Shelf-Break Areas with Leptometra Phalangium (Echinodermata: Crinoidea)
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Marine Biology (2004) 145: 1129–1142 DOI 10.1007/s00227-004-1405-8 RESEARCH ARTICLE F. Colloca Æ P. Carpentieri Æ E. Balestri G. D. Ardizzone A critical habitat for Mediterranean fish resources: shelf-break areas with Leptometra phalangium (Echinodermata: Crinoidea) Received: 8 October 2003 / Accepted: 10 May 2004 / Published online: 3 July 2004 Ó Springer-Verlag 2004 Abstract This paper considers the potential role of the structure of species occurring on the shelf break showed crinoid Leptometra phalangium as an indicator of highly that for some of them the selection of this area is related productive areas along the shelf break that can sustain to specific phases of their life cycle. Significant highest large biomasses of benthopelagic fish and recruits. The abundance of recruits and juveniles was observed for structure of fish assemblages in the central Mediterra- Merluccius merluccius, Helicolenus dactylopterus, Phycis nean Sea (central-western coast of Italy), analysed on blennoides, Parapenaeus longirostris and Capros aper in the basis of surveys carried out in summer and autumn at least one of the two seasons. Similarly, an increased from 1997 to 2001, revealed the presence of a well-de- abundance of spawners of red mullet Mullus barbatus fined group of species on the shelf break. This area, and four-spotted megrim Lepidorhombus boscii was ob- occurring at a depth of between 120 and 170 m, is served on the shelf break. Results of this study may have characterised by detritic organogenic sediments colon- important consequences for management of fish stocks ised by the crinoid L. phalangium, a suspension-feeding and assemblages in the central Mediterranean. The co- macro-epibenthic species confined in the Mediterranean occurrence of high densities of L. phalangium and ben- to the shelf-break area. Its abundance in the studied area thopelagic fish, occurring mainly with juveniles and can reach 12–15 ind. m)2. A total of 121 species spawners, strongly indicates a potential role of L. pha- belonging to 66 families of demersal organisms (crusta- langium as an indicator of highly productive areas cean decapods and stomatopods, cephalopods, selaceens around the shelf break. Such areas appear to play a and teleosteens) were caught at shelf-break stations from major role in the production of some of the most September to October. The species which typified the abundant and commercially important fish species, such assemblage were the fishes Trisopterus minutus capel- as the Mediterranean hake and red mullet. anus, Merluccius merluccius, Glossanodon leioglossus, Argentina sphyraena, Capros aper, Macroramphos scol- opax and Lepidotrigla cavillone, the crustacean decapod Parapenaeus longirostris and the cephalopods Illex coindetii and Todaropsis eblanae. Detritic shelf-break Introduction stations showed a higher abundance of demersal organisms than stations distributed on muddy bottoms The shelf break is a high-energy zone as a result of the in the same depth range (100–200 m). Such differences interplay of tides, storm waves, breaking internal waves appeared to be significant in September–October, when and bottom turbulence produced by fronts separating a clear increase in benthopelagic zooplanktivorus spe- shelf and slope waters (Herman 1972; Vanney and cies, such as Glossanodon leioglossus, Trachurus trachu- Stanley 1983). Intermittent mixing events may generate rus, Trachurus picturatus, was found. The length increased ammonium and nitrate concentrations in wa- ter columns, which, in turn, increase euphotic zone production (Soetaert et al. 2001). An enhanced phyto- Communicated by R. Cattaneo-Vietti, Genova plankton and zooplankton biomass associated with F. Colloca (&) Æ P. Carpentieri Æ E. Balestri Æ G. D. Ardizzone shelf-break fronts has been observed (see Mann and Department of Animal and Human Biology, Lazier 1996). The high production of plankton at the University of Rome ‘‘La Sapienza’’, V. le dell’Universita` shelf break makes it an important feeding ground for 32, 00185 Rome, Italy E-mail: [email protected] large shoals of fish (Williams et al. 2001), flocks of Tel.: +39-6-49914763 oceanic birds (Skov and Durinck 1998) and cetaceans Fax: +39-6-4958259 (Hain et al. 1985). 1130 Studies of communities in the Mediterranean shelf in the sensitive phases (recruitment, spawning, post- break have mostly focused on benthic assemblages in spawning) of the life cycle of demersal fish species. relation to sediment characteristics, which are muddy with a component of relict detritic sediments produced during Pleistocenic (20,000–8,000 years ago) fluctua- Materials and methods tions in sea levels (Pe´re` s and Picard 1964; Reyss 1973). Both eastern Atlantic and Mediterranean benthic com- Study area munities on the shelf break are characterised by high concentrations of macro-epibenthic filter-feeding and The area investigated covers 15,289 km2 off the central suspension-feeding organisms (Flach et al. 1998; Kal- western coasts of Italy (central Mediterranean Sea) be- lianotis et al. 2000). In the Mediterranean, where a well- tween 10 and 700 m depth (Fig. 1). The continental shelf defined benthic community on the shelf break has been extends up to 120–150 m depth between 15 and 30 km described (DL, Pe´re` s and Picard 1964), the dominant from the shoreline. The shelf break is characterised by megabenthic species is generally the crinoid Leptometra detritic organogenic sediments, with locally high abun- phalangium. This species is confined to the restricted dances of the crinoid Leptometra phalangium.This depth range that corresponds to the shelf-break and organism has a patchy distribution with a maximum canyon-head areas under bottom currents, where it can abundance of 12–15 ind. m)2 (Colloca 2003). ) reach a density of 30–50 ind. m 2 (Laborel et al.1961; Environmental conditions along the continental shelf Vaissie` re and Carpine 1964; Reyss and Soyer 1965; are heterogeneous due to the presence of both eutrophic Bourcier and Zibrowius 1973; Reyss 1973, 1974; Kal- areas, like those affected by the Tiber River’s waters lianotis et al. 2000). In the Atlantic Ocean, the vicariant (central Latium), and oligotrophic areas around the species Leptometra celtica characterises the shelf break Pontine Islands, lagoons connected to the sea, phaner- along the northwest Spanish and southwest Irish coasts ogam meadows, rocky and sandy shores. (Lavaleye et al. 2002). The limit between the upper and middle slopes ranges The impact of towed gear is particularly heavy on between 400 and 500 m depth (Carpine 1970; Relini these beds, both on L. phalangium, a fragile organism et al. 1986). The temperature down to 120–200 m depth that is easily destroyed by trawling, and on other epi- is generally constant at about 13°C (Ifremer 1997). A benthic organisms that have shown reduced abundance layer of Levantine intermediate water, characterised by in trawled areas (Smith et al. 2000). maximal salinity, flows anticlockwise along the slope Several studies have shown a relationship between the between 250 and 700 m depth (Serravall and Cristofalo distribution of macro-epibenthic and of demersal fish 1999). assemblages in the Mediterranean (Gaertner et al. 1999; Colloca et al. 2003a), indicating that the negative effects of fishing on the benthic community may be mirrored in Trawl survey data the distribution and abundance of fish species. The role played by the shelf break in the distribution Data were collected during eight trawl surveys, as part and dynamics of demersal species is very poorly under- of the MEDITS and GRUND projects, conducted, stood in the Mediterranean, even though previous respectively, in July and September–October from 1997 studies have revealed the occurrence of a defined fish to 2001. Both projects used a randomised, stratified assemblage on deeper shelf/shallow slope bottoms (Ga- sampling design based on depth (five bathymetric strata: ertner et al. 1999; Biagi et al. 2002; Colloca et al. 2003a). 10–50, 51–100, 101–200, 201–500 and 501–700 m) and It is worth stressing that this assemblage is characterised area. The depth limits were adopted to best cover the by high concentrations of juveniles of important com- distribution areas of the mainly exploited, or potentially mercially exploited species such as European hake exploitable, species. Locations of stations were selected (Merluccius merluccius). randomly within each stratum at the beginning of the The main aim of this paper is to evaluate the poten- projects (1996). The same stations, 61 for GRUND and tial role of L. phalangium beds as indicators of highly 46 for MEDITS, were sampled in the following years productive areas along the shelf break, which can sustain (see Relini 1998; Bertrand et al. 2002). Ten stations in all large biomasses of benthopelagic fish and recruits. For were located on detritic bottoms, with L. phalangium this purpose, the abundance and population structure of along the shelf break (crinoid stations). demersal species, including European hake, red mullet There were some differences between the two projects (Mullus barbatus), John dory (Zeus faber), poor cod as regards the vessels and equipment used for the sur- (Trisopterus minutus capelanus), horse mackerel (Tra- veys. The GRUND project was conducted using a vessel churus trachurus) and deep-water pink shrimp (Parape- (60 tons and 600 HP) equipped with an Italian otter naeus longirostris) were analysed by comparing data trawl mounting a 40-m head rope, a 40-mm ground obtained during stratified trawl surveys both on and off chain and 40- and 30-mm stretched mesh in the wing and the shelf break. cod-end, respectively. MEDITS was carried out from a The conservation of shelf-break fish assemblages is 88-ton, 800-HP vessel using an otter trawl net with a 40- also discussed with a view to reducing the fish mortality m head rope, an 8-m wing spread and a 20-mm mesh size 1131 Fig. 1 Trawl stations on the continental shelf and upper slope of the central western coasts of Italy (central Mediterranean Sea) (open circles Leptometra phalangium stations on the shelf break) at the cod-end (Fiorentini et al.