Environ Biol Fish (2011) 91:369378 DOI 10.1007/s10641-011-9796-0 Age and growth of the small red scorpionfish, Scorpaena notata Rafinesque, 1810, based on whole and sectioned otolith readings Giuseppe Scarcella & Mario La Mesa & Fabio Grati & Piero Polidori Received: 12 March 2010 / Accepted: 16 March 2011 / Published online: April 2011 # Springer Science+Business Media B.V 2011 Abstract Age and growth of Scorpaena notata from the northern Adriatic Sea were investigated by annual growth increment counts (annuli) Overall, age and growth were estimated from 538 specimens of S notata ranging between 47 and 199 mm TL No clear sexual dimorphism in size was observed Annual deposition of annuli and location of the first annulus have been validated by edge analysis and daily growth increment counts, respectively The estimated age range was between 016 years for female and 14 years for males The estimated values of Von Bertalanffy asymptotic length L1 (cm) and k (years1) were 16.3 and 0.46 for males and 15.6 and 0.35 for females Thus, males of S notata appeared to attain a slightly larger size at faster rate than females The growth performance index ranged between 1.9 and 2.1, which is in the middle of the range observed in other scorpaenids Comparing ageing data of S notata with other Mediterranean scorpaenids, a direct relationship between fish size and growth performance was observed Keywords Adriatic Sea Age and growth Scorpaenidae Scorpaena notata G Scarcella (*) : M La Mesa : F Grati : P Polidori Istituto di Scienze Marine, Consiglio Nazionale delle Ricerche, L.go Fiera della Pesca 2, 60125 Ancona, Italy e-mail: g.scarcella@ismar.cnr.it Introduction The small red scorpionfish, Scorpaena notata (Rafinesque 1810) (Pisces, Scorpaenidae), is a benthic sedentary species distributed in the Eastern Atlantic from the Bay of Biscay to Senegal, off Madeira, Azores and the Canary Islands, in the Mediterranean Sea and the Black Sea, where it is represented by the subspecies Scorpaena notata afimbria (Slasteneko 1939) (Hureau and Litvinenko 1986) Although S notata is considered to be rare in the northern Adriatic Sea (Hureau and Litvinenko 1986), it was frequently found in recent surveys close to natural and artificial hard bottom (Fabi et al 2004; Casellato and Stefanon 2008), playing likely a more important role in the benthic fish community of rocky habitat than previously thought The small red scorpionfish is generally less than 20 cm total length (TL), and inhabits preferably rocky bottoms inside crevices or sea grass meadows, but it is also captured by trawlers operating on sandy bottoms in the proximity of hard substrates (Hureau and Litvinenko 1986; Harmelin-Vivien et al 1989; Morte et al 2001; Relini et al 2007) Several biological aspects of S notata have been studied in the Mediterranean Sea, such as diet (Harmelin-Vivien et al 1989; Morte et al 2001), gonad morphology (Muủoz et al 1996, 2002a, b), fecundity and reproductive cycle (Muủoz et al 2005) and the relationships with artificial and natural habitats (Relini et al 2002; Ordines et al 2009) The only data concerning age and growth are 370 presented in a publication based on scales reading of specimens from the Gulf of Gabes (Bradai and Bouain 1990) and by a recent study of Ordines et al (2009) about the habitat preference of S notata in Balearic Islands The present paper aims to improve the knowledge of ageing procedure for S notata by means of otolith readings, also observed after sectioning, in order to reach a better understanding of the life span and the calculation of the Von Bertalanffy growth parameters of the species, applying also indirect age validation methods to support the reliability of age estimates Material and methods Samples of S notata were collected in the northern Adriatic Sea between July 2004 and November 2008 The study area included both natural reefs consisting of hard substrates and artificial structures, such as offshore gas platforms (Fig 1) Sampling was carried out in the proximity of natural reefs and artificial structures, using a beam trawl with 40 mm cod-end stretched mesh size and trammel nets with 70 and 400 mm stretched mesh size Hauls performed with beam trawl were randomly located over the whole sampling area The beam trawl was generally towed at Fig Map of northern Adriatic Sea with sampling locations Environ Biol Fish (2011) 91:369378 about 4.85.2 knots for 1530 on the bottom during daylight hours Conversely, trammel nets, positioned close to artificial structures and natural reefs, were set at dusk and pulled in at dawn, with an average fishing time of 12 h The size distribution of the sample could be affected by the selectivity of the gears employed Anyway, the beam trawl was towed at a speed of six knots and, hence, its selectivity was strongly reduced by the scarce opening of codend meshes It is acknowledged that codends of such gears are virtually non-selective for the sizes of most finfish (Rotherham et al 2008) In addition, the particular shape of S notata (round body with many spines) strongly reduces the probabilities of escaping thoughout the codend meshes A study on gillnet selectivity carried out in Mediterranean showed that this gear is scarcely selective for Scorpaena porcus (Stergiou and Erzini 2002) As a matter of fact, trammel nets are widely considered less selective if compared to gillnets, and, also in this case, the presence of many spines around the body of scorpenids favour the non-selective way of capture called entanglement On this basis, it can be realistic to assume that the samples collected during fishing survey may be representative of the pupulation at sea Environ Biol Fish (2011) 91:369378 In the laboratory, wet weight (W, 0.1 g), TL, measured to the lowest mm, and sex, determined by macroscopic examination of gonads were recorded for each specimen In small specimens, gonads were observed by a light microscope to aid sex determination The sagittal otoliths were removed from all specimens, cleaned and stored dry The lengthweight relationship of fish was calculated both for the whole population and for each sex The exponential equation W=aTLb, where W is total weight (0.1 g), TL is total length of fish (mm) and a and b are regression parameters, was fitted to the data The equation was linearized applying the log10 transformation of data to estimate the regression parameters An F-test was used to test for difference between males and females in allometric indices (b) (Sokal and Rohlf 1995) The weight of both left and right sagitta was recorded with an accuracy of 0.1 mg and compared using a t-test for paired comparisons (Sokal and Rohlf 1995) As no difference was found (Students t-test, d f.=806, t=1.647, p>0.05), the maximum (ODmax; 0.01 mm) and the minimum diameter (ODmin; 0.01 mm) were measured from one randomly selected sagitta (left or right) on the whole sample, under a stereomicroscope coupled to a video camera using an image analysis software (OPTIMAS 6.5) The relationship between TL and ODmax and ODmin was investigated by linear regression analysis Otoliths were immersed in ethanol with distal face up and the annuli were counted using a binocular microscope under reflected light against a dark background (magnification: ì25 and ì40) The nucleus and the opaque zones of otolith appeared as light rings and the translucent or hyaline zones as dark rings The combination of each opaque and subsequent translucent zone was considered to be an annulus, as observed in other scorpaenids (Massutớ et al 2000; Lúpez Abellỏn et al 2001) In larger fish, the annulation pattern was difficult to discriminate due to considerable otolith thickness As a consequence, these otoliths were embedded in epoxy resin and transversally sectioned Otolith sections were then polished with 0.05 m alumina paste and read under reflected light following the same aforementioned procedure To compare the two readings procedures also in smaller otoliths, a representative sample of them was read directly as a whole and then sectioned As the age estimates were 371 the same, the sectioning practice was carried out only on the thick otoliths of larger (older) fish The count path of annuli in whole otoliths was generally from the nucleus towards the tip of the rostrum, where the deposition of seasonal rings appeared to start Otoliths were firstly read by one reader, without any ancillary data on fish size A second reading was carried out a week later by the same reader When readings differed by one or more annuli, a third reading was made; if the third reading differed from the previuos two, the otolith was discarded The index of average percent error (APE) (Beamish and Fournier 1981), as well as the mean coefficient of variation (CV) (Chang 1982), were calculated to estimate the relative precision between readings To assess the annual nature of ring deposition, i.e the accuracy of age estimates, two different indirect or semidirect methods were applied To validate specimens aged 0, i.e fish with sagittae composed of only an opaque nucleus, some otoliths were randomly selected for microincrements (daily rings) counting Following other studies on scorpaenids (Laidig and Ralston 1995; Massutớ et al 2000), we assumed that microincrements were laid down daily, providing the true age (in days) of aged specimens Otoliths were set in moulds, embedded in epoxy resin and ground to obtain sagittal sections They were then polished with 0.05 m alumina paste and the microincrements counted under a light microscope at magnification ì400630 To validate the seasonality of deposition of opaque and translucent zones, the relative frequency of an opaque zone on the otolith margin was plotted by month (Beckman and Wilson 1995; Panfili and Morales-Nin 2002) The cycle in formation of the opaque and translucent zones should equal year in true annuli (Campana 2001) Once the age estimates were validated, the Von Bertalanffy growth function was fitted to the agelength data using the routine FISHPARM from the statistical package FSAS (Saila et al 1988), which implements the Marquardt algorithm for non-linear least squares parameter estimation The Von Bertalanffy growth parameters (L1 , k and t0) were calculated for each sex and for the whole population (including unsexed fish) Finally, the growth performance index (=2 log L1 ỵ logk) (Munro and Pauly 1983), was calculated to compare the growth of S notata with other scorpaenids 372 Environ Biol Fish (2011) 91:369378 Results Overall, 570 specimens of S notata were collected, 239 females, 302 males and 29 unsexed individuals The sex ratio differed significantly from 1:1, males being more abundant than females (2 =7.3, d.f.=1, p0.1), and nearly isometric Maximum and minimum otolith diameters (ODmax, ODmin) varied linearly with fish length, according to the following equations: ODmax ẳ 0:47 TL ỵ 0:55; n ẳ 570; r2 ẳ 0:89 ODmin ẳ 0:17 TL ỵ 0:65; n ẳ 570; r2 ẳ 0:76 Fig Photograph showing the annulations pattern of Scorpaena notata sagittal otolith, 3-years-old male, 112 mm total length (TL) were difficult to discriminate, so the transverse sections were helpful to obtain a more reliable count of annuli (Fig 4) Microincrement counts to validate fish aged (i.e young of the year) were carried out on ten specimens ranging between 47 and 96 mm TL The sagittae of S notata showed the typical pattern of light and dark alternated microincrements, representing daily growth rings (Fig 5) A continuous series of concentric rings of increasing size, ranging from 1.0 to 3.6 m, were observed from the core to the otolith margin Accessory primordia were also observed in some specimens The age estimates ranged from 140 to 300 days, validating all otoliths aged characterized by an opaque nucleus surrounded by a more or less developed translucent zone The annulation pattern of otoliths was composed of alternating opaque and translucent zones Outside the opaque nucleus, the first 23 rings were wide and easily recognizable, followed by rings of decreasing thickness towards the otolith margin (Fig 3) In larger fish (>150 mm), the outer rings close to the margin Fig Length frequency distribution of Scorpaena notata sampled in the northern Adriatic Sea Fig Photographs of (a) surface and (b) cross-section otolith of Scorpaena notata, 16-years-old female, 165 mm total length (TL) Vertical line indicate plane observed by cross-section Dots denote annulus from 1st to 16th Environ Biol Fish (2011) 91:369378 373 72% of males and females, respectively The Von Bertalanffy growth function was fitted to the agelength data pairs for each sex, and for the whole population (Table 2, Fig 7) The parameters estimated were significantly different between sexes only if considered together, with male showing L1 , k and t0 slightly higher than females (Table 3) The growth performance index (), calculated for each sex, is reported in Table Lengths-at-age, calculated from the Von Bertalanffy growth function, provides estimates of growth increments by age The annual growth rate ranged between 19.2 and 0.2 mm for females and 28.8 and 0.1 mm for males in the estimated age range (Table 4) Fig Photomicrograph of otolith microstructure in the core region, showing the typical pattern of alternating light and dark increments assumed to be deposited daily Scale bar = 10 m The edge analysis, performed on the whole fish sample, validated the annual deposition of each annulus formed by an opaque and translucent zone In particular, the opaque zones were laid down from April to August, whereas the translucent zones were laid down from September to March (Fig 6) Out of 570 otoliths examined, only 32 (approximately 5%) were discarded, because they were either unreadable or provided different age estimates between readings Counting variability indices, CV and APE, were both quite low (11.6% and 8.2%, respectively), indicating that the ageing procedure adopted gave a reasonable level of consistency (or reproducibility) between readings Age estimates ranged between and 14 years for males and between and 16 years for females (Table 1) However, the fish sample was mainly composed of 14-year-old fish, representing 80% and Fig Monthly change in relative frequency of the opaque zone and translucent zone on the otolith margin of Scorpaena notata Discussion Comparing the results of the age reading reported in the present paper with those obtained in other published works on small red scorpionfish, it is possible to discover evident discrepancies (Table 2) The ageing methodology (scales reading) adopted by Bradai and Bouain (1990), as well as the relatively narrow fish size range composed of small individuals, could have led to an underestimate of the life span of S notata from the Gulf of Gabes Indeed, differently from scales, otoliths are one of the few calcified structures that is nonskeletal and their growth is maintained even through periods when somatic growth is nonexistent (Maillet and Checkley 1990; Campana and Thorrold 2001) The advantage of a continuous growth pattern is most evident in studies of old fish, in which annulus counts from scales grossly underestimate those visible in the otolith (Beamish and McFarlane 1995) Instead, the study from Balearic Islands (Ordines et al 2009) was based on otolith readings carried out on a large fish sample (947 individuals), characterized by a wide size range comparable to the Adriatic sample However, age estimates were performed on whole otoliths, which again could have led to an underestimate of age in older (larger) specimens of S notata, considering that generally otoliths of these individuals are extremely opaque and too thick to allow a reliable estimate of rings close to the otolith margin Interestingly, S notata did not show a clear sexual size dimorphism in growth, although males may have a slightly greater size at age than females This is in 374 Environ Biol Fish (2011) 91:369378 Table Age length key of Sorpaena notata sampled in the northern Adriatic Sea, and the number of fish (n) Lenght class (mm) Females Age class (year) 40 10 11 12 13 14 15 16 50 60 70 80 90 3 11 100 110 13 13 120 15 18 1 2 130 10 11 1 140 1 2 150 3 2 1 2 1 1 2 160 170 1 1 180 1 1 190 n 27 44 56 35 11 6 3 1 Males 60 70 80 90 11 100 110 120 20 15 130 18 24 2 140 21 12 150 15 1 160 10 3 1 2 48 12 170 180 n 11 36 56 87 1 2 1 1 2 Table The von Bertalanffy growth parameters, number of specimens, growth performance indices (), size and age ranges observed for Scorpaena notata The asymptotic standard errors of the estimates are shown between brackets Authors Area Method Sex L mm k year1 t0 year n Age ranges years Size ranges mm Present study Northern Adriatic Sea Whole otoliths and sectioned otoliths M 163.2 (3.0) 0.46 (0.5) 1.17 (0.21) 285 2.08 014 65198 F 155.7 (3.3) 0.35 (0.5) 2.06 (0.43) 225 1.92 016 47199 Total 157.2 (2.1) 0.44 (0.4) 1.37 (0.19) 538 2.03 016 47199 Total 139.48 0.116 1.378 85 1.35 16 50112a Bradai and Bouain (1990) Ordines et al (2009) a Gulf of Gabes Scales Balearic Islands Whole otoliths Standard lengths M 179 0.236 1.742 476 1.88 08 40179 F 179 0.195 2.269 471 1.80 06 40179 Total 179 0.216 1.946 947 1.84 08 40179 Environ Biol Fish (2011) 91:369378 375 Table Likelihood ratio test comparison of Von Bertalanffy parameters estimated for males and females of Scorpaena notata sampled in the northern Adriatic Sea d.f = degree of freedom; * = significant at =0.01 Null hypothesis d.f Log residual sum of squares p-values L =L 0.004 2.275 0.131 k =k 0.004 2.223 0.136 t0 =t0 0.007 3.550 0.060 L, k =L, k 0.063 32.172 0.000* L, t0 =L, t0 0.036 18.554 0.000* k, t0 =k, t0 0.009 4.449 0.108 L, k, t0 =L, k, t0 0.093 47.340 0.000* and Casadevall 2002), could be the reason of the absence of sexual size dimorphism As reported for H dactylopterus (Massutớ et al 2000), S notata exhibited a double mechanism of formation of seasonal growth rings, corresponding to different stages of life, namely immature/juvenile and adult fish The change in deposition pattern was Table Estimated values of fish length-at-age for each sex of Scorpaena notata derived from the Von Bertalanffy equations Age Females Males (years) TL (mm) Annual growth TL (mm) Annual growth (mm) (mm) Fig von Bertalanffy growth curves fitted to the length-age data of Scorpaena notata a males, b females and c whole sample contrast to most other Scorpaeniformes, females being often larger than males (Wyllie Echeverria 1986; Lenarz and Wyllie Echeverria 1991).Generally, it is rather common in fish that a larger size in females increase their fecundity, while the same may not be true in males (Berglund et al 1986) In the case of S notata, the documented low fecundity (Muủoz et al 2005), compared to other Scorpaeniformes, such as Trigla lyra (Muủoz 2001) or H dactylopterus (Muủoz 89.1 108.3 19.2 113.2 84.4 28.8 121.9 13.7 131.5 18.3 131.7 9.7 143.1 11.6 138.6 6.9 150.5 7.4 143.5 4.9 155.1 4.7 147.0 3.5 158.2 3.1 149.5 2.5 160.1 1.9 151.3 1.8 161.2 1.1 152.6 1.3 161.9 0.7 10 153.5 0.9 162.4 0.5 11 154.1 0.6 162.7 0.3 12 154.6 0.5 162.9 0.2 13 154.8 0.3 162.9 0.1 14 154.9 0.3 163.0 0.1 15 155.1 0.2 16 155.3 0.2 376 observed at about 23 years, in concomitance with the attainment of first sexual maturity, which in the northern Adriatic Sea population took place between 10 and 14 cm TL (G Scarcella, pers comm.) Furthermore, we frequently found false rings within the first 34 true annuli, as commonly observed in some species of Helicolenus (Massutớ et al 2000; Sequeira et al 2009) Similarly, the need of otolith sectioning for ageing purposes has been already reported for other scorpaenids, such as Scorpaena elongata (Gancitano and Ragonese 2008), Helicolenus dactylopterus (White et al 1998; Allain and Lorance 2000), Sebastes marinus and Sebastes mentella (Stransky et al 2005) The relationships between otolith diameters and fish size was linear, as reported for other species of Scorpaena (i.e S elongata and S maderensis) (La Mesa et al 2005; Rizzo et al 2003) and for some Sebastidae (Wyllie Echeverria 1987), indicating a proportionality between the two mentioned dimensions The seasonal deposition of opaque and translucent zones in the whole sample, as well as the microincrement counts of young-of-the-year specimens, supported the validation of ageing fish by counting annuli The fall-winter deposition of the translucent zone, supposed to occur during a slow growth period, took place when local sea temperatures reach minimum values (Artegiani et al 1997) The precision of age estimates was comparable to that reported for species of similar longevity, such as Pontinus kuhlii (Lúpez Abellỏn et al 2001), and within the range suggested by Campana (2001) Considering the Von Bertalanffy growth parameters estimated for the northern Adriatic population of S notata, the negative t0 and the low value of L1 compared to the maximum size of fish caught were probably due to the relatively low abundance of large and small fish in the sample Anyway, the longevity of about 15 years can be considered a reliable estimate of the maximum age attainable by the species, as the maximum size of fish aged is close to that reported in several localities of the Mediterranean Sea (Dulcic and Kraljevic 1996; Merella et al 1997; Morey et al 2003; Karakulak et al 2006) This estimate falls within the wide range of longevity found in other Mediterranean scorpaenids, such as S porcus and S maderensis, which attain respectively 11 and years of age (Jardas and Pallaoro 1992; La Mesa et al 2005), and S elongata and H dactylopterus, which Environ Biol Fish (2011) 91:369378 attain more than 30 years (Massutớ et al 2001; Gancitano and Ragonese 2008) The index of growth performance () is a useful tool for comparing the growth curves of different populations of the same species and/or of different species belonging to the same order (Sparre et al 1987) In Mediterranean scorpaenids, the growth performance ranged from 1.68 in S maderensis (La Mesa et al 2005), to 2.03 in S notata (present study), 2.07 in H dactylopterus (Massutớ et al 2000), 2.16 in S porcus (Jardas and Pallaoro 1992), 2.41 in P kuhlii (Lúpez Abellỏn et al 2001) and 2.45 in S elongata (Gancitano and Ragonese 2008), indicating a direct relationship between fish size and growth performance, as observed elsewhere in the genus Sebastes (Love et al 2002) Acknowledgements We thank the staffs of Marine Environment Management Unit and Population Dynamic Unit of ISMAR-CNR Ancona who respectively contributed to the sample collection and otoliths processing We thank also Lesley Farley for the English revision of the text and two anonymous referees for their suggestions and criticism that clearly improved an earlier version of this manuscript References Allain V, Lorance P (2000) Age estimation and growth of some deep-sea fish from the northeast Atlantic Ocean Cybium 24:716 Artegiani A, Bregant D, Paschini E, Pinardi N, Raicich F, Russo A (1997) The Adriatic Sea general circulation Part I: air-sea interactions and water mass structure J Phys Oceanogr 27:14921514 Beamish RJ, Fournier DA (1981) A method for comparing the precision of a set of age determinations Can J Fish Aquat Sci 38:982983 Beamish RJ, McFarlane GA (1995) A discussion of the importance of aging errors, and an application to walleye pollock: the worlds largest fishery In: Secor DH, Dean JM, Campana SE (eds) Recent developments in fish otolith research University of South Carolina Press, Columbia, pp 545565 Beckman DW, Wilson CA (1995) Seasonal timing of opaque zone formation in fish otoliths In: Secor DH, Dean JM, Campana SE (eds) Recent developments in fish otolith research University of South Carolina Press, Columbia, pp 545565 Berglund A, Rosenqvist G, Svensson I (1986) Mate choice, fecundity and sexual dimorphism in two pipefish species (Syngnathidae) Behav Ecol Sociobiol 19:301307 Bradai MN, Bouain A (1990) Age et croissance de Scorpaena notata (Rafinesque, 1810) dans le Golfe de Gabes (Tunisie) Bull Stn Oceanogr Salammbo 17:3546 Environ Biol Fish (2011) 91:369378 Campana SE (2001) Accuracy, precision and quality control in age determination, including a review of the use and abuse of age validation methods J Fish Biol 59:197242 doi:10.1006/jfbi.2001.1668 Campana SE, Thorrold SR (2001) Otoliths, increments, and elements: keys to a comprehensive understanding of fish populations? 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abundant at many of our sampling sites, and have been shown to significantly influence algal standing crops and benthic macroinvertebrate abundance in Costa Rican streams (Barbee 2002) The mullet J pichardi was not as common in the small streams we sampled, but is much larger in size and has traditionally been the most important species for subsistence fishing by indigenous communities in the study area (Borge and Castillo 1997; McLarney and Mafla 2007) There is a clear need for more detailed ecological information on these and other diadromous fishes in the region, and life history studies that build on previous work (e.g Cruz 1987) would be an important first step The life history requirements and ecological role of diadromous fishes in Central American streams deserve important consideration as hydropower development continues to increase in the region (Anderson et al 2006; McLarney and Mafla 2007) Acknowledgements Funding for this research was provided by the National Science 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Heithaus MR (eds) Biology of sharks and their relatives CRC Press LLC, Boca Raton, pp 399–447 Compagno LJV (1984) Sharks of the world An annotated and illustrated catalogue of shark species known to date Part I (Hexanchiformes to Lamniformes) FAO, Rome Cortes E (1999) Standardized diet compositions and trophic levels of sharks ICES J Mar Sci 56:707–717 Dayton PK (2003) The importance of the natural sciences... Importance of estuarine nursery habitats The importance of nursery habitats for the early life of marine fishes, particularly for estuarine dependent species have been recognized worldwide (Beck et al 2001; Heck et al 2003; Dahlgren et al 2006) As suggested by the ‘nursery role hypothesis’, a habitat is a nursery for juveniles of a particular species if its contribution per unit area to the production of individuals... the recruitment of JSB compared to the vast tidal and intertidal habitats of the Ariake Bay Otolith microchemistry (Sr/Ca ratio) of JSB revealed that approximately 50% of the population that recruit to the adult stock each year from the Ariake Bay is derived from juveniles that use nursery areas of the Chikugo River estuary (Secor et al 1998; Ohta 2004) Considering that the remaining 50% of the recruits... the diet of first feeding larvae Diets of first feeding and early larvae should be restricted to a few prey taxa because of smaller mouth gape width (MGW) of the first feeding and early larvae MGW is significantly and linearly related with the size of larvae and juveniles (Nip et al 2003); the relationship is: MGW (mm)=0.8383*SL (cm)−0.0408 (R2 =0.950) From this relationship, the estimated MGW of first... equally sized panels of 25 cm, 22 cm, and 18 cm stretch mesh Gillnets were set for an average of approximately 60 min at depths less than 5 m Catch rates and biological parameters Sevengill shark catch per unit effort (CPUE) was expressed as the number of sharks caught, standardized by the median number of hooks (35) used per set over the duration of sampling, divided by the number of hours the longline... the use of DNA analysis to improve dietary information J Exp Mar Biol Ecol 393:188–192 387 Barnett A, Stevens JD, Frusher SD, Semmens JM (2010b) Seasonal occurrence and population structure of the broadnose shark Notorynchus cepedianus in coastal habitats of south-east Tasmania J Fish Biol 77:1688–1701 Baum JK, Myers RA (2004) Shifting baselines and the decline of pelagic sharks in the Gulf of Mexico... studies in the Cyclades (Aegean Sea): size selectivity of small-hook longlines and monofilament gill nets Fish Res 58:25–40 Environ Biol Fish (2011) 91:369–378 Stransky C, Gudmundsdóttir S, Sigurdsson T, Lemvig S, Nedreaas K, Saborido-Rey F (2005) Age determination and growth of Atlantic redfish (Sebastes marinus and S mentella): bias and precision of age readers and otolith preparation methods ICES J