

doi:10.1111/j.1095-8649.2011.03210.x, available online at wileyonlinelibrary.com 

Species, sex, size and male maturity composition 

of previously unreported elasmobranch landings 

in Kuwait, Qatar and Abu Dhabi Emirate 

A. B. M. Moore*†‡, I. D. McCarthy*, G. R. Carvalho§ and R. Peirce‖ 

*School of Ocean Sciences, Bangor University, Askew Street, Menai Bridge, Anglesey, LL59 

5AB, U.K., †RSK Environment Ltd., Spring Lodge, 172 Chester Road, Helsby, Cheshire, WA6 

0AR, U.K., §Molecular Ecology & Fisheries Genetics Laboratory, Environment Centre for 

Wales, Bangor University, Deiniol Road, Bangor, Gwynedd LL57 2UW, U.K. and ‖Shark 

Conservation Society, Dulverton House, 8 Crooklets, Bude, Cornwall, EX23 8NE, U.K. 

This paper presents data from the first major survey of the diversity, biology and fisheries of elasmobranchs 

in the Persian (Arabian) Gulf. Substantial landings of elasmobranchs, usually as gillnet 

by-catch, were recorded in Kuwait, Qatar and the Emirate of Abu Dhabi (part of the United Arab 

Emirates), although larger elasmobranchs from targeted line fisheries were landed in Abu Dhabi. The 

elasmobranch fauna recorded was distinctive and included species that are undescribed, rare and have 

a highly restricted known distribution. Numerical abundance was dominated by sharks (c. 80%), 

of which carcharhinids were by far the most important. The milk shark Rhizoprionodon acutus 

and whitecheek shark Carcharhinus dussumieri together comprised just under half of all recorded 

individuals. Around 90% of recorded sharks were small (50–90 cm total length, L T ) individuals, 

most of which were mature individuals of species with a small maximum size (

2 A. B. M. MOORE ET AL. 

29° 

Iraq 

Kuwait 

Sharq 

Fahaheel 

Shatt al Arab 

Iran 

Iraq 

Saudi Arabia 

Red Sea 

Yemen 

Gulf of Aden 

Iran 

Pakistan 

Gulf 

Oman 

Gulf of Oman 

Arabian Sea 

26° 

Bahrain 

Saudi Arabia 

Doha 

Qatar 

Al Khor 

The 

Gulf 

Abu Dhabi 

Strait of Hormuz 

Gulf of 

Oman 

0 50 100 

km 

200 

U.A.E. 

Oman 

49° 53° 56° 

Fig. 1. Map of The Gulf showing political boundaries and the locations sampled during the current survey. 

research is lacking in the Indian Ocean, despite high and increasing reported landings 

(Anderson & Simpfendorfer, 2005). In Arabian waters, Oman has led the efforts to 

characterize the diversity, biology and fisheries of its elasmobranch fauna in the Gulf 

of Oman and the Arabian Sea, both through landings-based and fisheries-independent 

data (Henderson et al., 2006, 2007, 2009; Henderson & Reeve, 2011). The elasmobranchs 

of The Gulf (Fig. 1) have received little attention, however, and are poorly 

understood (Moore, 2011). A number of concerns about Gulf elasmobranchs have 

been highlighted including the role of the Islamic Republic of Iran and the United 

Arab Emirates (U.A.E.) as major contributors to global elasmobranch landings and 

shark-fin exports, respectively, a possible increase in demand for meat and cartilage 

locally, as well as a possible fisheries-related change in elasmobranch community 

composition along the Iranian coast since the 1970s (Moore, 2011). 

In addition to research efforts, since 1999, the United Nations Food and Agriculture 

Organisation (FAO) has encouraged all states catching elasmobranchs in either 

targeted or by-catch fisheries to voluntarily participate in the International Plan of 

Action for the Conservation and Management of Sharks (IPOA-Sharks) and to have 

developed a ‘Shark-Plan’ by 2001 (FAO, 1999). The IPOA-Sharks is aimed at ensuring 

the sustainable use of all chondrichthyans and places particular emphasis on the 

importance of catch data to support this aim (FAO, 1999). All of The Gulf states (i.e. 

Bahrain, Iran, Iraq, Kuwait, Qatar, Saudi Arabia and the U.A.E.; excluding Oman 

here, whose coastline is largely non-Gulf) operate a range of fisheries for teleosts 

and invertebrates, including extensive gillnetting (Bishop, 2002). In addition, there 

are some reports of targeted elasmobranch fisheries in Gulf waters of the U.A.E. 

(Anderson & Simpfendorfer, 2005). As such, all Gulf states can be assumed to be 

© 2012 The Authors 

Journal of Fish Biology © 2012 The Fisheries Society of the British Isles, Journal of Fish Biology 2012, doi:10.1111/j.1095-8649.2011.03210.x

ELASMOBRANCHS IN KUWAIT, QATAR AND ABU DHABI 3 

catching elasmobranchs to some degree in their fishing activities, although to date 

none have adopted a Shark-Plan (FAO, 2011a). Kuwait and Qatar have either not 

reported elasmobranch landings or reported zero values to the FAO (FAO, 2011b). 

The U.A.E. has reported landings to the FAO since 1986 (FAO, 2011b), and Abu 

Dhabi Emirate publishes data on elasmobranch landings (by port, gear and vessel 

type, and month) as part of annual fisheries statistics (Environmental Research & 

Wildlife Development Agency, 2005; Environment Agency Abu Dhabi, 2010). The 

value of these data in assessing the composition of elasmobranch landings is compromised, 

however, by broad and unspecified reporting, such as of ‘sharks and rays’ 

or similar broad taxonomic groupings. 

Given the concerns noted above and the paucity of data and management locally, 

there is a clear need for basic information on the diversity, size composition, reproductive 

biology and fisheries of Gulf elasmobranchs (Moore, 2011). Surveys of fish 

markets and landing sites have provided a rapid and relatively inexpensive source of 

important data for elasmobranchs elsewhere on a range of aspects including reproductive 

biology, fisheries and new species (White & Dharmadi, 2007; Bizarro et al., 

2009; Last et al., 2010a). In order to address local data gaps, the current paper reports 

species, size and sex composition, male maturity and fisheries of elasmobranchs 

recorded at landing sites and markets at three locations approximately equidistant 

along the Arabian coastline of The Gulf, i.e. Kuwait (north-western Gulf), Qatar 

(southern) and Abu Dhabi Emirate, of the U.A.E. (south-eastern) (Fig. 1). The results 

comprise the first major survey of elasmobranch diversity, biology and fisheries in 

The Gulf and form an important addition to knowledge on the relatively poorly 

known status of elasmobranchs in the western Indian Ocean. 

STUDY REGION 

MATERIALS AND METHODS 

The Gulf (Fig. 1) is a shallow (average depth 35 m), semi-enclosed offshoot of the northwest 

Indian Ocean, connected to the much deeper Gulf of Oman (>3000 m), and subsequently 

the Arabian Sea, through the narrow Strait of Hormuz. The Gulf is a harsh environment, 

and in some cases can be subject to extremes of water temperature (4–39 ◦ C) and salinity 

(brackish to >70); the single major freshwater input discharges into the north-west Gulf 

through the Shatt al Arab near northern Kuwait (Sheppard et al., 1992; Carpenter et al., 

1997). In general, coastal waters of Qatar and the Emirate of Abu Dhabi (the largest in the 

U.A.E.) are characterized by extensive shallows of


E); in Qatar (12–29 April 2009) at Corniche, Doha (25 ◦ 17 ′ N; 51 ◦ 32 ′ E), Doha main 

wholesale market (25 ◦ 17 ′ N; 51 ◦ 32 ′ E) and Al Khor (25 ◦ 41 ′ N; 51 ◦ 31 ′ E); and in the 

U.A.E. (Abu Dhabi Emirate) (5–11 April 2010) at Abu Dhabi City (Mina Zayed) (24 ◦ 30 ′ 

N; 54 ◦ 22 ′ E) (Fig. 1). Elasmobranchs were openly sold in Kuwait in 2008, but in 2011, 

the Public Authority for Agriculture and Fisheries (PAAFR) was actively enforcing a ban on 

their sale in the market, so most sampling was performed at the quayside. Further brief SCS 

visits were made to Qatar (Doha wholesale market) in April of both 2010 and 2011, although 

these were not sampled and only photographs were taken for later evaluation of broad species 

composition. A planned SCS market survey to Bahrain in April 2011 was postponed as a 

result of political unrest. 

All specimens were identified and measured to the nearest cm using total length (L T , upper 

caudal fin lobe straightened along the body axis) for sharks and guitarfishes (i.e. Rhinidae, Rhinobatidae 

and Rhynchobatidae), or disk width (W D ) for rays (i.e. all non-guitarfish batoids). 

Sex was recorded and apart from a few instances [including eight pregnant whitecheek shark 

Carcharhinus dussumieri (Müller & Henle, 1839)], male and female elasmobranchs were not 

routinely dissected for macroscopic examination of the reproductive tract owing to logistical 

constraints. For male sharks, maturity was categorized based on a slightly adapted version 

of the scale in Henderson et al. (2006) but using only external examination of claspers. The 

classes used were juvenile (i.e. claspers undeveloped, not extending beyond posterior tips 

of pelvic fins), maturing (extending beyond posterior margin of pelvic fin, but flexible and 

not fully calcified) and mature (much longer than pelvic-fin rear margin, rigid). From this, a 

male maturity size range was derived for each species, from the size of the smallest maturing 

individual to the size above which all males were mature. These data were also used to construct 

maturity ogives and L T at 50% maturity (L T50 ). For rays, only the size above which 

all individuals were deemed mature (i.e. fully developed, rigid claspers) is presented here; 

male maturity data for guitarfishes are not presented as claspers can remain flexible in mature 

animals (A. Henderson, pers. comm.). 

DATA ANALYSIS 

For those species where a total of >100 individuals were recorded (seven sharks and three 

batoids) the size-frequency distributions for each sex were tested to see if data conformed 

to a normal distribution using an Anderson–Darling test. Dependent on whether data were 

distributed normally or not, intergender size differences were then investigated using either 

a two-tailed t-test or a Mann–Whitney U-test, to test for possible sex-based differences 

in fisheries landings. The Mann–Whitney U-test was further used to investigate possible 

geographic and temporal differences in landed size for both sexes between sampling events 

in the two most commonly recorded species, i.e. milk shark Rhizoprionodon acutus (Rüppell, 

1837) and C. dussumieri. Size-frequency distributions of males and females were compared 

using a χ 2 contingency test with the size distribution divided up into 5 or 10 cm size class 

intervals as appropriate to the L T of the species under consideration. To test the null hypothesis 

of sex ratios being at parity for each sampling event, a χ 2 test was performed for each species. 

For male sharks with a sample size of >200, L T50 was calculated using the logistic equation 

Y = x + (z − x)(1 + e −K(L T−L T50 ) ) −1 ,whereY is the percentage of males mature in the L T 

size class (cm) and K, x and z are constants. The equation was fitted using the non-linear 

curve fitting programme in Minitab v14 (www.minitab.com). The relationships between C. 

dussumieri maternal L T and both litter size and mean embryo L T were investigated using 

correlation analysis. 

RESULTS 

IDENTIFICATION AND TAXONOMY 

A total of 4649 elasmobranchs were examined during the survey (Table I). The 

vast majority of individuals were successfully identified to species level, with two 




Table I. Elasmobranch taxa recorded in surveys of markets and harbours during April in Kuwait (2008 and 2011), Qatar (2009) and Abu Dhabi 

(2010): number, percentage of total of all species, size range by sex (sharks and guitarfishes LT, raysWD; mean ± S.D. cm) and male maturity. 

For sharks, the range of the smallest maturing size above which all males were mature, excluding anomalous outliers, and for rays, only the size 

above which all males were mature are presented 

Taxon 

Kuwait 

2008 

Number of individuals 

(% total of all species combined) 

Kuwait 

2011 Qatar 

Abu 

Dhabi 

Total all 

sites 

Male (♂) and female (♀) size range 

(mean ± s.d. cm) 

Male 

maturity 

(cm) 

Sharks 

Hemiscyllidae 

Chiloscyllium arabicum 20 

(1·32) 

Triakidae 

Mustelus mosis 15 

(0·99) 

Hemigaleidae 

Chaenogaleus macrostoma 39 

(2·57) 

Hemipristis elongata 1 

(0·07) 

Paragaleus randalli 5 

(0·33) 

Carcharhinidae 

Carcharhinus spp. A 62 

(4·09) 

Carcharhinus amblyrhynchoides 5 

(0·33) 

105 

(15·02) 

12 

(1·72) 

17 

(2·43) 

5 

(0·25) 

2 

(0·10) 

30 

(1·49) 

— 1 

(0·05) 

5 

(0·72) 

35 

(1·73) 

— 2 

(0·10) 

3 

(0·43) 

— 130 

(2·80) 

5 

(1·20) 

1 

(0·24) 

34 

(0·73) 

87 

(1·87) 

— 2 

(0·04) 

23 

(5·54) 

68 

(1·46) 

— 64 

(1·38) 

— — 8 

(0·17) 

♂ 56–77 (68·6 ± 4·4) 

♀ 56–80 (68·8 ± 4·9) 

♂ 65–84 (72·3 ± 4·9) 

♀ 63–83 (76·0 ± 5·0) 

♂ 70–90 (78·1 ± 5·2) 

♀ 58–92 (80·3 ± 7·8) 

♂ NA 

♀ 117–119 (118·0 ± 1·4) 

♂ 61–81 (73·2 ± 4·5) 

♀ 56–83·6 (70·3 ± 8·3) 

♂ 67–130 (84·1 ± 19·4) 

♀ 63–140 (86·4 ± 15·3) 

♂ 80–88 (85·4 ± 4·8) 

♀ 82–166 (100·0 ± 36·9) 

62–68 

NAmin –65 

NAmin –70 

NA 

61–64 

NA 

NA 




Table I. Continued 



Taxon 

Kuwait 

2008 

Kuwait 

2011 Qatar 

Abu 

Dhabi 

Total all 

sites 

Carcharhinus amboinensis 32 

(2·11) 

Carcharhinus brevipinna 23 

(1·52) 

Carcharhinus dussumieri 338 

(22·30) 

13 

(1·86) 

1 

(0·14) 

139 

(19·89) 

3 

(0·15) 

22 

(5·30) 

— 5 

(1·20) 

525 

(26·00) 

Carcharhinus falciformis — — — 1 B 

Carcharhinus leiodon 25 

(1·65) 

Carcharhinus leucas 26 

(1·72) 

Carcharhinus limbatus 32 

(2·11) 

Carcharhinus macloti 16 

(1·06) 

7 

(1·00) 

16 

(2·29) 

20 

(2·86) 

7 

(1·00) 

2 

(0·48) 

(0·24) 

70 

(1·51) 

29 

(0·62) 

1004 

(21·60) 

1 

(0·02) 

— — 32 

(0·69) 

1 

(0·05) 

27 

(1·34) 

— 43 

(0·92) 

39 

(9·40) 

— 6 

(1·45) 

Carcharhinus melanopterus — — — 1 

(0·24) 

Carcharhinus sorrah 179 

(11·81) 

5 

(0·72) 

79 

(3·91) 

Loxodon macrorhinus — — 63 

(3·12) 

25 

(6·02) 

155 

(37·35) 

118 

(2·54) 

29 

(0·62) 

1 

(0·02) 

288 

(6·19) 

218 

(4·69) 

Male(♂) and female (♀) size range 

(cm) (mean ± s.d. cm) 

Male 


(cm) 

♂ 57–227 (119·0 ± 47·6) 

♀ 69–246 (129·7 ± 44·5) 

♂ 71–214 (100·9 ± 46·1) 

♀ 72–92 (83·4 ± 6·4) 

♂ 36–96 (76·0 ± 10·5) 

♀ 36–100 (75·5 ± 14·2) 

206–227 

NAmin –204 

63–80 

♀∼60 NA 

♂ 66–132 (80·6 ± 21·1) 

♀ 68–142 (88·6 ± 24·3) 

♂ 75–158 (109·6 ± 18·2) 

♀ 82–183 (113·8 ± 25·8) 

♂ 58–210 (113·4 ± 45·8) 

♀ 55–223 (119·9 ± 44·2) 

♂ 49–83 (62·1 ± 8·6) 

♀ 59–94 (71·8 ± 12·1) 

♂ NA 

♀ 73 

♂ 52–152 (82·4 ± 17·0) 

♀ 50–166 (90·4 ± 24·4) 

♂ 53–79 (68·8 ± 5·1) 

♀ 53–84 (69·3 ± 6·2) 

NAmin –123 

NA 

164–184 

75–83 

NA 

85–110 

61–71 







Taxon 

Kuwait 

2008 

Kuwait 

2011 Qatar 

Abu 

Dhabi 

Total all 

sites 

Rhizoprionodon acutus 185 

(12·20) 

Rhizoprionodon oligolinx 172 

(11·35) 

18 

(2·58) 

54 

(7·73) 

931 

(46·11) 

112 

(26·99) 

1246 

(26·80) 

— — 226 

(4·86) 

Sphyrnidae 

Sphyrna lewini — — — 1 

(0·24) 

Sphyrna mokarran 20 

(1·32) 

— 3 

(0·15) 

Guitarfish 

Rhinidae 

Rhina ancylostoma — — — 1 

(0·24) 

Rhynchobatidae 

Rhynchobatus cf. djiddensis C 7 

(0·46) 

Rhinobatidae 

Rhinobatos granulatus 13 

(0·86) 

10 

(1·43) 

130 

(18·60) 

2 

(0·10) 

1 

(0·02) 

— 23 

(0·49) 

1 

(0·02) 

— 19 

(0·41) 

— — 143 

(3·08) 

Rhinobatos halavi — — — 13 

(3·13) 

13 

(0·28) 



♂ 47–88 (65·1 ± 7·0) 

♀ 43–89 (64·1 ± 10·1) 

♂ 45–64 (57·8 ± 4·5) 

♀ 45–85 (66·4 ± 5·4) 

♂ NA 

♀ 87 

♂ 72–137 (102·0 ± 31·7) 

♀ 72–214 (144·5 ± 42·2) 

♂ NA 

♀ 180 

♂ 81–177 (138·2 ± 25·6) 

♀ 73–149 (91·2 ± 32·4) 

♂ 47–120 (74·7 ± 17·7) 

♀ 39–175 (107·8 ± 45·3) 

♂ 67–94 (81·8 ± 9·0) 

♀ 76–94 (81·7 ± 8·7) 

Male 


(cm) 

54–68 

45–53 

NA 

NA 

NA 

NA 

NA 

NA 







Taxon 

Kuwait 

2008 

Kuwait 

2011 Qatar 

Abu 

Dhabi 

Total all 

sites 

Rhinobatos cf. punctifer — 1 

(0·14) 

3 

(0·15) 

Rays 

Dasyatidae 

Himantura fai — — — 1 

(0·24) 

Himantura imbricata 3 

(0·20) 

Himantura sp. B 17 

(1·12) 

Himantura uarnak species complex 14 

(0·92) 

Pastinachus sephen 72 

(4·75) 

Gymnuridae 

Gymnura cf. poecilura 9 

(0·59) 

Myliobatidae 

Aetobatus flagellum 13 

(0·86) 

Aetobatus cf. ocellatus 7 

(0·46) 

11 

(1·57) 

42 

(6·01) 

9 

(1·29) 

26 

(3·72) 

15 

(2·15) 

23 

(3·29) 

2 

(0·10) 

118 

(5·84) 

1 

(0·05) 

4 

(0·20) 

6 

(0·30) 

— 3 

(0·15) 

— 4 

(0·09) 

1 

(0·02) 

— 16 

(0·34) 

— 177 

(3·81) 

1 

(0·24) 

25 

(0·54) 

— 102 

(2·19) 

— 30 

(0·65) 

— — 36 

(0·77) 

— 10 

(0·22) 



♂ NA 

♀ 77–87 (82·0 ± 4·0) 

♂ NA 

♀ 124 

♂ 15–23 (19·3 ± 2·8) 

♀ 14–26 (19·9 ± 4·0) 

♂ 24–54 (41·9 ± 7·9) 

♀ 25–62 (40·6 ± 10·9) 

♂ 45–106 (87·8 ± 13·0) 

♀ 40–121(88·4 ± 20·1) 

♂ 35–96 (50·2 ± 11·0) 

♀ 32–89 (56·3 ± 11·2) 

♂ 33–69 (54·7 ± 7·9) 

♀ 37–94 (61·3 ± 19·2) 

♂ 27–58 (42·1 ± 9·7) 

♀ 33–74 (54·3 ± 14·5) 

♂ 105–155 (124·0 ± 15·5) 

♀ 61–125 (90·7 ± 32·3) 

Male 


(cm) 

NA 

NA 

18 

43 

84 

54 

48 

53 

119 







Taxon 

Kuwait 

2008 

Kuwait 

2011 Qatar 

Abu 

Dhabi 

Total all 

sites 



Aetomylaeus cf. milvus 16 

(1·06) 

Aetomylaeus nichofii 24 

(1·58) 

— 9 

(0·45) 

1 

(0·14) 

Rhinopteridae 

Rhinoptera javanica — 2 

(0·29) 

Rhinoptera jayakari 5 

(0·33) 

Rhinoptera spp. 121 

(7·98) 

6 

(0·86) 

1 

(0·14) 

69 

(3·42) 

— 25 

(0·54) 

— 94 

(2·02) 

— — 2 

(0·04) 

2 

(0·10) 

91 

(4·51) 

Mobulidae 

Mobula cf. eregoodootenkee — — 2 

(0·10) 

All species combined 1516 

(100) 

699 

(100) 

2019 

(100) 

— 13 

(0·28) 

1 

(0·24) 

214 

(4·60) 

— 2 

(0·04) 

415 

(100) 

4649 

(100) 

♂ 48–90 (68·7 ± 14·1) 

♀ 47–123 (76·0 ± 25·9) 

♂ 35–72 (45·6 ± 6·1) 

♀ 24–72 (51·1 ± 11·8) 

♂ NA 

♀ 99–122 

(110·5 ± 16·3) 

♂ 58–60 (63·5 ± 7·8) 

♀ 43–116 (80·2 ± 17·5) 

♂ 45–85 (69·6 ± 11·0) 

♀ 43–96 (77·1 ± 11·6) 

♂ 105 

♀ ∼100 

NA, not applicable; NAmin, minimum size not available. 

A Probably comprising C. amblyrhynchoides –C. leiodon –C. limbatus and C.limbatus –C. brevipinna not accurately identified. 

B Excised embryo. 

C Unsexed specimens of c. 140–200 cm L T recorded. 

Male 


(cm) 

69 

40 

NA 

NA 

71 

105 




exceptions: (1) two subtly different species of cownose rays Rhinoptera sp., for 

which field identification characters (Last et al., 2010b) only became available at the 

end of the survey, and (2) a number of morphologically similar Carcharhinus species 

in the initial part of the first survey (Kuwait in 2008). Undescribed taxa [whipray 

Himantura sp. B sensu Manjaji, 2004; guitarfish Rhinobatos cf. punctifer (‘RHY’ 

sensu Henderson et al., 2007)] were also recorded, as was a cryptic species complex 

[Himantura uarnak (Gmelin 1789)]. Preliminary examination of barcode data, 

photographs and specimens indicates that taxonomic clarification is also required for 

several more batoid species collected in this survey (White et al., 2010; R. D. Ward, 

W. White, P. Last, pers. comms.). 

TAXONOMIC COMPOSITION 

A total of 39 elasmobranch species (including the H. uarnak species complex) 

were recorded during the survey, comprising 21 shark, five guitarfish and 13 ray 

species (Table I). These belonged to a total of 13 families, of which the most 

species-rich were whaler sharks (Carcharhinidae, 14 species) and stingrays (Dasyatidae, 

five species). The vast majority (c. 80%) of elasmobranchs were sharks, with 

rays comprising c. 16%; guitarfishes were relatively unimportant overall (c. 4%). 

Amongst the sharks, carcharhinids dominated (c. 73% of total individuals), with 

weasel sharks (Hemigaleidae) and carpetsharks (Hemiscyllidae) comprising c. 3% 

each. Houndsharks (Triakidae) and hammerhead sharks (Sphyrnidae) each comprised 


(a) 

(b) 

(c) 

(d) 

Fig. 2. Broad taxonomic composition of elasmobranchs recorded at each sampling event: (a) Kuwait 2008, 

(b) Qatar 2009, (c) Abu Dhabi 2010 and (d) Kuwait 2011. , Chiloscyllium arabicum; , Triakids & 

Hemigaleids; , small (100 cm maximum 

L T ) carcharhinids and sphyrnids; , guitarfishes; ,rays. 

further species was recorded beyond approximately three quarters (c. 1500) of the 

total individuals recorded, suggesting that these surveys adequately recorded species 

richness of landings at the time of the survey. Conversely, new taxa were recorded 

throughout the much smaller survey in Abu Dhabi, indicating that species richness 

was insufficiently sampled at this location. 

SIZE, MATURITY AND SEX COMPOSITION 

Table I presents details of species abundance, size ranges for males and females 

and male maturity for the individuals sampled in the 2008–2011 surveys. The vast 

majority (c. 90%) of all sharks recorded were small with an L T between 50 and 

90 cm, and this pattern was broadly consistent across the four sampling events 

(Kuwait 2008 c. 85%; Qatar c. 94%; Abu Dhabi c. 87%; Kuwait 2011 c. 87%) 

(Fig. 2). The sample comprised mostly of species with a small reported maximum 

size of 200 cm L T [comprising 

pigeye Carcharhinus amboinensis (Müller & Henle 1839), spinner Carcharhinus 




(a) 

100 

Small 

non-carcharhinids 

75 

50 

Larger sharks 

Common small 

100 cm max L T 

Mature males (%) 

(b) 

25 

0 

C. arabicum (64) 

M. mosis (19) 

C. macrostoma (36) 

P. randalli (39) 

100 

Benthic 

R. acutus (901) 

R. oligolinx (92) 

L. macrorhinus (151) 

C. dussumieri (732) 

C. macloti (15) 

C. amblyrhynchoides (3) 

C. brevipinna (14) 

C. leucas (20) 

C. amboinensis (26) 

C. leiodon (14) 

C. limbatus (52) 

C. sorrah (150) 

Carcharhinus spp. (36) 

S. mokarran (6) 

Bentho-pelagic 

75 

50 

25 

0 

H. sp. B (110) 

P. sephen (36) 

H. uarnak complex (21) 

G. cf. poecilura (19) 

A. flagellum (22) 

A. cf. ocellatus (7) 

A. nichofii (47) 

A. cf. milvus (15) 

Rhinoptera spp. (83) 

Fig. 3. Percentage of male (a) sharks and (b) rays (see Table I) that were assessed as mature ( )orimmature 

( ). The number of males assessed for each species are presented in parentheses. L T , total length. 

brevipinna (Müller & Henle 1839), blacktip shark Carcharhinus limbatus (Müller 

& Henle 1839) and great hammerhead Sphyrna mokarran (Rüppell 1837)] were relatively 

unimportant in terms of overall abundance (


Table II. Sex ratios of elasmobranchs recorded in surveys of markets and harbours at locations 

in The Gulf deviating significantly from parity in favour of one sex 

Species Male Female 

Mustelus mosis Kuwait 2011*** 

Chaenogaleus macrostoma Kuwait 2011* Qatar** 

Carcharhinus dussumieri Kuwait 2008***, Qatar*** 

Loxodon macrorhinus 

Abu Dhabi*** 

Rhizoprionodon acutus 

Kuwait 2008***, Qatar*** 

Rhizoprionodon oligolinx Kuwait 2011*** 

Himantura sp. B Qatar**, Kuwait 2011* 

Pastinachus sephen Kuwait 2008** 

Gymnura cf. poecilura Kuwait 2011* 

Aetobatus flagellum Kuwait 2011* 

Aetomylaeus nichofii Kuwait 2008** Qatar* 

Rhinoptera spp. Qatar*** Kuwait 2008***, 

Kuwait 2011* 

*P


100 

(a) 

80 

60 

40 

20 

0 

41–50 

80 

(b) 

70 

51–60 

61–70 

71–80 

81–90 

91–100 

101–110 

111–120 

121–130 

131–140 

141–150 

151–160 

161–170 

60 

Frequency 

50 

40 

30 

20 

10 

0 

25 

(c) 

31–35 

36–40 

41–45 

46–50 

51–55 

56–60 

61–65 

66–70 

71–75 

76–80 

81–85 

86–90 

20 

15 

10 

5 

0 

0–10 

11–20 

21–30 

31–40 

41–50 

51–60 

61–70 

71–80 

81–90 

91–100 

101–110 

111–120 

L T (cm) 

121–130 

131–140 

141–150 

151–160 

161–170 

171–180 

181–190 

191–200 

Fig. 4. Total length (L T )-frequency distributions for male ( ) and female ( ) elasmobranchs for 

(a) Carcharhinus sorrah, (b) Rhizoprionodon oligolinx and (c) Rhinobatus granulatus. 

The landed size of both sexes of the commonest shark species varied between sampling 

events. Males of C. dussumieri landed in Qatar were significantly larger compared 

to those in Kuwait in both 2008 and 2011 (Mann-Whitney U-test, P


Mature males (%) 

100 (a) 

100 

(b) 

90 

90 

80 

80 

70 

70 

60 

60 

50 

50 

40 

40 

30 

30 

20 

20 

10 

10 

0 

0 

30 35 40 45 50 55 60 65 70 75 80 85 90 95 50 55 60 65 70 75 80 

100 

90 

80 

70 

60 

50 

40 

30 

20 

10 

0 

(c) 

50 55 60 65 70 75 80 

100 

90 

80 

70 

60 

50 

40 

30 

20 

10 

0 

(d) 

40 45 50 55 60 65 70 

L T (cm) 

Fig. 5. Male maturity ogives for (a) Carcharhinus dussumieri, (b)Loxodon macrorhinus, (c)Rhizoprionodon 

acutus and (d) Rhizoprionodon oligolinx. The sample size (n), total length (L T ) at 50% maturity (L T50 ) 

and coefficients for the logistic model Y = x + (z − x)(1 + e −K(L T −L T50 ) ) −1 used to determine L T50 are 

(a) n = 730, L T50 = 72·1 cm,K = 0·619, x = 0·5, z = 99·6(r 2 = 0·991); (b) n = 147, L T50 = 64·0 cm, 

K = 0·441, x =−3·2, z = 100·2 (r 2 = 0·947); (c) n = 811, L T50 = 61·7 cm,K = 0·714, x = 0·1, z = 

99·1 (r 2 = 0·993); (d) n = 92, L T50 = 53·0, K = 18·45, x =−4·7, z = 97·0 (r 2 = 0·990). 

those in the same location in 2008 (Mann–Whitney U-test, P


between maternal L T and litter size (Pearson r = 0·785, n = 8, P0·05). 

The L T of the embryos and that of the smallest animals recorded in landings indicate 

that birth size locally is c. 36–38 cm L T . A female R. oligolinx of 67·4 cmL T 

contained three embryos (males of 23·6 and 28·2 and a female of 26·2 cmL T ). 

An 83·5 cm L T M. mosis from Abu Dhabi contained 11 fully formed embryos 

(six males 25·5–28·4 cmL T ; five females 26·5–26·7 cmL T ). In addition to five 

embryos, the right uterus also contained a yolk-coloured oval ovum of solid waxy 

material c. 5 cm long, which is unusual (A. Henderson, pers. comm.). Other notable 

opportunistic observations from females of elasmobranch species whose biology is 

poorly known were an aborted recently formed candle (i.e. eggs within a common 

shell) from a 180 cm L T bowmouth guitarfish Rhina ancylostoma Bloch & Schneider 

1801 (Abu Dhabi); mature oocytes of ≤1·4 cm diameter in a 23·2 cmW D scaly 

whipray Himantura imbricata (Bloch & Schneider 1801) in Qatar; several female 

cowtail stingray Pastinachus sephen (Forsskål 1775) of ≥61 cm W D with emergent 

fully formed embryos (Kuwait 2008); several females of Rhinoptera sp. (probably 

Rhinoptera jayakari Boulenger 1895) of ≥80 cm W D with emergent fully formed 

embryos and aborted embryos of 22·5–28·5 cmW D (Kuwait 2008). 

FISHERIES, BY-CATCH, DISCARDING AND UTILIZATION 

The majority of elasmobranchs recorded in this study were landed by small 

(c. 7–10 m) open speedboats operating gillnets to target teleosts in local, coastal 

waters. In addition to small gillnetters, larger (c. 15–20 m) dhows also operated, targeting 

teleosts with either hemispherical wire fish traps (gargoor) or gillnets, but these 

vessels appeared to contribute less to the overall elasmobranch landings recorded. 

In Abu Dhabi, these dhows (reportedly operating in Gulf waters) also landed large 

elasmobranchs from hook-and-line fishing as a targeted but apparently supplementary 

activity. While all quayside landings recorded at Abu Dhabi were caught in The 

Gulf, the retail market sold both locally caught elasmobranchs and those imported 

overland originating from the Gulf of Oman. A small number of small sharks at each 

survey location had apparently been caught by handline, presumably opportunistically 

during gillnetting. In addition, recent footage from Bahrain demonstrated that 

large gargoor (c. 250 cm height) can occasionally capture quite large (c. 150 cm L T ) 

rhynchobatids (A. B. M. Moore & R. Peirce, pers. obs.). 

All elasmobranchs recorded were landed and marketed whole, with no direct evidence 

of removal of fins at sea. Fins were observed being removed from sharks 

and guitarfishes in Kuwait and Qatar both by fishers at the quayside and by stallholders 

upon sale for consumption, apparently for onward sale. Large sharks and 

guitarfishes landed whole in Abu Dhabi were transported overland to other Emirates 

for processing, presumably for the fin market (A. B. M. Moore pers. obs.; R. Jabado 

pers. comm.). While prices were not recorded, sharks were marketed at relatively 

low prices compared to teleosts. Rays appeared to be of particularly low value and 

catches in Kuwait were commonly observed being discarded for disposal (either 

back at sea, in the harbour or in refuse), although rays picked from the gillnets of 

incoming vessels at Doha Corniche, Qatar, were always marketed. Rhynchobatid 

guitarfishes were the only commonly encountered batoid taxa that were apparently 

of some value, both as meat and apparently for fins. Weather conditions appeared 




to have a notable effect on elasmobranch discarding practices in Kuwait, with the 

unsettled weather in April 2011 resulting in nets being emptied at the quayside rather 

than at sea. This factor is likely to explain the high number of low-value C. arabicum 

and R. granulatus recorded in 2011. 

OVERVIEW 

DISCUSSION 

The current study is the first detailed, large-scale survey of the diversity, biology 

and fisheries of Gulf elasmobranchs and provides a foundation for more refined 

characterization. This is despite its limitations in terms of seasonal coverage and relatively 

small sample size compared with other recent works sampling elasmobranch 

landings at (sub-) tropical locations (White & Dharmadi, 2007; Bizarro et al., 2009). 

Detailed assessment of reproductive status was not recorded in this study. The findings 

confirm that the elasmobranch landings in the parts of The Gulf covered by this 

survey are dominated by a few species of common, small carcharhinids which are 

caught largely as by-catch, and the species and size composition of these landings 

are notably different from those reported in adjacent Oman, which also has targeted 

fisheries for large pelagic species (Henderson et al., 2007, 2009). These data also 

confirm regional differences in the reproductive biology of common fisheries species, 

as well as sex-based and sampling event-based differences in the landed size of some 

common species, which may be due to fisheries or biological factors. Together, these 

highlight the need for local characterizations of elasmobranch fisheries in the western 

Indian Ocean. 

This study further confirms that significant numbers of multiple species of elasmobranchs 

are landed in Kuwait and Qatar, although the FAO does not list any landings 

of elasmobranchs from these countries. Furthermore, Kuwait, Qatar and the U.A.E. 

routinely catch and land elasmobranchs and should therefore strive to develop a 

shark plan, although it should be noted that all other Gulf states (Bahrain, Iran, Iraq 

and Saudi Arabia) have some documentary evidence of elasmobranchs occurring 

in fisheries, particularly as by-catch (Moore, 2011). Given the multiple nations that 

surround The Gulf, a shark plan may be best achieved with a regional approach, as 

was done for the Mediterranean Sea (UNEP, 2003). 

TAXONOMY AND SPECIES RECORDS 

The current study has provided valuable information on the diversity of elasmobranchs 

in the north-western Indian Ocean, an area which is in general poorly 

sampled despite indications that its elasmobranch fauna has significant biogeographic 

and taxonomic interest (Moore, 2011). While 39 taxa were recorded, many of these 

occurred infrequently, and the commercial harvest was dominated by a small number 

of small-sized sharks. The surveys provided a number of new records, the most 

notable of which was the rediscovery of C. leiodon in Kuwait, previously known 

only from a single specimen collected over 100 years ago from 3000 km away in 

Yemen (Moore et al., 2011). The surveys also provided significant range extensions 

for a number of shark (Moore et al., 2010) and batoid species, including the first 




substantiated record of a devil ray (Mobulidae) from The Gulf (unpubl. data). In 

addition, the surveys provided material and data on several taxa in need of description 

or taxonomic resolution that will contribute to an improved understanding of 

Indo-Pacific Ocean elasmobranch diversity. 

PATTERNS OF DIVERSITY AND DISTRIBUTION 

Local patterns of diversity from this study are difficult to compare quantitatively 

when based on samples of differing sizes from landings of different fleets. Based on 

a qualitative comparison of the data (and excluding those species recorded on only a 

handful of occasions), Kuwait was notable in that a number of species (C. leiodon, 

R. oligolinx, R. granulatus and A. flagellum) were only recorded there, which may 

be due to the proximity of the Shatt al Arab. These data and those from other studies 

indicate geographical differences in the abundance of common small carcharhinid 

species such as the prevalence of C. dussumieri along the coasts of Kuwait, Qatar 

(this study) and Iran (Blegvad, 1944 as Carcharias menisorrah), but relative rarity 

off the Omani coast (Henderson et al., 2007; A. Henderson, pers. comm.). Loxodon 

macrorhinus was absent from Kuwait in both 2008 and 2011 but relatively abundant 

in Qatar, Abu Dhabi (this study) and in Oman’s landings (Henderson et al., 2007). 

This may be related to this species apparently preferring low turbidity environments 

(Gutteridge et al., 2011). Rhizoprionodon oligolinx was only recorded (but relatively 

abundant) in Kuwait, where the morphologically similar L. macrorhinus was absent. 

Excluding two species recorded as single individuals from Abu Dhabi that had 

probably originated from the Gulf of Oman [silky shark Carcharhinus falciformis 

(Müller & Henle 1839) and scalloped hammerhead Sphyrna lewini (Griffith & Smith 

1834)] and the H. uarnak species complex, the number of elasmobranch species 

recorded in the current study from The Gulf was 36. Again, excluding H. uarnak, 

this is notably less than the 56 reported from Oman (Henderson & Reeve, 2011) and 

largely reflects the lack of species associated with deeper waters in the shallow Gulf. 

Notable temporal differences were observed in the abundance of two species of 

carcharhinid (R. acutus and C. sorrah) that were common in Kuwait in 2008 but 

were rare or absent at this location in 2011. This absence could be due to a large 

number of variables, possibly including windy conditions at sea in 2011 reducing 

gillnet catches of these species, or temporal variation in estuarine discharges affecting 

distribution. Surface water temperature was probably not a factor, being similar at 

the time of both surveys (22·2 ◦ C in 2008 and 21·4 ◦ C in 2011, measured at the 

same station off east Failaka Island; F. Al-Yamani, unpubl. data). 

SHARK BIOLOGY 

There was a highly significant bias towards (mostly mature) males for both 

C. dussumieri and R. acutus, in both Kuwait in 2008 and Qatar. This may be related 

to spatial segregation from males by birthing or near-term pregnant females, as both 

species are reported as having peaks in parturition in spring or summer in the region 

(Assadi, 2001; Henderson et al., 2006). Interpretation of elasmobranch sex ratio data 

from landings is, however, problematic, as any significant differences may be due 

to a number of confounding factors (e.g. gear bias) as well as natural segregation, 

which is commonly reported in elasmobranchs (reviewed in Sims, 2005). 




Carcharhinus dussumieri in this study attained the maximum reported L T of 

100 cm (Compagno et al., 2005), consistent with previous work locally off Iran 

(Blegvad, 1944, as C. menisorrah) and larger than the maximum of 93·7 cmL T 

recorded off Indonesia (White, 2007). Male maturity size range of C. dussumieri in 

the current study was 63–80 cm L T (all mature animals ≥66 cm L T and an L T50 

of 72·1 cm) and broadly consistent with male size at first maturity (67 cm L T ) and 

L T50 (69 cm) recorded off Iran’s Gulf of Oman coast (Assadi, 2001) but slightly 

smaller than that of c. 74cmL T reported from Indonesia (White, 2007). 

The size at male maturity recorded for C. sorrah in this study (i.e. all mature 

by c. 110 cm L T ) is broadly consistent with that for Indonesia and most other 

locations [recorded and reviewed by White, 2007)]. One female C. sorrah from 

Qatar of 166 cm L T extends the reported maximum size for this species (160 cm 

L T ; Compagno et al., 2005; White, 2007, Henderson et al., 2009). Landings of this 

species were conspicuous in their dominance by juveniles of the 70–85 cm size 

class (c. 77% of all C. sorrah recorded), indicating that this group is particularly 

vulnerable to capture locally in spring. Given a reported birth size of 45–60 cm 

(Compagno et al., 2005) and rapid growth of neonates of c. 20 cm in the first year 

off northern Australia (Davenport & Stevens, 1988), it is probable that this group 

represents individuals in their first year or so of life. 

In Chiloscyllium arabicum maximum L T has been reported as 70 cm (Compagno 

et al., 2005), yet, in this study, males and females were often larger than this (≤77 and 

≤80 cm L T , respectively) which accords with 78 cm L T reported in an earlier study 

from The Gulf (Goubanov & Shleib, 1980). Maturity in this species was reported to 

occur between 44·6 and 54·1 cmL T (Dingerkus & DeFino, 1983 as C. confusum, 

sex not specified), yet in this study all stages were notably larger than this (juvenile 

56 and 62 cm L T , n = 2; maturing 62 and 68 cm L T , n = 2; mature 60–77 cm L T , 

n = 60). In this study, C. arabicum was notable in that it was the only species whose 

size frequency distribution (pooled samples for both males and females) was normal, 

indicating that a wide range of life stages are vulnerable to capture locally in April. 

Loxodon macrorhinus size ranges from the current study are similar to those 

recorded from Oman (Henderson et al., 2009), although the maximum reported size 

from both of these studies is smaller than that of 99 cm L T recorded in Indonesia 

(White, 2007). Further regional differences in the biology of this species are the 

size at male maturity, which in this study (61–71 cm L T ;64·0 L T50 ) is notably 

smaller than that recorded from Indonesia (80–83 cm L T ,81·9 cmL T50 ; White, 

2007) and southern Africa (73–75 cm L T ; Bass et al., 1975). The highly significant 

bias towards (large, mature) males in the April sample at Abu Dhabi is consistent 

with notable sex and size segregation reported for this species from longline surveys 

in the Maldives (Anderson & Ahmed, 1993). 

The average size of R. acutus at Abu Dhabi (probably originating from the Gulf 

of Oman) was significantly greater than at other locations, and the mean size for 

both males and females of R. acutus from Oman (Henderson et al., 2009) was also 

greater than that found in this study. While this may simply represent gear bias, 

it may also indicate that populations of this species from The Gulf and the Gulf 

of Oman have different size characteristics. The L T50 for males of R. acutus from 

the current study (61·7 cmL T ) was also smaller than that from Oman (64·7 cmL T ; 

Henderson et al., 2006), Indonesia (71·8 cmL T ) and other locations in the Indo-West 

Pacific (White, 2007). Maximum size of R. acutus in this study (89 cm L T ) is at 




least c. 5 cm smaller than that recorded in several other Indo-West Pacific Ocean 

locations (Henderson et al., 2009). 

Rhizoprionodon oligolinx has few published studies on its biology. Males from 

this study were found to mature at a notably larger size (45–53 L T , with L T50 

at 53 cm) than previously reported (i.e. in Indonesia, 43–45 cm L T , with L T50 at 

44·6 cm; White, 2007; 29–38 cm L T , Compagno et al., 2005). Furthermore, females 

from this study attained a greater maximum size (85 cm L T ) than previously reported 

(70 cm L T ; Compagno et al., 2005). The near-term embryo 28·2 cmL T recorded is 

of a similar size to the smallest mature individuals reported elsewhere (Compagno 

et al., 2005). The average size of females of this species was significantly larger 

than males, and in Kuwait in 2011, females (at least some of which were pregnant) 

dominated recorded landings. 

All shark landings were dominated by individuals of 50–90 cm L T , with relatively 

few recorded above or below this size. The elasmobranch most commonly recorded 

(R. acutus) is known to have localized abundance of free-swimming neonates of 


were often discarded as unmarketable and took considerable effort to remove from 

gillnets by fishers (A. B. M. Moore, pers. obs.), suggesting that any measures developed 

to reduce this by-catch would be favourably received. Excluding guitarfishes, 

the rarity of rays encountered at Abu Dhabi is consistent with year-round fisheries 

statistics for this port (Environment Agency Abu Dhabi, 2010) and probably reflects 

bias from fisheries or discarding practices. Species composition of batoids in Iranian 

waters of the easternmost Gulf was reported by Vossoughi & Vosoughi (1999), who, 

notwithstanding probable nomenclatural errors, found a broadly similar suite of taxa 

to that reported in this study. 

Although from separate years and potentially confounded by species identification, 

the highly significant sex biases recorded for Rhinoptera spp. towards females 

(in Kuwait in 2008) and males (Qatar) are of interest and may reflect real geographical 

differences. Indicative data from this study suggests male and female maturity 

at c. 70 and 80 cm W D, respectively. Data from Kuwait in 2008 and Qatar are 

notable in that smaller (and probably immature) males and females of 75 cm W D of just one sex occurred. In 

Kuwait in 2008, all animals >75 cm W D were female and were an order of magnitude 

more abundant than males of any size. As several of these females were 

pregnant with well-developed embryos, this observation may be related to parturition. 

In Rhinoptera species elsewhere, sexual segregation of adults around the time of 

parturition has been reported (Smith & Merriner, 1987), as has migration into areas 

at similar latitudes in April (Goodman et al., 2011) and large schools consisting 

mostly of pregnant females off southern India (James, 1962). The schooling nature 

of Rhinoptera species, as well as limiting life-history characters (e.g. low fecundity, 

Bizarro et al., 2006) may make them particularly vulnerable to fisheries impacts. 

FISHERIES MONITORING AND MANAGEMENT 

These data suggest that species, sex and size composition of elasmobranch landings 

in The Gulf can vary markedly both geographically and between years at the 

same location. Additionally, the wide range of water temperature in The Gulf means 

that seasonal variation in abundance and distribution is likely to occur (Moore, 2011). 

With a view to collating data for input to a shark plan, a starting point for basic 

and inexpensive elasmobranch monitoring by Gulf states could therefore be a similar 

survey approach to that of this study in each of the four seasons and on an 

annual basis, but with appropriate consideration of geographical scale where necessary 

(e.g. the extensive coastline of Iran). This might, however, still underestimate 

total fishing mortality by not recording at-sea factors such as discards, as well as possibly 

under-representing valuable large sharks that may be finned at sea and landed 

elsewhere. These data highlight that recording at a species level is essential in any 

future monitoring. While largely accurate in itself, reporting of higher taxa such as 

Carcharhinidae, as is done for Abu Dhabi (Environment Agency Abu Dhabi, 2010) 

will fail to differentiate between small species and juveniles of larger species that 

are likely to be more vulnerable to fisheries. Species-level identification locally is 

especially important as rare, threatened and geographically restricted species occur, 

such as of juveniles of C. leiodon, alongside abundant small carcharhinid species in 

Kuwait. 




The market surveys were supported in Kuwait in 2008 by the Kuwait Ministry of Interior 

(S. Al-Fahad and J. Failkawi), Y. Al-Khorafi, the Kuwait Scientific Centre, Gulf Telecom 

and the Kuwait Institute for Scientific Research (KISR); in Qatar by Qatar University Environmental 

Studies Centre (M. Al-Ansi, I. Al-Maslamani), Qatar Ministry of Environment, 

Department of Fisheries (M. Mohannadi); and in Kuwait in 2011 by the Kuwait Environmental 

Research and Awareness Centre (KERA) and H. Muraad (PAAFR). We thank W. White 

and P. Last (CMAR, Australia), A. Henderson (SQU, Oman), L. Compagno (Shark Research 

Center, Iziko-South African Museum) for confirming identifications of problematic specimens 

and informative discussions; A. Marriott (Bangor University) for statistical help; K. Samimi- 

Namin (Netherlands Centre for Biodiversity Naturalis, Leiden) for the map and F. Al-Yamani 

(KISR) for temperature data. For invaluable assistance with market sampling, we thank 

D. Almojil and A. Alhafez (KERA), R. Jabado (U.A.E. University), A. Reeve (SCS/SQU 

Oman) and volunteers of the SCS (M. Barnes, T. Bennett, M. Boothman, S. Benzie, M. 

Bradfield, S. Collins, G. Gilgannon, D. Green, E.-L. Nichols, S. Nicholls, J. Peirce, C. Sayle, 

M. Sharland, A. Sweeney and M. Webb). Finally, we thank the many fishers and market 

traders for their patience, humour and generous help and the editor and referees for their 

helpful comments on an earlier draft version of the paper. The contents of this paper do not 

necessarily represent the views of the RSK Group. 

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