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1272 Lophius in the world: a synthesis on the common features and life strategies A. C. Fariña, M. Azevedo, J. Landa, R. Duarte, P. Sampedro, G. Costas, M. A. Torres, and L. Cañás Fariña, A. C., Azevedo, M., Landa, J., Duarte, R., Sampedro, P., Costas, G., Torres, M. A., and Cañás, L. 2008. Lophius in the world: a synthesis on the common features and life strategies. – ICES Journal of Marine Science, 65: 1272– 1280. Keywords: Atlantic Ocean, biological traits, life history, Lophius, NW Pacific, phylogeny. Received 11 January 2008; accepted 9 August 2008. A. C. Fariña, P. Sampedro, M. A. Torres, and L. Cañás: Instituto Español de Oceanografı́a, Paseo Marı́timo Alcalde Francisco Vázquez N8 10, 15001 A Coruña, Spain. M. Azevedo and R. Duarte: Instituto Nacional de Recursos Biológicos, INRB/L-IPIMAR, Avenida de Brası́lia, 1449-006 Lisboa, Portugal. J. Landa: Instituto Español de Oceanografı́a, Promontorio de San Martı́n s/n, 39004 Santander, Spain. G. Costas: Instituto Español de Oceanografı́a, Apartado 1552, 36200 Vigo, Spain. Correspondence to A. C. Fariña: tel: þ34 981 205362; fax: þ34 981 229077; e-mail: celso. [email protected]. Introduction The order Lophiiformes (Pisces: Teleostei) contains a highly diverse group of strictly marine fish distributed throughout the world’s oceans. The order includes 65 genera and 18 families, distributed among five suborders: Lophioidei, Antennarioidei, Chaunacoidei, Ogcocephaloidei, and Ceratioidei (Pietsch, 1984; Pietsch and Grobecker, 1987). The suborder Lophioidei (reviewed by Caruso, 1981, 1983), contains a single family, the Lophiidae, consisting of four genera (Sladenia, Lophiodes, Lophiomus, and Lophius) and 25 species (Caruso, 1985). The Lophiidae are found in temperate, tropical, and subtropical waters of the Atlantic, Indian, and western Pacific, but oddly enough not in the eastern Pacific. They are characterized by the dorso-ventrally compressed morphology of the head and body, a wide and cavernous mouth, thin skin, an absence of scales and swimbladder, and a modified first dorsal fin ray (illicium) with a terminal esca, which serves as a lure. Worldwide, seven species of Lophius (goosefish, monkfish, or anglerfish) are known, six being found along both coasts of the Atlantic Ocean, one extending to the western Indian Ocean, and the other in the Northwest Pacific (Figure 1). Lophius americanus is distributed in the Northwest Atlantic from the northern Gulf of St Lawrence and the Grand Banks, south to Cape Hatteras, NC (Steimle et al., 1999). Lophius gastrophysus lives in the western Atlantic from NC, USA, and the northern Gulf of Mexico to Argentina. The distribution of Lophius vomerinus extends from northern Namibia in the Southeast Atlantic to Durban on the east coast of South Africa, and in the northern and western Indian Ocean (FAO, 2008). Lophius vaillanti is found in the eastern Atlantic from north of Walvis Bay (238S) to the Gulf of Guinea (Maartens and Booth, 2005). Lophius piscatorius inhabits the Northeast Atlantic from Iceland and the southwestern Barents Sea to the Strait of Gibraltar, including the Mediterranean and the Black Sea (Caruso, 1986; Solmudsson et al., 2007). Lophius budegassa coexists with L. piscatorius over most of the range of the latter, although it has a more southerly distribution, from the British Isles to Senegal (Caruso, 1986). Finally, Lophius litulon occurs in the Northwest Pacific from Hokkaido to Kyushu, in the Gulf of Po-Hai, the Yellow Sea, and the East China Sea (Yoneda et al., 1997). The presence of the genus in the Pacific is restricted to those waters, because it is absent from the central and eastern Pacific Ocean. The most common habitat of the genus Lophius is bathydemersal over the continental shelf and upper slope down to depths .1000 m, on soft to hard sand and gravel substrata. The fish stay in the water column as eggs and larvae, then shift to a benthic existence as juveniles and adults. Anglerfish are caught in bottom trawl mixed or target fisheries, or as target species using gillnets. They are highly prized for human consumption and usually marketed fresh or frozen. # 2008 International Council for the Exploration of the Sea. Published by Oxford Journals. All rights reserved. For Permissions, please email: [email protected]. Downloaded from http://icesjms.oxfordjournals.org/ at Pennsylvania State University on February 27, 2014 Seven species of Lophius are known worldwide, six in the Atlantic Ocean and just one in the Northwest Pacific. The genus supports valuable fisheries (except for Lophius vaillanti), most for a long time, though the exploitation of Lophius gastrophysus along the coast of Brazil is relatively recent. The manuscript reviews the current knowledge of phylogeographic and biological traits of Lophius species, pointing out common aspects in the life histories. Within the Lophiidae, the genus Lophius is phylogenetically the most derived, vicariance and dispersal having played a significant role in driving speciation. Life histories seem to have followed similar adaptive processes from a common ancestor along with similar environmental characteristics. The genetic structure of populations is poorly known, and usually, genetic differentiation is limited. Life-history aspects (age, growth, reproductive cycle, early stages, and feeding ecology) are addressed, and fisheries are reviewed. However, knowledge of many aspects of the biology and ecology (e.g. validation of the growth pattern, maturation processes, spawning areas and periodicity, recruitment processes, mortality, stock identification, and habitat needs) remains limited. 1273 Synthesis of the common features and life strategies of Lophius spp. Figure 1. World map showing the distribution of Lophius species. La, L. americanus; Lb, L. budegassa; Lg, L. gastrophysus; Ll, L. litulon; Lp, L. piscatorius; Lv, L. vaillanti; and Lvo, L. vomerinus. Phylogeny and biogeography The origins of the related species of Lophius scattered throughout the world’s oceans are linked to geological and biogeographical events, such as tectonic activity, genetic divergence, colonization, and geographic isolation, regression and expansion. The emergence of natural barriers as a result of geological changes led to speciation by vicariance or dispersal (Grant and Leslie, 1993). Phylogenies based on morphological characters and allozymes have been used to propose derivations of the Lophiidae, identifying sister taxa and explaining the current geographic distribution of the species of Lophius (Grant and Leslie, 1993). Caruso (1985) concluded that the genus Lophius is the most derived of the four genera of the family Lophiidae, based on external osteology and other morphological characters. In formulating hypotheses for the phylogeny of the genus, Caruso (1977) proposed that small numbers of pectoral and dorsal fin rays and vertebrae represented the primitive state and large numbers the derived state. Among species of Lophius, L. vaillanti has the fewest pectoral fin rays (19–24), so according to Caruso’s hypothesis, it may be phylogenetically the most primitive species of the genus. Lophius americanus and L. piscatorius have relatively large numbers of vertebrae (30–31) and dorsal fin rays, so may represent the most derived types, although the similarity of the morphological characters could be the result of convergence processes between distantly related species (Caruso, 1983; Leslie and Grant, 1994). The following sequences of historical biogeographic and divergence events for the species of Lophius were suggested by Grant and Leslie (1993). The genus Lophius arose from a common ancestor of Lophiomus [a monotypic genus widely distributed throughout the western Pacific and Indian Ocean (Caruso, 1983)] by the closure of the Téthys Sea. The tropical eastern Atlantic L. vaillanti represents the most morphologically and genetically primitive of the Lophius species. Tectonic separation of the South American and African Plates split the ancestral Population structure and migrations The population structure of Lophius species remains poorly known. Most studies are restricted to North Atlantic species. Overall, both European anglerfish show limited genetic structure, low genetic variation having been detected off the west coast of Scotland (Crozier, 1988) and between populations from the Irish Sea and the west of Scotland (Crozier, 1987). In contrast, Blanco et al. (2006) reported high levels of microsatellite polymorphism for both L. piscatorius and L. budegassa from the Cantabrian Sea, north of Spain. For the northernmost populations of L. piscatorius, however, O’Sullivan et al. (2006) found no spatial or temporal genetic differentiation. Charrier et al. (2006) obtained similar results throughout the Northeast Atlantic and the Mediterranean Sea, but did find a significant, though weak, differentiation between Atlantic and Mediterranean populations of L. budegassa. They attributed this contrast between species to a combination of a phylogeographic barrier across the Almerı́a –Orán oceanographic front and the possibility of ancient colonization of the Mediterranean Sea by L. budegassa. Also for L. budegassa in the Mediterranean, Garoia et al. (2003) described a moderate level of genetic variation in polymorphic microsatellite loci. Populations of L. americanus are relatively homogeneous genetically along the US east coast, with the level of polymorphism within populations as low as that between populations (Chikarmane et al., 2000). Smith and Fujio (1982) also reported a very low level of genetic variation in L. litulon, but high levels have been observed in L. vomerinus off South Africa (Leslie and Grant, 1990). The lack of differentiation in the diverse species of Lophius suggests unrestricted gene flow over large areas. A common ecological strategy can be drawn, through potential broad dispersal capacity by an extended larval pelagic phase during which they are passively transported by currents before settling on the seabed (Leslie and Grant, 1990; Hislop et al., 2001). Moreover, juveniles and adults can move considerable distances, though the goal of such movements is not clear. For the Northeast Atlantic, Pereda and Landa (1997) and Laurenson et al. (2005) documented both spatial stability and extensive displacements of L. piscatorius. Laurenson et al. (2005) reported displacements of up to 876 km by an immature female, from a release location near the Shetland Islands to the southeast of Iceland, showing that large Downloaded from http://icesjms.oxfordjournals.org/ at Pennsylvania State University on February 27, 2014 Aspects of anglerfish life history and fisheries are well documented for some species, particularly for those from the North Atlantic and for L. litulon, but many aspects of the biology and ecology remain poorly known. For example, little is known about maturation, reproduction, spawning time or location, or the larval phase. The goal of this paper is to review the current knowledge and recent literature on the main biological characteristics and fisheries of the species of the genus, and to identify the common life strategies of the various species, pointing out the gaps and research needs from a perspective of fisheries management. tropical population and led to the appearance of L. gastrophysus in the West and the South Atlantic. A northward expansion of the range and vicariance or dispersal from ancestral South Atlantic populations gave rise to L. americanus. Grant and Leslie (1993) went on to suggest that an ancestral European species of Lophius arose from an ancestral North American species by a northeasterly expansion or dispersal of larvae or adults. Palaeo-oceanographic events in the Mediterranean Sea allowed the appearance of the two European species, Lophius piscatorius and L. budegassa. Dispersal of a European ancestor along the West African coast led to the appearance of L. vomerinus in the eastern South Atlantic. This species is considered a probable sister taxon of L. budegassa. Grant and Leslie (1993) also theorized that L. litulon arose by long-distance dispersal from the North Atlantic to the western North Pacific through a warmer Arctic Ocean after the Pliocene opening of the Bering Strait. Lophius piscatorius and L. litulon are sister taxa. The lack of colonization in the eastern North Pacific or posterior extinctions may explain the non-existence of Lophius in other areas of the Pacific. 1274 Feeding ecology Anglerfish are opportunistic, non-selective feeders, with a common feeding strategy. They are sit-and-wait predators, luring their prey by raising and moving the illicium. Food habits and diet composition are known for most species, although the information is limited for L. vaillanti and L. gastrophysus. The diet spectrum is size-dependent, with a similar pattern throughout the genus: invertebrates (crustaceans and cephalopods) make up a significant part of the diet of small juveniles, the consumption of invertebrates decreases with age, and larger juveniles and adults are mainly ichthyophagous, which includes a wide range of pelagic and benthic fish prey. Larger fish eat larger prey, but prey size selection can be attributed as much to the size and morphology of the mouth as to visual or sensory factors (Gordoa and Macpherson, 1990). In addition to overall ontogenetic shifts, the diet depends largely on predator size and geographic area, and like most ambush feeders, anglerfish diet varies seasonally to reflect spatio-temporal patterns in prey availability and abundance (Kosaka, 1966; Crozier, 1985; Laurenson and Priede, 2005). Adult L. americanus prey primarily on teleosts (red hake, Urophycis chuss, and sand lance, Ammodytes spp.; Armstrong et al., 1996), but there is also seasonal predation on squid (Staudinger, 2006). Over its geographical range, large L. piscatorius consume different prey items, with Norway pout (Trisopterus esmarkii) and blue whiting (Micromesistius poutassou) being the main prey in northern and southern European waters, respectively (Crozier, 1985; Pereda and Olaso, 1990; Laurenson and Priede, 2005), and other prey significant in certain areas at certain times. For example, whiting (Merlangius merlangus) and Norway lobster (Nephrops norvegicus) are significant components of anglerfish diet in the Irish Sea (Crozier, 1985), lesser sandeel (Ammodytes marinus) is seasonally important around Shetland (Laurenson and Priede, 2005), and cephalopods are important in the Cantabrian Sea (Velasco et al., 1998). Blue whiting and Phycis blennoides are major food items for middle and older age classes of L. budegassa in the Cantabrian Sea (Preciado et al., 2006). Dragonets (Paracallionymus costatus) and Cape hake (Merluccius capensis and Merluccius paradoxus) dominate the diet of L. vomerinus (Macpherson, 1985; Walmsley et al., 2005). Prey organisms of L. litulon are mainly fish, with Pseudosciaena manchurica being cited as dominant throughout the year and for all predator sizes (Cha et al., 1997). Empty stomachs are often found in Lophius, at a variable rate throughout the year, suggesting a low frequency of feeding (Crozier, 1985; Armstrong et al., 1996; Walmsley et al., 2005; Preciado et al., 2006). There does, however, tend to be an increase in the frequency of empty stomachs in autumn (up to 70% in L. budegassa) and with depth (Preciado et al., 2006). Seasonal variation in feeding intensity, with a decrease in autumn, has also been recorded for L. litulon (Kosaka, 1966), and Crozier (1985) reported major feeding activity of L. piscatorius in autumn and winter. It has been suggested that larger anglerfish only move to capture prey in response to internal rhythms independent of food availability (Macpherson, 1985) or when guaranteed a return, and that they do not eat again until the previous prey is completely digested (Kosaka, 1966). This could be a strategy to ensure maximum energy intake to offset the expense of prey capture (Walmsley et al., 2005). Very low incidences of cannibalism have been reported for Lophius (Crozier, 1985; Cha et al., 1997; Laurenson and Priede, 2005; Walmsley et al., 2005), except in L. americanus, for which predation by older conspecifics is relatively common (Armstrong et al., 1996). Reproduction and early life Typically, the morphology of anglerfish ovaries differs markedly from that of most other teleosts. Ovarian structure consists of very long ribbons of a gelatinous matrix, within which individual mature eggs float in separate chambers (Armstrong et al., 1992; Afonso-Dias and Hislop, 1996). In ripe females, the egg ribbons may be .10-m long and may contain more than a million eggs (Armstrong et al., 1992; Yoneda et al., 2001). The gonad mass of a mature female in spawning condition forms up to 35 –50% of total body mass (Armstrong et al., 1992; Yoneda et al., 2001; Walmsley et al., 2005) representing a considerable energetic contribution to reproduction. Females spawn buoyant gelatinous egg masses. Fertilization is external, but spawning behaviour and areas are poorly documented for the Atlantic. During the spawning season, L. piscatorius produces a single batch (AfonsoDias and Hislop, 1996), but L. litulon appears to spawn several batches (Yoneda et al., 2001). Female Lophius mature at a larger size than males, and spawning seasonality varies between species and geographic area (Table 1). The eggs and the larvae are pelagic. Hislop et al. (2001) indicated that the pelagic phase of L. piscatorius lasted for up to 4 months after hatching, but the early life stages are poorly known. The stages of embryonic and larval development of L. americanus were described by Everly (2002), and in varying degrees of detail for L. piscatorius (several reviews in the first part of the 20th century; Laurenson, 2006), L. gastrophysus (Matsuura and Yoneda, 1986), and L. litulon (Kim, 1976). The phylogenetic proximity between L. gastrophysus and L. americanus is supported by the same pattern and sequence of dorsal fin spine development at the larval stage (Everly, 2002). The common reproductive strategy of Lophius in releasing eggs in single veils may facilitate their dispersion and that of the larvae over great distances, while allowing for their protection against Downloaded from http://icesjms.oxfordjournals.org/ at Pennsylvania State University on February 27, 2014 displacements are not restricted to mature anglerfish. Other species (L. americanus, L. budegassa, and L. litulon) exhibit seasonal onshore–offshore movements in response to thermal conditions, prey availability, or in relation to spawning (Steimle et al., 1999; Landa et al., 2001a; Yoneda et al., 2002). Vertical displacements of immature and mature L. piscatorius from the seabed to near the surface have been recorded in the Northeast Atlantic (Hislop et al., 2000); the cause of this behaviour is unknown, but it may relate to spawning or feeding. There is no strong genetic evidence for stock discrimination in North Atlantic Lophius species. However, for management purposes, different areas have been defined based either on geographical barriers or on different operating fleets. Two stocks are defined (northern and southern) for L. americanus (Steimle et al., 1999) and L. budegassa (ICES, 2006), and three for L. piscatorius (ICES, 2006). For both L. piscatorius and L. budegassa, morphometric characters provide reasonable discrimination among populations from western and southern European waters (Duarte et al., 2004), but as described above, genetic and other biological traits make the current geographical boundary for separation of northern and southern stocks questionable (Fariña et al., 2004). Morphological evidence for stock structure among areas of the distribution range has also been documented for L. vomerinus by Leslie and Grant (1990). A. C. Fariña et al. 1275 Synthesis of the common features and life strategies of Lophius spp. Table 1. Length-at-first-maturity by sex and spawning season for species of the genus Lophius. Species Length-at-first-maturity (cm) Spawning season Source predators (Armstrong et al., 1992). Pelagic stages may intermix over wide areas, but in L. piscatorius there is evidence of transitory segregation (Swan et al., 2004), which is probably related to the overall limited genetic differentiation and the morphological components detected. Age and growth There have been many studies on age and growth of anglerfish, most undertaken in the past 20 years and with emphasis on European species. Again, data are particularly scarce for L. gastrophysus and L. vaillanti. Different hard parts (otoliths, illicia, and vertebrae) have been used for the purpose. Whole or sectioned sagittal otoliths were used in earlier studies (Tsimenidis and Ondrias, 1980; Griffiths and Hecht, 1986; Crozier, 1989) but their opacity, irregular margins, and incidence of multiple growth zones made interpretation difficult. An alternative option was available in the interpretation of sectioned illicia. This technique, developed by Dupouy et al. (1986) for L. piscatorius, has been widely used for age determination of European (Duarte et al., 1997; Quincoces et al., 1998a, b; Landa et al., 2001b; Garcı́a-Rodrı́guez et al., 2005) and other species, for example L. vomerinus (Maartens et al., 1999; Walmsley et al., 2005). Illicia have the advantage of being simple to collect from commercial samples, so allowing high sampling intensity. Vertebrae have been used in studies on the age and growth of L. americanus and L. litulon (Armstrong et al., 1992; Yoneda et al., 1997; Cha et al., 1998; Cullen et al., 2007). Regardless of preference for the structure used, all authors reported great difficulties in age interpretation, mainly because of the location of the first annulus and the existence of false annuli. Comparative studies between structures are scarce, although Wright et al. (2002) analysed the microstructure of lapilli otoliths of L. piscatorius to determine the position of the first annulus and to compare them with the microstructure of illicia and sagittae. Results indicated that the first increment on illicia might not be annual. Additionally, recent mark-recapture experiments applied to L. piscatorius provided evidence of faster growth than inferred from reading illicia (Laurenson et al., 2005; Landa et al., 2008). Woodroffe et al. (2003) showed that development of the marginal increments on otoliths and illicia has a similar pattern, and that the precision of age determination was better for sagittae sectioned through the sagittal plane than for whole otoliths or for illicia. However, it is necessary to take into account the fact that precision and agreement in age estimation among readers is related to the experience and familiarity of the reader with each structure. A comparative study between structures of the same fish, with readers experienced in the use of both illicia and otoliths, was performed at a workshop held in 2004 for L. piscatorius and L. budegassa. Results were that agreement between experienced readers was better for illicia for both species and that ages from illicia tended to be higher than for otoliths (Duarte et al., 2005). Among the different studies whose growth parameters are summarized in Table 2, some general patterns can be extracted. First, the northern species (L. americanus, L. piscatorius, and L. litulon) show faster growth rates than austral (L. vomerinus) and European species with a more southern distribution (L. budegassa). Second, growth rates differ significantly between sexes, particularly in older fish (Figure 2). Females attain greater length and, according to age readings, greater age than males. Of the Lophius species, the L1 for females ranged from 110 to 166 cm and for males from 68 to 129 cm, with age estimates up to 25 and 21 years, respectively (Table 2). Finally, length-at-age increases approximately linearly until ages 11 –15 in L. piscatorius, L. budegassa, and L. americanus (Cullen et al., 2007; Duarte et al., 2007; Landa et al., 2007). Fisheries The genus Lophius is exploited worldwide. Historically, they have been bycatch in mixed fisheries, but an increase in their economic value together with the overexploitation of other groundfish species has led to the development of targeted anglerfish fisheries. Anglerfish are caught with trawl and fixed nets, mostly gillnets. According to FAO (2007), the world catch of the three main species (L. piscatorius, L. americanus, and L. vomerinus) in 2007 levelled off at .100 000 t. In general, anglerfish are managed through one or a combination of a total allowable catch (TAC), effort control, mesh size restrictions, and seasonal closures. Lophius piscatorius and L. budegassa have been exploited in northern European waters for at least a century as a valuable bycatch (Hislop et al., 2001), and since the 1980s they have also become an important fishery resource in Iberian waters (Piñeiro Downloaded from http://icesjms.oxfordjournals.org/ at Pennsylvania State University on February 27, 2014 Females Males Lophius piscatorius 73.2 – 98.0 48.9 –58.0 November –May Afonso-Dias and Hislop (1996) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . January –June Duarte et al. (2001) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . May–June Quincoces et al. (1998b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Laurenson et al. (2001) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lophius budegassa 54.8 – 64.5 34.5 –37.6 October –March Duarte et al. (2001) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . May–June Quincoces et al. (1998a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lophius americanus 44.0 – 48.5 36.9 –40.0 May–June Armstrong et al. (1992) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Almeida et al. (1995) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lophius vomerinus 58.2 39.9 Austral spring Maartens and Booth (2005) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36.9 37.6 Walmsley et al. (2005) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lophius litulon 56.7 36.2 February –May Yoneda et al. (2001) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lophius gastrophysus 50.0 – Austral spring and summer Valentim et al. (2007) 1276 A. C. Fariña et al. Table 2. Growth parameters (L1, k, and t0) for male and female Lophius spp. from studies based on readings of illicia and vertebrae. et al., 2001; ICES, 2006). Both species are caught on the same grounds by the same fleets and are marketed together, but L. piscatorius is the most abundant. Landings increased sharply with the development of directed fisheries and the expansion to deeper water with improvements in fishing technology. ICES provides advice for the management of three stocks in European waters: North Sea (Divisions IIIa, IVa –c, and VIa,b), Northern (Divisions VIIb –k and VIIIa,b,d), and Iberian (Divisions VIIIc and IXa) stocks. Total landings from these stocks ranged, in the past two decades, from 50 000 to 70 000 t (ICES, 2008a, b). Figure 2. Mean length-at-age for a selection of growth studies on Lophius spp. Lophius americanus is commercially exploited by Canada and the USA. In Canada, the most important fishery is on the Scotian Shelf, and the species is an important bycatch species in the fisheries on the Grand Banks and in the Gulf of St Lawrence (Kulka and Miri, 2003). The US commercial fishery operates in deep water of the Gulf of Maine, Georges Bank, and off southern New England. The development of this fishery in the 1980s was related to the opening of the international markets of Europe and Asia, and led to a peak in landings in 1998, of some 29 000 t (NOAA, 2004). After the implementation of a management plan in the USA in 2000, landings have remained at 20 000 t (NOAA, 2004). Although both L. vomerinus and L. vaillanti are found in waters of the Southeast Atlantic, L. vomerinus is the most abundant (94% of Namibian anglerfish landings) and commercially valuable (Maartens and Booth, 2001a, b). FAO total official catches for L. vomerinus in 1999 were 21 000 t, with Namibia and South Africa being the countries recording highest catches. In Namibia until the early 1990s, anglerfish were taken as bycatch in the trawl fishery targeting Cape hake, but since independence in 1991, the anglerfish-directed fishery has grown substantially, from ,2000 to 12 000 t in 1994 (Booth and Quinn, 2006). In 2001, a management plan was introduced, and as well as other management measures, a TAC of 13 000 t was established. In South Africa, where anglerfish is still just a bycatch of the trawl fishery targeting Cape hake (Walmsley et al., 2005), catches increased from some 5000 t in the 1980s to 10 700 t in 2001 (Booth, 2004). A catch limit of 7000 t was introduced in 2005 (R. W. Leslie, pers. comm.). Lophius gastrophysus, known locally as “frog-fish” is one of the main resources of deep-sea fisheries off the south and southeast coasts of Brazil. Historically, it has been a bycatch of the industrial shrimp fishery (Jablonski et al., 1998; Vianna and Almeida, 2005), but in 2001 it became the target of the trawl fishery operating south of Brazil and of a foreign fleet operating with “rasco”, a gillnet specially designed to catch the species (Bruno et al., 2001; Perez et al., 2002a, b). The efficiency of the “rasco” underpinned Downloaded from http://icesjms.oxfordjournals.org/ at Pennsylvania State University on February 27, 2014 Species Area Method Sex L1 (cm) k t0 Source Lophius piscatorius Celtic Sea and Bay of Biscay Illicia Male 129.50 0.11 0.54 Dupouy et al. (1986) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Female 166.60 0.08 0.40 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Iberian Atlantic waters Illicia Male 110.50 0.11 0.25 Landa et al. (2001b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Female 163.50 0.06 20.44 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lophius americanus Southern New England and Virginia Vertebrae Male 105.90 0.16 0.20 Armstrong et al. (1992) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Female 157.60 0.10 0.16 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lophius litulon East China Sea and Yellow Sea Vertebrae Male 113.00 0.08 0.40 Yoneda et al. (1997) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Female 154.70 0.06 0.35 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Southwest Korean waters Vertebrae Male 82.23 0.18 20.64 Cha et al. (1998) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Female 127.60 0.12 20.39 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lophius budegassa Celtic Sea and Bay of Biscay Illicia Male 84.76 0.10 0.56 Dupouy et al. (1986) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Female 111.20 0.07 0.50 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Iberian Atlantic waters Illicia Male 71.50 0.13 0.05 Landa et al. (2001b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Female 93.50 0.10 0.50 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lophius vomerinus Namibian waters Illicia Male 72.30 0.14 20.30 Maartens et al. (1999) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Female 112.00 0.08 20.36 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . West South African coast Illicia Male 68.50 0.10 21.69 Walmsley et al. (2005) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Female 110.20 0.05 21.54 Synthesis of the common features and life strategies of Lophius spp. 1277 the development of a national gillnet fleet targeting anglerfish (Perez et al., 2002a). In 2001, the anglerfish landings from southeastern Brazil were some 9000 t, and 37% of this was from the gillnet fleet (Perez et al., 2005). Because of the heavy exploitation and economic importance of L. gastrophysus, a federal regulation with several measures aimed at sustainable exploitation was established in 2005 (Valentim et al., 2007). Fishery information on L. litulon is scarce. The species is caught by bottom trawlers from Japan, China, and Korea during winter, but other species commercially exploited in the same area are considered to be more important than anglerfish (Yoneda et al., 2001). † information on the maturation processes, on the role of the gelatinous veil, spawning behaviour, spawning areas, and fecundity; Final remarks † basic biological studies and data on populations of L. gastrophysus and L. vaillanti; † basic data (length composition, abundance index, size distribution, etc.) on several species of Lophius; † for European anglerfish (L. piscatorius and L. budegassa), more accuracy in age determination; † for other species of Lophius, age and growth data collected routinely for age-based analysis; † environmental effects of fishing on population dynamics; † determination of the spawning components and stock structure, to identify appropriate management units. Acknowledgements We thank the European Commission, Directorate General Fisheries, for financial support through Study Contract 99/013 and FISH/2004/03-22, U. Autón for technical assistance, and the Instituto Español de Oceanografı́a for financial support to MAT through a research grant. References Afonso-Dias, I., and Hislop, J. R. G. 1996. The reproduction of anglerfish Lophius piscatorius Linnaeus from the north-west coast of Scotland. Journal of Fish Biology, 49: 18– 39. Almeida, F. P., Hartley, D. L., and Burnett, J. 1995. Length – weight relationships and sexual maturity of goosefish off the northeast coast of the United States. North American Journal of Fisheries Management, 15: 14 – 25. Armstrong, M. P., Musick, J. A., and Colvocoresses, J. A. 1992. Age, growth, and reproduction of the goosefish Lophius americanus (Pisces: Lophiiformes). Fishery Bulletin US, 90: 217 – 230. Armstrong, M. P., Musick, J. A., and Colvocoresses, J. A. 1996. Food and ontogenetic shifts in feeding of the goosefish, Lophius americanus. Journal of Northwest Atlantic Fishery Science, 18: 99 – 103. Blanco, G., Borrell, Y. J., Cagigas, E., Vázquez, E., and Sánchez Prado, J. A. 2006. A new set of highly polymorphic microsatellites for the white and black anglerfish (Lophiidae). Molecular Ecology Notes, 6: 767– 769. Booth, A. J., and Quinn, T. J. 2006. Maximum likelihood and Bayesian approaches to stock assessment when data are questionable. Fisheries Research, 80: 169– 181. Booth, T. 2004. South African monkfish (Lophius vomerinus) stock assessment. Marine and Coastal Management Demersal Working Group Document WG/05/04/D: A7. 18 pp. Bruno, I., Fariña, A. C., Landa, J., and Morlán, R. 2001. The gillnet fishery for anglerfish (Lophius piscatorius) in deep-water in the Northwest of Iberian Peninsula. NAFO SCR Document 01/ 99. 5 pp. Caruso, J. H. 1977. The systematics of the fish family Lophiidae. PhD thesis, Tulane University, New Orleans, USA. 219 pp. Caruso, J. H. 1981. The systematics and distribution of the lophiid anglerfishes. 1. A revision of the genus Lophioides with the description of two new species. Copeia, 1981: 522 –549. Caruso, J. H. 1983. The systematics and distribution of the lophiid anglerfishes. 2. Revisions of the genera Lophiomus and Lophius. Copeia, 1983: 11 – 30. Caruso, J. H. 1985. The systematics and distribution of the lophiid anglerfishes. 3. Intergeneric relationships. Copeia, 1985: 870 – 875. Caruso, J. H. 1986. Lophiidae. In Fishes of the North-eastern Atlantic and the Mediterranean, pp. 1362– 1363. Ed. by P. J. P. Whitehead, Downloaded from http://icesjms.oxfordjournals.org/ at Pennsylvania State University on February 27, 2014 This very general overview shows that species of the genus Lophius have similar traits throughout the world. However, some biological and life-history aspects, as well as fisheries and exploitation patterns, have different levels of knowledge, depending on species and geographic area. Growth studies based on illicia (L. piscatorius and L. budegassa) and vertebrae (L. americanus) show a linear relation between age and total length, which is not consistent with the von Bertalanffy growth model (von Bertalanffy, 1938). Information on growth rate is crucial because estimation of stock/population productivity and resilience is partly dependent on it. For all Lophius species, there is a great need for information related to validation of the methods of age determination. Among others, Wright et al. (2002) and Landa et al. (2008) provide information on this topic for L. piscatorius, and it suggests that the current reading criteria applied to illicia may be biased, which might underpin the apparently linear growth pattern. Therefore, undertaking similar studies for other species of Lophius and in other areas is important, because clarification may be obtained for all species of Lophius from such research. Studies on the reproductive biology of Lophius show general sampling problems related to sampling females close to the spawning season. Such seasons are generally broadly defined, and related parameters such as age- and length-at-first-maturity are difficult to estimate, particularly for females. More investigation of this topic could provide more insight into the location of spawning grounds, and knowledge of spawning behaviour would be of benefit to management of all Lophius species. The exploitation of Lophius species showed a similar development worldwide, with commercial interest being relatively recent compared with the more traditional groundfish species. 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