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Biology of the thornback ray (Raja clavata Linnaeus, 1758) in the North Aegean Sea

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Figure 1

Length–frequency distribution by sex of thornback ray
Length–frequency distribution by sex of thornback ray

Figure 2

Central part of the longitudinal section of R. clavata (by Koray Cabbar)
Central part of the longitudinal section of R. clavata (by Koray Cabbar)

Figure 3

von Bertalanffy length growth curves for R. clavata
von Bertalanffy length growth curves for R. clavata

Figure 4

Seasonal frequency distribution of R. clavata for females (A) and males (B) at maturity stages
Seasonal frequency distribution of R. clavata for females (A) and males (B) at maturity stages

Figure 5

Relationship between total length and clasper length, testicle length, testicle width for males of R. clavata
Relationship between total length and clasper length, testicle length, testicle width for males of R. clavata

Figure 6

Relationship between total length and ovary length (A), ovary width (B) for females of R. clavata
Relationship between total length and ovary length (A), ovary width (B) for females of R. clavata

Figure 7

Relationship between follicle length (cm) and follicle weight (g) for females of R. clavata
Relationship between follicle length (cm) and follicle weight (g) for females of R. clavata

Figure 8

Gonadosomatic index of thornback ray
Gonadosomatic index of thornback ray

Frequency of prey groups for each size class of thornback ray

N Size class (cm) Annelida Cephalopoda Crustacea Teleostei Other
2 40–50 0 0 0.15 0 0.23
29 50–60 0.001 1.71 4.62 9.23 2.8
35 60–70 0.004 2.03 11.91 18.58 4.99
27 70–80 0.002 0.97 6.48 20.56 3.69
11 80–90 0.004 0.28 3.76 7.18 0.84

Diet composition of R. clavata presented as percentage numerical composition (N%), percentage weight of prey items (W%), percentage frequency of occurrence (F%) and percentage index of relative importance (IRI%)

Prey N% W% F% IRI%
Nematoda
  Unidentified nematode 3.37 0.01 5.00 0.80
Cephalopoda
  Loligo sp. 2.25 5.64 5.00 1.88
  Octopus sp. 0.16 0.08 0.45 0.01
Crustacea
Decapoda
  Aegaeon cataphractus 0.16 0.11 0.45 0.01
Natantia
  Caridea 24.88 12.41 17.73 31.42
  Parapenaeus longirostris 4.82 2.63 2.73 0.97
  Solenocera membranacea 0.80 0.29 0.45 0.02
Stomatopoda
  Squilla mantis 4.17 7.14 3.64 1.96
  Erugosquilla massavensis 0.16 0.02 0.45 0.00
Brachyura
  unidentified Brachyura 29.53 7.71 10.45 18.51
  Medorippe lanata 0.96 0.79 1.36 0.11
  Gonoplax rhomboides 0.32 0.19 0.45 0.01
  Anapagurus petiti 0.48 0.07 0.45 0.01
  Athelecyclus roduntatus 0.16 0.19 0.45 0.01
Teleostei
  Boops boops 0.48 3.47 0.45 0.09
  Merluccius merluccius 0.48 1.27 0.91 0.08
  Gobius sp. 0.32 0.70 0.45 0.02
  Trachurus trachurus 1.77 9.12 4.55 2.35
  Micromesistius poutassou 0.64 4.29 0.91 0.21
  Mullus barbatus 0.96 10.27 2.73 1.46
  Mullus surmuletus 0.16 1.37 0.45 0.03
  Sardina pilchardus 0.32 2.23 0.91 0.11
  Spicara smaris 0.32 3.18 0.45 0.08
  Trachurus mediterraneus 0.16 0.30 0.45 0.01
  Engraulis encrasicolus 1.12 1.28 0.45 0.05
  Gadiculus argenteus 0.16 0.22 0.45 0.01
  Triglidae 0.48 1.87 0.45 0.05
  Serranus hepatus 0.16 0.62 0.45 0.02
  Unidentified fish 8.19 9.20 12.27 10.14
Other 12.04 13.32 24.55 29.59

Von Bertalanffy growth parameters for R. clavata

Sex Growth parameters
L (cm) (95% CI) K (year−1) (95% CI) t0 (year) (95% CI)
Males 101.71 (53.67−149.76) 0.18 (−0.060 to 0.410) −0.07 (−2.910 to 2.779)
Females 106.54 (73.18−139.89) 0.16 (0.029−0.286) −0.28 (−1.981 to 1.427)

Comparison of von Bertalanffy growth parameters by region

Region N Sex L (cm) K (year−1) t0 (year) Maximum age (year) Study
UK waters M 88.3 0.22 −1.30 Taylor & Holden 1964
F 127.3 0.10 −2.50
Irish Sea and Bristol Channel M 85.6 0.21 −0.60 Holden 1972
F 107.0 0.13 −0.60
Carmarthen Bay, United Kingdom M + F 139.2 0.09 −2.63 12 Ryland & Ajayi 1984
Irish Sea, Northeast coast M + F 105.0 0.22 0.45 Brander & Palmer 1985
Irish Sea, southeast coast M 96.8 0.19 −1.36 Fahy 1991
F 107.8 0.15 −1.01
Italy, Sicily Channel 200 M 116.7 0.106 −0.41 Cannizzaro et al. 1995
224 F 126.5 0.098 −0.51
North Sea M 98.0 0.17 −0.43 Walker 1999
F 118.0 0.14 −0.88
Irish Sea 258 M 106.5 0.14 −1.74 8 Gallagher et al. 2005
F 139.5 0.093 −1.84 8
Caernarfon Bay, North of Wales 54 M 100.9 0.18 −0.95 7 Whittamore & McCarthy 2005
135 F 117.6 0.16 −0.70 9
Portugal 85 M 121.5 0.11 −0.11 Serra-Pereira et al. 2005
115 F 130.5 0.10 −0.13
200 M + F 130.5 0.10 −0.14 10
North-East Mediterranean 53 M 61.9 0.22 −1.79 Yeldan et al. 2008
37 F 95.7 0.92 −2.60
90 M + F 79.7 0.13 −2.70 8
North Aegean Sea, Saros Bay 98 M 79.99 0.23 −1.09 7 Yığın 2010
128 F 95.00 0.15 −1.10 8
226 M + F 99.99 0.15 −1.08 8
Gulf of Gabes, Tunisia 530 M 100.8 0.14 1.13 12 Kadri et al. 2014a
750 F 114.6 0.11 1.23 15
West Algerian coast 166 M 100.0 0.35 −0.68 Adda-Hanifi et al. 2017
397 F 105.0 0.46 −0.20
North Aegean Sea 28 M 101.71 0.18 −0.07 8 this study
76 F 106.54 0.16 −0.28 8

Age frequency distribution for R. clavata from the North Aegean Sea

Age Males Females Combined
Groups n % n % n %
4 5 25 12 19.3 17 20.7
5 6 30 20 32.3 26 31.7
6 2 10 12 19.4 14 17.1
7 3 15 11 17.7 14 17.1
8 4 20 7 11.3 11 13.4

Length–weight relationship parameters of thornback ray by region

Study Region Sex N Length(cm) Weight (g) Relationship Parameters
Min. Max Min. Max a b r2
Filiz & Mater 2002 North Aegean Sea M + F 31 20.5 99.0 28.9 2614.3 0.0016 3.29 0.93
Borges et al. 2003 Southern Portugal M + F 13 13.7 54.0 7.8 758.2 0.0014 3.36 0.99
Mendes et al. 2004 Coast of Portugal M + F 63 30.6 86.2 110 4590 0.0025 3.23 0.96
Demirhan et al. 2005 Southeast Black Sea M 23 48.0 95.0 620 5000 0.0005 3.02 0.96
F 29 34.3 88.2 170 5450 0.0003 3.70 0.94
Rosa et al. 2006 Azores Archipelago M + F 404 37.0 89.0 270.0 5110.0 0.0058 3.02 0.93
İşmen et al. 2007 Gulf of Saros M + F 112 6.0 60.0 5 4610 0.01300 3.12 0.99
İlkyaz et al. 2008 Aegean Sea M + F 24 29.0 51.3 0.0335 2.89 0.96
Yeldan et al. 2008 Northeast Mediterranean M + F 90 29.3 64.6 110 1120 0.0034 3.10 0.99
Yığın & İşmen 2009 North Aegean Sea M 98 11.0 76.0 6.30 2900 0.00146 3.34 0.99
F 128 10.0 88.0 5.00 4622 0.00181 3.31 0.99
M + F 226 10.0 88.0 5.00 4622 0.00163 3.32 0.99
Bök et al. 2011 Marmara Sea M + F 24 12.2 70 0.015 2628 0.00001 2.87 0.89
Yemişken 2017 Gulf of Iskenderun M + F 66 6.1 42.6 2 1387 0.0100 3.16 0.99
M 27 6.8 33.8 4 648 0.110 3.12 0.99
F 39 6.1 42.6 2 1387 0.0096 3.17 0.99
Around Gökçeada M + F 19 6.8 46.3 6 2650 0.0127 3.13 0.80
In this study North Aegean Sea M + F 104 43.1 89.9 588.7 5192.8 0.0041 3.10 0.93
M 28 43.1 82.7 588.7 3492 0.0223 2.68 0.88
F 76 50.2 89.9 654.1 5192.8 0.003 3.18 0.95
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Chemistry, other, Geosciences, Life Sciences