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Review Article Mediterranean Marine Science Indexed in WoS (Web of Science, ISI Thomson) and SCOPUS The journal is available on line at http://www.medit-mar-sc.net DOI: http://dx.doi.org/10.12681/mms.659

Age at maturity of Mediterranean marine A.C. TSIKLIRAS1 AND K.I. STERGIOU1,2 1 Laboratory of Ichthyology, Department of Zoology, School of Biology, Aristotle University of Thessaloniki, UP Box 134, 541 24, Thessaloniki, Greece 2 Institute of Marine Biological Resources and Inland Waters, Hellenic Centre for Marine Research, Aghios Kosmas, 16604, Athens, Greece Corresponding author: [email protected] Handling Editor: Konstantinos Tsagarakis Received: 3 October 2103; Accepted: 22 July 2014; Published on line: 26 September 2014.

Abstract

In this review, we collected data on the age at maturity (tm) and maximum reported age (tmax) for 235 stocks of Mediterranean marine fishes, belonging to 82 , 37 families, 12 orders and 2 classes ( and Elasmobranchii). Among Actinop- terygii (mean tm ± SD = 2.20 ± 1.43 y, n = 215), tm ranged from 0.3 y, for the common goby Pomatoschistus microps, to 12 y, for the dusky grouper Epinephelus marginatus, while among Elasmobranchii (mean tm ± SD = 5.94 ± 2.47 y, n = 20), tm ranged between

2.7 y, for brown ray Raja miraletus, and 12 y for the picked dogfish Squalus acanthias. Overall, tmax ranged between 1 y, for the transparent goby Aphia minuta, and 70 y, for the americanus. Mean tmax of Actinopterygii (tmax ± SD = 10.14

± 9.42 y) was lower than that of Elasmobranchii (tmax ± SD = 14.05 ± 8.47 y); tm exhibited a strong positive linear relation with 2 tmax for both Actinopterygii (logtm = 0.58 x logtmax – 0.25, r = 0.51, P < 0.001) and Elasmobranchii (logtm = 0.67 x logtmax – 0.006, 2 r = 0.51, P = 0.007). Mean tm/tmax did not differ significantly with sex within Actinopterygii (ANOVA: F = 0.27, P = 0.60, n = 90; females: mean ± SD = 0.276 ± 0.143; males: mean ± SD = 0.265 ± 0.138) and Elasmobranchii (ANOVA: F = 1.44, P = 0.25, n = 10; females: mean ± SD = 0.499 ± 0.166; males: mean ± SD = 0.418 ± 0.133). Finally, the dimensionless ratio tm/tmax was significantly lower (ANOVA: F = 31.04, P < 0.001) for Actinopterygii (mean ± SD = 0.270 ± 0.135, n = 180) than for Elasmobranchii, (mean ± SD = 0.458 ± 0.152, n = 20), when stocks with combined sexes were excluded from the analysis. Keywords: Age at maturity, maximum age, empirical equations, life history.

Introduction because it requires either ageing of caught individuals or knowledge of the local growth parameters in order for t The reproductive life history characteristics of stocks, m to be estimated from the corresponding length at maturity. such as spawning period (Tsikliras et al., 2010), length Therefore, the literature on the variability of tm among at maturity (Tsikliras & Stergiou, 2014) and fecundity families and orders of is rather scarce. Jennings et al. (Despoti & Stergiou, 2013) are important for assessing (1998) analyzed the life history traits of 18 fish stocks and the effects of fishing on populations and ecosystems report that stocks with high t , such as rays and skates, (Jennings et al., 1998). Age at maturity (t ) is a key m m tend to decline more than expected from their rates of fish- element of the life history strategies of fishes and has been ing mortality. Recently, Drazen & Haedrich (2012) ana- widely used in modelling and grouping fish species based lyzed the life history characteristics of 41 deep- demer- on their traits (Winemiller & Rose, 1992; Rochet, 2000; sal fishes and report that mt increases with depth. King & McFarlane, 2003), as well as a stress indicator This review article is the third in the series of reviews for fisheries (Trippel, 1995). Olsen et al. (2004) have on the reproductive biology of Mediterranean marine fish- shown that simultaneous decreases in growth rate and es, the first being by Tsikliras et al. (2010) on their spawn- tm are evidence of fishery-induced evolution in Atlantic ing period, and complements the review on the size at ma- cod (Gadus morhua) following population decrease that turity of Mediterranean fishes (Tsikliras & Stergiou, 2014). has been attributed to overfishing. The mt is also essential The aim of the present work was to collect the avail- in demographic analyses (Chen & Yuan, 2006) being able data on the age at maturity (tm, y) and maximum re- the lower limit of generation time (= the mean age of ported age (tmax, y) for Mediterranean marine fish stocks the spawning stock), which is highly correlated with the and study: (a) the phylogenetic (between classes), sexual intrinsic rate of population growth (Ainsley et al., 2011). (between sexes) and habitat (among habitats - only for

The availability of tm estimates is limited compared Actinopterygii) variability of tm, (b) the relationship be- to length at maturity data (Tsikliras & Stergiou, 2014) tween tm and tmax per class and (c) the dimensionless ratio

Medit. Mar. Sci., 16/1, 2015, 5-20 5 tm/tmax per class and sex, which shows the proportion of Results lifespan that is covered before and after maturation. The were the most represented order (128 compilation of such a dataset is also important for setting stocks; 54.5% of the total) of the dataset, the historical baselines against which future estimates of t m most represented family (38 stocks; 16% of the total) and can be compared in order to define trends in t with time m Trachurus trachurus the most represented species (17 (i.e. shifting baselines: Pauly, 1995). stocks) (Table 1).

Overall, we collected data on tm for 235 stocks, be- Materials and Methods longing to 82 species, 37 families, 12 orders and 2 class- es (Table 1). Among Actinopterygii (n = 215), t ranged The available and accessible information on t of m m from 0.3 y, for the common goby Pomatoschistus mi- Mediterranean marine fishes were extracted from - jour crops, to 12 y for the dusky grouper Epinephelus mar- nals, conference proceedings, theses, and technical re- ginatus, while among Elasmobranchii (n = 20), it ranged ports for the period up to December 2013. The dataset between 2.7 y, for the brown ray Raja miraletus, and 12 also includes some stocks from the Marmara Sea, SE y for the picked dogfish Squalus acanthias. The mean , Suez Canal, and Bay of Cadiz because of their tm ± SD was 2.20 ± 1.43 y for Actinopterygii and 5.94 close proximity to the openings of the ± 2.47 y for Elasmobranchii (Table 1). The tmax ranged (Table 1). Fish were classified in classes (Actinopterygii: between 1 y, for the transparent goby Aphia minuta, and actinopterygians or ray-finned fishes; Elasmobranchii: 70 y, for the wreckfish Polyprion americanus. Mean tmax elasmobranchs or sharks, rays and skates), orders and was lower for Actinopterygii (tmax ± SD = 10.14 ± 9.42 y) families based on Eschmeyer (2012). Valid species compared to Elasmobranchii (tmax ± SD = 14.05 ± 8.47 y). names and authorities were according to FishBase (www. The frequency distribution of tm was unimodal for both fishbase.org; Froese & Pauly, 2014). All ages were con- classes, with a peak at 2 y for Actinopterygii and a peak verted to years (y). at 5 y for Elasmobranchii (Fig. 1).

In the literature, there are several measures or defi- The tm exhibited a strong positive linear relation with nitions of sexual maturity (Binohlan, 1998; Tsikliras & tmax for both Actinopterygii and Elasmobranchii (Fig. 2). Stergiou, 2014). When it comes to age at maturity, the The slopes of the relations did not differ between Actin- most common definition is the median or mean age at opterygii and Elasmobranchii (ANCOVA: F = 0.16, P = which 50% of the population becomes mature for the 0.69) but the intercepts did (ANCOVA: F = 72.45, P < first time, i.e. the age of first maturity (Tsikliras et al., 0.001). The empirical equations on the logarithmic val- 2013a). ues were (the non-transformed equations are also given We also collected information on t (also in y) ei- for comparability purposes): max Actinopterygii logt = 0.58 x logt – 0.25, r2 = ther from the original article or when not available from m max 0.51, P < 0.001, n = 215 FishBase (Froese & Pauly, 2014) and consequently we (tm = 0.11 x tmax + 1.10) computed the dimensionless ratio tm/tmax. Additional in- formation on the sampling procedure and habitat of each stock was recorded and, for the vast majority of stocks, is given in detail in Tsikliras & Stergiou (2014). Such in- formation included country and specific location of sam- pling, duration and frequency of sampling, sex of speci- mens, number of individuals collected and sampling gear (Table 1). The habitat of each species was assigned based on the habitat information provided in FishBase (Froese & Pauly, 2014): demersal (D), pelagic (P), reef-associat- ed (R), benthopelagic (BEP), bathydemersal (BAD) and bathypelagic (BAP). Although there are some inaccura- cies regarding habitat allocation in FishBase, we decided to follow the habitat preferences exactly as they appear in FishBase for comparability purposes with previous ar- ticles and FishBase itself. Analysis of variance (ANOVA) and covariance (AN- COVA) were used to compare means and regression lines, respectively. Fisher’s Least Significance Differ- Fig. 1: Percentage frequency distribution of age at maturity (tm, ence (LSD) test was used to test for differences in the y) of Mediterranean marine stocks per class. Grey bars refer to means between variables in multiple comparisons. Actinopterygii (n=215) and black bars to Elasmobranchii (n=20).

6 Medit. Mar. Sci., 16/1, 2015, 5-20 (continued)

: maximum maximum : max (2006) (2001) (2011) (2011) (2004) (1996) et al. et al. et al. et al. et al. et al. Reference : age at maturity (y), t (y), maturity at age : D’Onghia Ismen (2003a) Ismen (2003a) Uckun Akyol Mouhoub (1986) Mouhoub (1986) & Koutrakis (2013) Tsikliras & Koutrakis (2013) Tsikliras Antonopoulou (2006) & Tsikliras Antonopoulou (2006) & Tsikliras Bouaziz Osman Osman Millán (1999) Millán (1999) Millán (1999) Millán (1999) Millán (1999) Millán (1999) Millán (1999) Millán (1999) Bouaziz & Bennoui (2004) Froglia & Gramitto (1981) Politou & Papaconstantinou (1991) Politou & Papaconstantinou (1991) Froglia (1981) Zoubi (2001) m / m max t t 0.38 0.13 0.14 0.25 0.25 0.25 0.25 0.25 0.22 0.21 0.27 0.30 0.67 0.67 0.25 0.25 0.25 0.25 0.25 0.25 0.25 0.25 0.50 0.13 0.24 0.22 0.20 0.46 max t 8 8 7 8 4 4 4 4 4 5 5 5 3 3 4 4 4 4 4 4 4 4 4 8 5 6 5 9 m t 3.0 1.0 1.0 2.0 1.0 1.0 1.0 1.0 0.9 1.0 1.3 1.5 2.0 2.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 2.0 1.0 1.2 1.3 1.0 4.1 max L 19.2 30.0 29.0 57.5 15.6 19.5 17.5 17.3 16.2 24.8 24.3 25.5 25.0 25.0 18.0 18.0 18.0 18.0 18.0 18.0 18.0 18.0 19.0 32.0 25.0 22.3 25.0 66.0 N 3342 234 368 240 1271 2849 1993 500 500 5467 656 737 1326 469 505 751 1181 564 612 1502 1011 887 423 Sex F F M F C F M F M F M F F M F F F F M M M M C C F M C C values were taken from FishBase (Froese & Pauly, 2014). Pauly, & (Froese FishBase from taken were values

Year max 2000 1999-2000 1999-2000 1997 1993-1996 2000-2002 2000-2002 2000-2002 2000-2002 1995-1996 2008 2008 1989 1990 1991 1992 1989 1990 1991 1992 1986-1988 1986-1988 1995-1998 and t and max Habitat BAD R R P P P P P P P P P P P P P P P P P P P P BAP BEP BEP BEP D Location Ionian Sea Levantine Sea Levantine Sea Aegean Sea Aegean Sea Aegean Sea Aegean Sea Aegean Sea Aegean Sea Alexandria coast Alexandria coast Bay of Cadiz Bay of Cadiz Bay of Cadiz Bay of Cadiz Bay of Cadiz Bay of Cadiz Bay of Cadiz Bay of Cadiz Aegean Sea Aegean Sea Adriatic Sea coast Mediterranean Species Chlorophthalmus agassizi Chlorophthalmus Saurida undosquamis Saurida undosquamis Belone belone Sardina pilchardus Sardina pilchardus Sardina Sardina pilchardus Sardina pilchardus Sardina pilchardus Sardina aurita aurita Sardinella aurita Sardinella Etrumeus sadina Etrumeus sadina Engraulis encrasicolus Engraulis encrasicolus Engraulis encrasicolus Engraulis encrasicolus Engraulis encrasicolus Engraulis encrasicolus Engraulis encrasicolus Engraulis encrasicolus Engraulis encrasicolus poutassou Micromesistius capelanus Trisopterus capelanus Trisopterus Merluccius merluccius ratio. Nomenclature based on FishBase. Italicized L Italicized FishBase. on based Nomenclature ratio. max /t m Family ) at maturity of Mediterranean marine fish stock and location of study. Habitat (D: demersal, P: pelagic, R: reef-associated, BEP: benthopelagic, BAD: bathydemersal, BAP: BAP: bathydemersal, BAD: benthopelagic, BEP: reef-associated, R: pelagic, P: demersal, (D: Habitat study. of location and stock fish marine Mediterranean of maturity at ) /Order/ m Age (t Age CLASS ACTINOPTERYGII Aulopiformes Chlorophthalmidae Synodontidae Beloniformes Belonidae Engraulidae Merluccidae Νο 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 bathypelagic), Year: years of sampling, Sex (F: females, M: males, C: combined sexes), N: number of specimens examined; Lmax: maximum length (cm), t (cm), length maximum Lmax: examined; specimens of number N: sexes), 1. combined C: Table males, M: females, (F: Sex sampling, of years Year: bathypelagic), t the and (y), age reported

Medit. Mar. Sci., 16/1, 2015, 5-20 7 (continued)

(1998) (1998) (2001) (1998) (1998) (1995) (1995) (2010) (2012) (2012) et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. Reference Bouaziz Garcia-Rodriguez & Esteban (1995) Garcia-Rodriguez & Esteban (1995) Recasens Recasens Mugahid & Hashem (1982) Mugahid & Hashem (1982) Garcia-Rodriguez & Esteban (1995) Garcia-Rodriguez & Esteban (1995) Bouaziz Bouaziz Hotos (1999) Hotos (1999) Minos Okumuş & Başçinar (1997) Okumuş & Başçinar (1997) Koutrakis (1994) Minos (1996) Minos (1996) (2000) Ergene (2000) Ergene Koutrakis (1994) Koutrakis (1994) Katselis (1996) Katselis (1996) Klein & Raventos (2007) Klein & Raventos (2007) Azevedo & Homem (2002) Sley Sley Marino Marino Karlou-Riga (1995) / m max t t 0.32 0.30 0.31 0.44 0.38 0.44 0.44 0.29 0.29 0.38 0.25 0.61 0.70 0.33 0.20 0.13 0.30 0.30 0.30 0.30 0.30 0.75 0.75 0.73 0.52 0.20 0.20 0.40 0.25 0.22 0.27 0.23 0.14 max t 9 9 7 8 8 9 9 7 7 8 8 3 3 15 15 15 10 10 10 10 10 4 4 4 4 5 5 3 11 11 15 15 10 m t 2.9 2.7 2.2 3.5 3.0 4.0 4.0 2.0 2.0 3.0 2.0 1.8 2.1 5.0 3.0 2.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 2.9 2.1 1.0 1.0 1.0 2.8 2.4 4.0 3.5 1.4 max L 65.5 68.0 52.5 50.0 42.0 44.0 41.0 68.0 52.5 60.0 60.0 66.7 50.0 50.0 74.4 66.7 38.6 54.8 42.0 36.9 36.9 30.5 30.5 30.0 24.0 12.1 12.1 14.1 41.4 41.6 150.0 150.0 39.3 N 73 955 502 308 619 81 198 955 502 393 312 32 87 79 39 80 64 114 87 123 55 217 272 122 127 777 891 211 205 369 Sex F F M F M F M F M F M F F M C M F F M F M F M F M F M F M F M F Year 1991-1993 1991-1993 1989-1991 1989-1991 1972 1972 1991-1993 1991-1993 1992-1994 1992-1994 2003-2009 1995 1995 1989-1990 1990-1995 1990-1995 1992-1994 1992-1994 1989-1990 1989-1990 1991-1995 1991-1995 1998-1999 1998-1999 2004-2006 2004-2006 1990-1992 1990-1992 1989-1991 Habitat D D D D D D D D D D D P P P D D P P P P P D D D D R R D R R R R P Location Algerian coast - - Gulf of Lions Gulf of Lions Libyan coast Libyan coast Balearic Sea Balearic Sea Algerian coast Algerian coast Black Sea Ionian Sea Ionian Sea Aegean Sea Black Sea Aegean Sea Ionian Sea Ionian Sea Levantine Sea Levantine Sea Aegean Sea Aegean Sea Ionian Sea Ionian Sea Mediterranean NW Mediterranean NW Gulf of Gabes Gulf of Gabes S Mediterranean S Mediterranean Aegean Sea Species Merluccius merluccius Merluccius merluccius Merluccius merluccius Merluccius merluccius Merluccius merluccius Merluccius merluccius Merluccius merluccius Merluccius merluccius Merluccius merluccius Merluccius merluccius Merluccius merluccius Liza haematocheila Liza aurata Liza aurata Liza haematocheila Liza haematocheila Liza ramada Liza ramada Liza ramada Liza ramada Liza ramada Liza saliens Liza saliens Liza saliens Liza saliens imberbis Apogon imberbis Parablennius ruber Caranx crysos Caranx crysos Seriola dumerili Seriola dumerili mediterraneus Trachurus Family /Order/ (continued) CLASS

Mugiliformes Mugilidae Perciformes Blennidae Carangidae Νο 29 30 31 32 33 34 35 36 37 38 39 43 40 41 42 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 Table 1. Table

8 Medit. Mar. Sci., 16/1, 2015, 5-20 (continued)

(1993) (2003) (2003) (2003) (2003) (2003) (2003) (2003) (2003) (1996) (1996) (2010) (2010) et al. (1993) (1993) (1993) (2011b) (2011b) . (1993) et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al et al. et al. et al. Reference Karlou-Riga (1995) Alegria (1990) Alegria (1990) Alegria (1990) Alegria (1990) Alegria (1990) Alegria (1990) Korichi (1988) Korichi (1988) Abaunza Abaunza Abaunza Abaunza Abaunza Abaunza Abaunza Abaunza Mytilineou (1988) Mytilineou (1988) Soykan Soykan Stergiou Stergiou Iglesias & Morales-Nin (2001) Iglesias & Morales-Nin (2001) La Mesa (2001) La Mesa (2001) Metin Metin Kovacic (2007) Kovacic (2007) Fouda Fouda Bouchereau Fouda Fouda / m max t t 0.26 0.21 0.26 0.27 0.28 0.25 0.27 0.10 0.10 0.20 0.20 0.19 0.27 0.26 0.09 0.18 0.18 0.20 0.20 0.40 0.40 0.38 0.43 0.70 0.60 0.42 0.42 0.40 0.40 0.35 0.35 0.50 0.50 0.15 0.50 0.50 max t 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 5 5 5 5 5 6 1 1 1 1 5 5 2 2 2 2 2 2 2 m t 2.6 2.1 2.6 2.7 2.8 2.5 2.7 1.0 1.0 2.0 2.0 1.9 2.7 2.6 0.9 1.8 1.8 1.0 1.0 2.0 2.0 1.9 2.6 0.7 0.6 0.4 0.4 2.0 2.0 0.8 0.8 1.0 1.0 0.3 1.0 1.0 max L 33.9 60.0 4.4 9.2 5.4 32.0 32.0 32.0 32.0 32.0 32.0 27.5 27.5 22.0 22.0 37.0 43.0 34.0 37.0 39.0 37.0 14.7 16.8 20.0 20.0 40.0 4.0 3.3 4.1 9.2 5.3 5.6 5.6 4.7 5.2 5.2 N 595 154 150 134 155 201 180 33 67 43 41 85 67 29 60 1870 885 1766 398 1763 1588 168 182 114 100 527 470 402 302 311 115 7363 300 178 Sex F F M F M F M F M F M F M F M F M F M F M F M F M F M F M F M F M C F M Year 1989-1991 1986-1988 1985-1993 2004-2007 2001-2002 1986 1986 1987 1987 1988 1988 2001 2001 2001 2001 2001 2001 2001 2001 1984-1985 1984-1985 2004-2007 2004-2007 1986-1988 1985-1993 1996 1996 2004-2007 2001-2002 1986-1987 1986-1987 1985-1989 1986-1987 1986-1987 Habitat P D P D D P P P P P P P P P P P P P P P P P P P P D P D D D D D D D D D Location Aegean Sea Aegean Sea Balearic Sea Aegean Sea Adriatic Sea Adriatic Sea Adriatic Sea Adriatic Sea Adriatic Sea Adriatic Sea Adriatic Sea Balearic Sea Balearic Sea Ionian Sea Ionian Sea Aegean Sea Aegean Sea Alboran Sea Alboran Sea Ionian Sea Ionian Sea Aegean Sea Aegean Sea Aegean Sea Balearic Sea Adriatic Sea Adriatic Sea Aegean Sea Adriatic Sea Suez Canal Suez Canal Mauguio Lagoon Suez Canal Suez Canal Species Trachurus trachurus Trachurus macrophthalma Aphia minuta Deltentosteus quadrimaculatus vittatus Trachurus trachurus Trachurus trachurus Trachurus trachurus Trachurus trachurus Trachurus trachurus Trachurus trachurus Trachurus trachurus Trachurus trachurus Trachurus trachurus Trachurus trachurus Trachurus trachurus Trachurus trachurus Trachurus trachurus Trachurus trachurus Trachurus trachurus Trachurus trachurus Trachurus Spicara flexuosa Spicara flexuosa Spicara maena Spicara maena Aphia minuta Crystallogobius linearis Crystallogobius linearis Deltentosteus quadrimaculatus Gobius vittatus Pomatoschistus marmoratus Pomatoschistus marmoratus Pomatoschistus microps aegyptia Silhouettea aegyptia Family /Order/ (continued) CLASS

Gobiidae Centracanthidae Νο 62 84 85 90 91 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 86 87 88 89 92 93 94 95 96 97 Table 1. Table

Medit. Mar. Sci., 16/1, 2015, 5-20 9 (continued)

(1988) (1988) et al. et al. (2003) (1978) (1978) (1995) (1995) (2010) (2010) (2010) (2012) . (2001) . (2001) (2006) (2006) et al. (2008) et al. et al. et al. (2013) (2013) et al. et al. et al. et al. et al et al et al. et al. et al. et al. et al. et al. Reference Franco Kara (1997) Kara (1997) Quignard Quignard Wassef & El Emary (1989) Wassef & El Emary (1989) Wassef Zoubi (2001) Jukic & Piccinetti (1981) Jukic & Piccinetti (1981) Lamrini (2010) Lamrini (2010) Mehanna (2009) & Papaconstantinou Vassilopoulou (1995) & Papaconstantinou Vassilopoulou (1995) Renones Renones Ozvarol Ozvarol Ismen (2006) Ismen (2006) Carbonara Dhieb Dhieb Papaconstantinou Papaconstantinou Gil Gil Chakroun-Marzouk & Ktari (2003) Chakroun-Marzouk & Ktari (2003) Ates Corriero et al (2005) Kara & Derbal (1999) Renones Marino Marino / m max t t 0.18 0.20 0.13 0.30 0.17 0.27 0.13 0.29 0.18 0.13 0.33 0.40 0.25 0.33 0.20 0.17 0.20 0.20 0.20 0.20 0.20 0.11 0.21 0.27 0.13 0.25 0.08 0.06 0.12 0.13 0.25 0.20 0.14 0.13 0.10 0.24 max t 5 15 15 15 15 15 15 7 6 8 6 5 6 6 5 6 5 5 5 5 5 70 9 9 8 8 42 42 21 21 4 15 50 50 50 50 m t 0.9 3.0 2.0 4.5 2.5 4.0 2.0 1.9 1.0 1.0 2.0 2.0 1.4 2.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 7.5 1.9 2.4 1.0 2.0 3.5 2.7 2.6 2.6 1.0 3.0 7.0 6.3 5.0 12.0 max L 20.0 60.0 60.0 56.0 56.0 67.0 44.4 24.9 15.4 18.0 30.0 29.0 28.0 29.9 26.4 32.0 32.0 21.1 16.4 17.0 15.1 131.0 34.5 34.5 32.5 32.5 59.2 53.0 71.0 290.3 119.0 129.0 72.0 100.3 N 171 300 227 467 121 232 104 179 113 1385 157 245 343 324 292 34 288 157 163 116 115 127 110 694 501 219 59 44 196 Sex F F M F M F M C M F F M C F M F M F M F M C F M F M F M F M C F M F F Year 2001 1990-1991 1990-1991 1973-1975 1973-1975 1981-1982 1990-1992 2002-2003 1981-1982 1995-1998 1972-1975 1972-1975 2009 2009 2007-2008 1991-1992 1991-1992 1990-1992 2002-2003 1999-2000 1999-2000 1998-2000 1998-2000 1985-1986 1985-1986 2007-2011 2007-2011 1995-1996 1995-1996 2003-2005 1998-2004 1994-1996 1994-1996 1993-1994 1998-2004 Habitat D D D D D D D R D D D D D D D D D D R D D D P P R R BEP BEP D D P P R R R R Location Venice Lagoon Venice Gulf of Annaba Gulf of Annaba Gulf of coast Tunisian Tunisian coast Tunisian Alexandria coast Balearic Sea Levantine Sea Alexandria coast Mediterranean coast Adriatic Sea Adriatic Sea Mediterranean coast Mediterranean coast Mediterranean coast Aegean Sea Aegean Sea Balearic Sea Levantine Sea Levantine Sea Levantine Sea Ionian Sea Gulf of Gabès Gulf of Gabès Aegean Sea Aegean Sea Balearic Islands Balearic Islands Tunis Gulf of Gulf of Tunis Gulf of Sea of Marmara entire Algerian coast Balearic Sea S Mediterranean S Mediterranean Species Zosterisessor ophiocephalus labrax Dicentrarchus labrax Dicentrarchus labrax Dicentrarchus Dicentrarchus labrax Dicentrarchus Dicentrarchus labrax Dicentrarchus Mullus surmuletus Upeneus moluccensis Dicentrarchus labrax Dicentrarchus Mullus barbatus Mullus barbatus Mullus barbatus Mullus surmuletus Mullus surmuletus Mullus surmuletus Mullus surmuletus Mullus surmuletus Mullus surmuletus Upeneus moluccensis Upeneus pori Upeneus pori Polyprion americanus Pomatomus saltatrix Pomatomus saltatrix Sparisoma cretense Sparisoma cretense Argyrosomus regius Argyrosomus regius Argyrosomus Sciaena umbra Sciaena umbra sarda Sarda Thunnus thynnus Epinephelus marginatus Epinephelus marginatus Epinephelus marginatus Epinephelus marginatus Family /Order/ (continued) CLASS

Moronidae Mullidae Polyprionidae Pomatomidae Scaridae Scombridae Serranidae Νο 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 129 132 133 130 131 Table 1. Table

10 Medit. Mar. Sci., 16/1, 2015, 5-20 (continued)

(2007) (2007) (2005) (2005) (2003) (2003) et al. et al. (2007) (2007) (2007) et al. et al. (2011a) (2003, 2011) (2003, 2011) (2003, 2011) (2003, 2011) et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. Reference Dulcic Zoubi (2001) Alegria-Hernández (1990) Alegria-Hernández (1990) Chali-Chabane (1988) Kallianiotis (1992) Kallianiotis (1992) Kallianiotis (1992) Kallianiotis (1992) Morales-Nin & Moranta (1997) Morales-Nin & Moranta (1997) Matić-Skoko Matić-Skoko Mouine Mouine Benchalel & Kara (2010) Benchalel & Kara (2010) Benchalel & Kara (2013) Benchalel & Kara (2013) Beltrano Beltrano Hammoud & Saad (2007) Hammoud & Saad (2007) Akyurt (2003) Türkmen & Akyurt (2003) Türkmen & Vitale Vitale Vitale Vitale Kallianiotis Kallianiotis Zoubi (2001) Somarakis & Machias (2002) Somarakis & Machias (2002) Cherabi (1987) Metin / m max t t 0.29 0.29 0.40 0.30 0.40 0.20 0.20 0.20 0.20 0.08 0.24 0.20 0.16 0.40 0.40 0.40 0.40 0.40 0.40 0.21 0.29 0.29 0.29 0.16 0.14 0.14 0.14 0.13 0.13 0.30 0.21 0.23 0.30 0.29 0.17 0.29 max t 7 5 5 5 5 5 5 5 5 28 28 13 13 10 10 10 10 10 10 7 7 7 7 12 12 12 12 12 12 12 12 8 6 7 6 7 m t 2.0 1.4 2.0 1.5 2.0 1.0 1.0 1.0 1.0 2.2 6.7 2.6 2.1 4.0 4.0 4.0 4.0 4.0 4.0 1.5 2.0 2.0 2.0 2.0 1.7 1.7 1.7 1.6 1.6 3.6 2.5 1.9 1.8 2.0 1.0 2.0 max L 9.0 30.0 34.6 34.0 35.0 35.0 34.1 25.6 23.0 20.0 23.5 18.0 18.0 18.0 18.0 80.0 75.0 23.0 20.0 37.2 34.6 34.6 24.0 30.0 30.0 30.0 30.0 27.7 22.8 34.0 28.8 25.5 27.1 27.1 36.5 24.9 N 1218 37 98 230 142 188 821 110 335 440 778 597 695 456 210 745 780 108 143 98 143 235 209 1612 1626 221 477 101 1190 1717 Sex C M F M F M F C F M F F M F M F M F M F M F M F M F M F M F M C F M F F Year 2002-2003 2002-2004 1997-1998 1997-1998 1997-1998 1997-1999 1995-1998 1957-1958 1957-1958 1988-1990 1988-1990 1988-1990 1988-1990 1993-1995 1993-1995 2000-2002 2000-2002 2002-2004 2005-2006 2005-2006 1997-1999 1997-1999 1999-2001 1999-2001 1998-1999 1998-1999 1997-1998 1997-1999 1995-1998 1988-1991 1988-1991 2002-2007 Habitat D D D D D D D D D D D D D D D BEP BEP BEP BEP D D D D BEP BEP BEP BEP D D D D BEP BEP BEP BEP BEP Location Adriatic Sea Tunis Gulf of Algerian coast Sicilian Channel Sicilian Channel Sicilian Channel Aegean Sea Mediterranean coast Adriatic Sea Adriatic Sea Algerian coast Cretan Sea Cretan Sea Cretan Sea Cretan Sea Balearic Sea Balearic Sea Adriatic Sea Adriatic Sea Tunis Gulf of Algerian coast Algerian coast Algerian coast Sicilian Channel Sicilian Channel Syrian coast Syrian coast Levantine Sea Levantine Sea Sicilian Channel Aegean Sea Mediterranean coast Cretan Sea Cretan Sea Alger Gulf of Aegean Sea Species Serranus hepatus Boops boops Boops boops Boops boops Boops boops Boops boops Boops boops Boops boops Boops boops dentex Dentex dentex annularis sargus Diplodus sargus sargus Diplodus sargus sargus Diplodus sargus sargus Diplodus sargus sargus Diplodus sargus sargus Diplodus sargus Diplodus vulgaris Diplodus vulgaris Diplodus vulgaris Lithognathus mormyrus Lithognathus mormyrus Lithognathus mormyrus Lithognathus mormyrus Lithognathus mormyrus Lithognathus mormyrus Lithognathus mormyrus Lithognathus mormyrus Pagellus acarne Pagellus erythrinus Pagellus erythrinus Pagellus erythrinus Pagellus erythrinus Family /Order/ (continued) CLASS

Sparidae Νο 134 148 149 160 161 162 163 135 136 137 138 139 140 141 142 143 144 145 146 147 150 151 152 153 154 155 156 157 158 159 164 165 166 167 168 169 Table 1. Table

Medit. Mar. Sci., 16/1, 2015, 5-20 11 (continued)

(1997) (1997) (2005) (2009) . (2009) (2003) (2003) et al. et al. (2008) . (2008) (2012) (2012) (2011) (2003) (2003) et al et al. et al. (1988) (1988) (1988) (1988) (2010) (2010) et al. (2011a) et al. et al et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. Reference Metin Pallaoro Mouine Wadie Wadie Wadie Zouari-Ktari Zouari-Ktari Wadie Wadie Wadie Kartas & Bondka (1986) Kartas & Bondka (1986) Orsi Relini Vassilopoulou Vassilopoulou Cau & Deiana (1983) Cau & Deiana (1983) Ilkyaz Ilkyaz Giovanardi & Piccinetti (1981) Afonso-Dias Cau & Deiana (1983) Cau & Deiana (1983) Cau & Deiana (1983) Cau & Deiana (1983) Nouar (2003) Nouar (2003) Ragonese Bradai & Bouain (1991) Bradai & Bouain (1991) Ordines Ordines Colloca Colloca Boudaya / m max t t 0.43 0.13 0.31 0.13 0.13 0.25 0.25 0.13 0.13 0.40 0.20 0.40 0.15 0.09 0.24 0.20 0.15 0.15 0.08 0.36 0.17 0.06 0.40 0.24 0.10 0.10 0.27 0.30 0.30 0.20 0.20 0.10 0.15 0.20 max t 7 15 13 8 8 8 8 8 8 5 5 10 13 11 12 10 13 13 13 8 13 13 9 8 40 40 30 10 10 5 5 21 13 15 m t 3.0 2.0 4.0 1.0 1.0 2.0 2.0 1.0 1.0 2.0 1.0 4.0 2.0 1.0 2.9 2.0 2.0 2.0 1.0 2.9 2.2 0.8 3.6 1.9 4.0 4.0 8.0 3.0 3.0 1.0 1.0 2.0 2.0 3.0 max L 27.8 36.8 27.0 31.6 27.0 28.3 24.9 42.0 42.0 33.0 26.0 334.5 29.5 23.5 16.5 14.7 11.6 10.1 14.5 37.6 14.1 10.7 39.5 38.9 32.0 28.0 57.0 22.9 20.8 12.8 12.8 27.0 27.0 36.0 N 136 601 330 432 516 537 395 3400 347 1847 1422 1009 563 623 664 540 684 85 90 195 Sex M M F F M F M F M F M F F M F M F M C F F M F M F M F F M F M F M F Year 2002-2007 2004 2005-2006 2003-2005 2003-2005 2004-2007 1990-2001 1990-1992 1990-1992 2004-2007 1998-1999 1985-1998 2005-2010 2005-2010 1994-1997 1994-1997 2003-2004 Habitat BEP BEP P BEP P P P P P D D D D P D D BAP BAP D D D D D D BAD BAD D D D D D D D D Location Aegean Sea Adriatic Sea Mediterranean coast Gulf of Tunis Gulf of Mediterranean coast Gulf of Gabès Gulf of Gabès Mediterranean coast Mediterranean coast Tunisian coast Tunisian Aegean Sea Adriatic Sea Tunisian coast Tunisian Ligurian Sea Aegean Sea Aegean Sea Sardinia Sardinia Aegean Sea Algarve Coast Sardinia Sardinia Sardinia Sardinia Algerian coast Algerian coast Sicilian channel Gulf of Gabès Gulf of Gabes Balearic Islands Balearic Islands Sea Tyrrhenian Sea Tyrrhenian Gulf of Gabes Species Pagellus erythrinus Sarpa salpa Spondyliosoma cantharus Sphyraena chrysotaenia Sphyraena chrysotaenia Sphyraena chrysotaenia Sphyraena sphyraena Sphyraena sphyraena Uranoscopus scaber Buglossidium luteum Buglossidium luteum Uranoscopus scaber Xiphias gladius boscii Lepidorhombus Bathysolea profundicola Bathysolea profundicola Buglossidium luteum azevia Microchirus Monochirus hispidus Monochirus hispidus Synapturichthys kleinii Synapturichthys kleinii Helicolenus dactylopterus Helicolenus dactylopterus Scorpaena elongata Scorpaena porcus Scorpaena porcus Scorpaena loppei Scorpaena loppei Chelidonichthys cuculus Chelidonichthys cuculus Chelidonichthys lucerna Family /Order/ (continued) CLASS

Sphyraenidae Xiphiidae Pleuronectiformes Sebastidae Scorpaenidae Νο 170 171 173 172 174 175 176 177 178 179 180 181 182 183 184 185 186 187 188 189 190 191 192 193 194 195 196 197 198 199 200 201 202 203 Table 1. Table

12 Medit. Mar. Sci., 16/1, 2015, 5-20 (2009) . (2009) (1995) (1995) (2007, 2010) (2007, 2010) et al et al. . (2008) (1997) (1997) (2012) (2012) (2010) (2004) (2013) (2013) et al. et al. (2013) (2013) (2012) (2012) . (2010) . (2004) et al. et al. et al et al. et al. et al. et al. et al et al. et al. et al. et al. et al. et al. et al. et al et al. Reference Boudaya Ismen Ismen Eryilmaz & Meric (2005) Eryilmaz & Meric (2005) Papaconstantinou (1982) Colloca Colloca Ilkyaz Ilkyaz Ismen Ismen Ismen (2003b) Ismen (2003b) & Ismen (2010) Yigin & Ismen (2010) Yigin Kadri Kadri Kadri Kadri Coehlo & Erzini (2006) Coehlo & Erzini (2006) Enajjar Enajjar Demirhan & Seyhan (2007) Demirhan & Seyhan (2007) Cannizzaro Cannizzaro Marouani Marouani Megalofonou Megalofonou / m max t t 0.11 0.13 0.14 0.20 0.21 0.30 0.40 0.40 0.40 0.40 0.24 0.22 0.45 0.45 0.89 0.67 0.49 0.50 0.49 0.39 0.64 0.57 0.36 0.29 0.32 0.28 0.36 0.25 0.53 0.37 0.46 0.41 max t 14 15 14 15 14 5 5 5 5 5 17 18 10 10 9 9 12 9 9 7 14 14 14 10 38 38 14 13 14 13 12 12 m t 1.5 2.0 2.0 3.0 3.0 1.5 2.0 2.0 2.0 2.0 4.0 4.0 4.5 4.5 8.0 6.0 5.9 4.5 4.4 2.7 9.0 8.0 5.1 2.9 12.0 10.5 5.1 3.3 7.4 4.8 5.5 4.9 max L 26.0 30.3 21.2 41.5 37.0 16.0 14.0 14.0 14.0 15.0 52.8 42.9 88.0 73.0 100.0 86.5 80.0 65.0 56.0 58.0 88.2 83.2 198.0 198.0 150.0 150.0 92.0 73.5 100.0 100.0 349 330 N 91 199 143 98 45 1429 308 2196 824 60 603 97 110 146 89 90 550 400 95 85 93 94 812 1038 81 71 178 323 Sex M F M F M F F M F M F M F M F M F M F M F M F M F M F M F M F M Year 2003-2004 1999-2000 1999-2000 1996-1997 1996-1997 1976-1978 1985-1995 1985-1995 2004-2007 2006-2008 2004-2007 2006-2008 1999-2000 1999-2000 2005-2007 2005-2007 2007 2007 2007 2007 1999-2000 1999-2000 1985-1991 1985-1991 2004-2005 2004-2005 1998-2003 1998-2003 Habitat D D D D D D D D D BEP D BEP D D BAD BAD D D D D D D D D BEP BEP D D D D P P Location Gulf of Gabes Iskenderun Bay Iskenderun Bay Marmara Sea Marmara Sea Aegean Sea Sea Tyrrhenian Sea Tyrrhenian Aegean Sea Aegean Sea Aegean Sea Aegean Sea Levantine Sea Levantine Sea Aegean Sea Aegean Sea Gulf of Gabes Gulf of Gabes Gulf of Gabes Gulf of Gabes Algarve Coast Algarve Coast Gulf of Gabes Gulf of Gabes Black Sea Black Sea Sicilian Channel Sicilian Channel Gulf of Gabès Gulf of Gabès E Mediterranean E Mediterranean Species Chelidonichthys lucerna Chelidonichthys lucerna Chelidonichthys lucerna Chelidonichthys lucerna Chelidonichthys lucerna cavillone Lepidotrigla cavillone Lepidotrigla cavillone Lepidotrigla cavillone Zeus faber Lepidotrigla cavillone Zeus faber Dasyatis pastinaca Dasyatis pastinaca Dipturus oxyrinchus Dipturus oxyrinchus Raja radula Raja radula Raja miraletus Raja miraletus Raja undulata Raja undulata Rhinobatos cemiculus Rhinobatos cemiculus Squalus acanthias Squalus acanthias Squalus blainville Squalus blainville Squalus blainville Squalus blainville Prionace glauca Prionace glauca Family /Order/ (continued) CLASS

Zeiformes ELASMOBRANCHII Rajiformes Dasyatidae Rajidae Rhinobatidae Squaliformes Squalidae Carcharhiniformes Carcharhinidae Νο 204 205 206 207 208 209 210 211 212 213 214 215 216 217 218 219 220 221 222 223 224 225 226 227 228 229 230 231 232 233 234 235 Table 1. Table

Medit. Mar. Sci., 16/1, 2015, 5-20 13 Fig. 2: The relationship of age at maturity (tm, y) with maxi- Fig. 3: Variability of age at maturity (tm, y) to maximum age mum reported age (tmax, y) of Mediterranean marine fish stocks (tmax, y) ratio per sex (F: females, M: males) within each class per class. Grey dots and dashed trend line refer to Actinop- (Actinopterygii and Elasmobranchii). The rectangular part of terygii (n=215) and black dots and continuous trend line to the plot extends from the lower quartile to the upper quartile. Elasmobranchii (n=20). All values have been logarithmically The horizontal line within each box shows the location of the transformed. median and the cross the location of the mean. The whiskers extend from the box to the minimum and maximum values and

2 circles indicate outliers. Elasmobranchii logtm = 0.67 x logtmax + 0.006, r = 0.51, P = 0.007, n = 20 (D and P) stocks, whereas reef-associated stocks, which (t = 0.23x t + 2.72) m max had the lowest mean t /t , differed significantly from Mean t /t was not significantly different (ANOVA: m max m max demersal and pelagic ones (Fig. 5). F = 0.27, P = 0.60) between female (mean ± SD = 0.276 ± 0.143, n = 90) and male (mean ± SD = 0.265 ± 0.138, Discussion n = 90) Actinopterygii (Fig. 3). Similarly, mean tm/tmax The ranges of tm values reported in the present work was not significantly different (ANOVA: F = 1.44, P = for Mediterranean marine fishes fall within the previously 0.25) between female (mean ± SD = 0.499 ± 0.166, n = reported range of tm values for actinopterygian and elas- 10) and male (mean ± SD = 0.418 ± 0.133, n = 10) Elas- mobranch fishes. Actinopterygii generally mature ear- mobranchii (Fig. 3). Consequently, sexes were combined lier in life compared to Elasmobranchii (Goldman et al., per class and mean tm/tmax was then compared between classes. Mean tm/tmax was significantly higher (ANOVA: F = 31.04, P < 0.001) in Elasmobranchii (mean ± SD = 0.458 ± 0.152, n = 20) compared to Actinopterygii (mean ± SD = 0.270 ± 0.135, n = 180) (Fig. 4).

With respect to habitat, the mean tm/tmax of Actinop- terygii ranged between 0.06 and 0.43 for benthopelagic (BEP: mean ± SD = 0.22 ± 0.086, n = 23), between 0.06 and 0.75 for demersal (D: mean ± SD = 0.29 ± 0.142, n = 107), between 0.09 and 0.70 for pelagic (P: mean ± SD = 0.28 ± 0.143, n = 63), and between 0.10 and 0.27 for reef-associated stocks (R: mean ± SD = 0.19 ± 0.053, n = 16) (Fig. 5). Elasmobranchii (n = 20) as well as bathydemersal (BAD: n = 3) and bathypelagic (BAP: n = 3) actinopterygian stocks were excluded from this comparison because of their small sample size. Overall, Fig. 4: Variability of age at maturity (tm, y) to maximum age mean tm/tmax values were significantly different among the (tmax, y) ratio per class (Actinopterygii and Elasmobranchii), sex- four habitat categories of Actinopterygii (ANOVA: F = es combined. The rectangular part of the plot extends from the 3.67, P = 0.013). Based on Fisher’s LSD test, there were lower quartile to the upper quartile. The horizontal line within no differences in the means between benthopelagic and each box shows the location of the median and the cross the loca- reef-associated (BEP and R), between benthopelagic and tion of the mean. The whiskers extend from the box to the mini- pelagic (BEP and P) and between demersal and pelagic mum and maximum values and circles indicate outliers.

14 Medit. Mar. Sci., 16/1, 2015, 5-20 lived counterparts (Frisk, 2010). Indeed, elasmobranchs are located at the upper end of that spectrum (Fig. 2). A

significant positive relation between longevity (tmax) and

tm has been reported previously for female and male ba-

toids (females: tm = 0.57 · tmax + 1.02; males: tm = 0.57 ·

tmax + 0.47) (Frisk, 2010). The empirical relation between

tm and tmax can be used for estimating tm for species for

which only tmax is known.

The dimensionless tm/tmax ratio shows that actinoptery-

gians mature very early, at around 1/3 of tmax, within their lifespans. In contrast, elasmobranchs mature later, close

to 1/2 of their tmax. Similar results have been reported for several shark and skate stocks with this ratio being gener- ally higher for skates than for other elasmobranch groups and generally (Frisk, 2010). In general, the onset of maturity may depend mainly on age for short-lived Fig. 5: Variability of age at maturity (t , y) to maximum age m species that mature early, and mainly on size for their (tmax, y) ratio of Actinopterygii per habitat category (BEP: benthopelagic, D: demersal, P: pelagic, R: reef-associated). longer-lived, late-maturing counterparts, with devel- Elasmobranchii and the categories bathydemersal (BAD) and opmental or genetic constraints more evident in longer bathypelagic (BAP) were excluded because of their small sam- lived species (Archibald et al., 1983; Roff, 1983). There ple size. The rectangular part of the plot extends from the lower are benefits and costs associated with reproducing early quartile to the upper quartile. The horizontal line within each or late in life. Benefits associated with early maturity in- box shows the location of the median and the cross the location clude increased probability of surviving to reproduce and of the mean. The whiskers extend from the box to the minimum reduced generation time, while the costs of early maturity and maximum values and circles indicate outliers. include reduced survival and fecundity later in life (Roff, 1992). High adult mortality, as a result of fishing, may

2012). For the latter, tm is variable, and has been reported favour earlier maturation (Rochet, 1998). Such a cost of to range from 2 to 26 y in 18 populations of 15 species reproduction, i.e. a trade-off between present reproduc- (Chen & Yuan, 2006). The same range has been reported tive effort and future age-specific reproductive success, by Cailliet & Goldman (2004), with a bimodal distribu- is evident across (Williams, 1966; Bell, 1980). tion with one peak at 5–6 y and a second one at 15–25 y. Although tm might differ between sexes, the fact that,

Higher tm values have been reported for long-lived elas- in the present work, we found no difference in the tm/tmax mobranchs, such as the whale shark Rhincodon typus (> ratio between males and females within both classes, may

22 y) and the picked dogfish Squalus acanthias (25-26 y) simply be attributed to the fact that tm and tmax vary simul- (Chen & Yuan, 2006; Dulvy et al., 2008). Similarly, for taneously and, within species, are determined by the cor- long-lived actinopterygians, such as the bluemouth rock- responding growth and mortality patterns (e.g. Tsikliras fish Helicolenus dactylopterus, ages at maturity higher et al., 2007; Gislason et al., 2010). For instance, a male than 15 y are known (Kelly et al., 1999). In their work, that matures earlier will experience higher future mortali- that reviewed the life history traits of 41 actinopterygian ties and will have a shorter lifespan compared to a female species, Drazen & Haedrich (2012) report that tm ranges that will mature later (e.g. Tsikliras & Koutrakis, 2013), between 2 and 36 y, with six species having tm that ex- i.e. age at maturity may co-vary with maximum age and, ceeds 20 y and five species having maxt that exceeds 100 y. in the same way, size at maturity co-varies with asymptotic

A similar range of ages at maturity (0.5 to 35.5 y) has been length (Beverton, 1992). Thus, the tm/tmax ratio will remain reported by He & Stewart (2001) in their review of 235 the same and bi-maturism will be absent, as has been re- fish stocks, including marine and freshwater finfishes, as ported to occur for the length at maturity to maximum re- well as a few elasmobranchs. It seems that, at least regard- ported length (Lm/Lmax) ratio (Tsikliras & Stergiou, 2014). ing elasmobranchs, and considering their small sample With the exception of reef-associated species, which size in the present work, long-living, late-maturing sharks exhibited the lowest tm/tmax values, the tm/tmax ratio of ac- are rare in the Mediterranean Sea. Given that the larger tinopterygians remained rather constant among the re- elasmobranchs are more susceptible to overfishing (Dulvy maining habitat categories (Fig. 5) and had similar values et al., 2014), this under-representation may be due to their with the overall tm/tmax ratio for actinopterygians. A simi- overexploitation that is known to occur in the Mediterra- lar constancy among habitat categories has been report- nean (Stergiou & Tsikliras, 2011; Tsikliras et al., 2013b). ed for the Lm/Lmax ratio of Mediterranean marine fishes The positive relation between tm and tmax (Fig. 2) (Tsikliras & Stergiou, 2014). Although there are several confirms the general pattern that longer-lived species large-sized, long-lived, and late-maturing reef-associated mature later and grow slower compared to their shorter- species (e.g. groupers) in the Mediterranean Sea, in the

Medit. Mar. Sci., 16/1, 2015, 5-20 15 present dataset these species were rare in favour of small- sar commun Diplodus sargus sargus (Sparidae) des cotes de l’est and medium-sized reef inhabitants (Table 1). This might algerien. Rapport du Congrès de la Commission Internationale pour l’Exploration Scientifique de la Mer Méditerranée, 39, 451. explain the lower tm/tmax ratio for reef-associated species. Drazen & Haedrich (2012) report that t increases with Benchalel, W., Kara, M.H., 2013. Age and growth and reproduc- m tion of the white seabream Diplodus sargus sargus (Linneaus, depth but in our dataset there are no real deep-water spe- 1758) off the eastern coast of Algeria. Journal of Applied Ich- cies given that most of the species are commercial living thyology, 29, 64-70. on the continental shelf. Thus, we could not test this hy- Beverton, R.J.H., 1992. Patterns of reproductive strategy param- pothesis. Besides deep-water species, special or complex eters in some marine fishes. Journal of Fish Biology, cases of reproductive strategies, such as hermaphrodism 41 (Suppl. B), 137-160. and strategies involving spawning migrations were also Binohlan, C., 1998. The MATURITY table. p. 176-179. In: Fish- rare in the dataset. Such strategies could also result in Base 98: Concepts, Design and Data Sources. Froese R., deviations from the general pattern that is presented here. Pauly, D. (Eds). ICLARM, Manila, Philippines. Bouaziz, A., Bennoui, A., 2004. Etat d’exploitation de l’anchois Finally, it should be noted that length at maturity and Engraulis encrasicolus (Linné, 1758) dans la baie d’Alger. tm should be independently estimated in reproductive biol- Rapport du Congrès de la Commission Internationale pour ogy, instead of tm being indirectly estimated from the von l’Exploration Scientifique de la Mer Méditerranée, 37, 318. Bertalanffy growth equation. This is because the growth Bouaziz, A., Bennoui, A., Brahmi, B., 2001. Sur l’estimation trajectory may differ as a result of quicker or slower rates del’etat d’exploitation du merlu Merluccius merluccius of approaching asymptotic lengths among individuals (He (Linnaeus 1847) de la region centre de la cote algerienne. & Stewart, 2001). Hence, maturity can be reached at same Rapport du Congrès de la Commission Internationale pour length and different age and vice versa (Kozlowski, 1996). l’Exploration Scientifique de la Mer Méditerranée, 36, 243. Bouaziz, A., Bennoui, A., Djabali, F., Maurin, C., 1998. Repro- duction du merlu Merluccius merluccius de la région de bou- References Ismail. Cahiers Options Méditerranee, 35, 109-117. Abaunza, P., Gordo, L., Karlou-Riga, C., Murta, A., Eltink, Bouchereau, J.-L., Quignard, J.-P., Joyeux, J.-C., Tomasini, J.-A., A.T.G.W. et al., 2003. Growth and reproduction of horse 1993. Structure du stock des géniteurs de la population de , Trachurus trachurus (Carangidae). 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