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Feeding Ecology of Demersal Elasmobranchs from the Shelf and Slope Off the Balearic Sea (Western Mediterranean)

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Feeding Ecology of Demersal Elasmobranchs from the Shelf and Slope Off the Balearic Sea (Western Mediterranean) Scientia Marina 75(4) December 2011, 633-639, Barcelona (Spain) ISSN: 0214-8358 doi: 10.3989/scimar.2011.75n4633 Feeding ecology of demersal elasmobranchs from the shelf and slope off the Balearic Sea (western Mediterranean) MARIA VALLS, ANTONI QUETGLAS, FRANCESC ORDINES and JOAN MORANTA Instituto Español de Oceanografía, Centre Oceanogràfic de les Balears, Moll de Ponent s/n, 07015 Palma, Spain. E-mail: [email protected] SUMMARY: The feeding ecology of eight demersal elasmobranchs, three sharks (Etmopterus spinax, Scyliorhinus canicula and Galeus melastomus) and five batoids (Myliobatis aquila, Leucoraja naevus, Raja polystigma, R. miraletus and R. clavata), from the Balearic Sea (western Mediterranean) was analyzed. For each species, the diet and feeding habits were characterized by depth strata using quantitative indices such as diet overlap, diet breadth and prey diversity. Diet variation with size and depth were also tested for the most abundant species. For shelf-living species, natantian and reptantian crustaceans together with teleosts were the most important preys. On slope bottoms, euphausiids were the preferential prey for S. canicula and G. melastomus, while E. spinax fed mainly on cephalopods. The most specialist and generalist diet corresponded to G. melastomus living on the upper slope and S. canicula from the continental shelf, respectively. High overlap was found between all the skates on the continental shelf and the sympatric sharks S. canicula and G. melastomus on the slope. Significant overlap was also found between S. canicula and R. clavata on the continental shelf. Size was found to significantly affect the diet of S. canicula, G. melastomus and R. clavata, whereas depth affected exclusively S. canicula. Keywords: Elasmobranch, batoid, shark, feeding habits, diet overlap, niche breadth, Mediterranean. RESUMEN: Ecología alimentaria de elasmobranquios demersales de plataforma y talud en aguas de las Islas Baleares (Mediterráneo Occidental). – Se estudia la ecología alimentaria de ocho elasmobranquios demersales, tres tiburones (Etmopterus spinax, Scyliorhinus canicula y Galeus melastomus) y cinco rayas (Myliobatis aquila, Leucoraja naevus, Raja polystigma, R. miraletus y R. clavata), del mar Balear (Mediterráneo Occidental). Para cada especie se carac- terizó la dieta y la estrategia alimentaria por estratos de profundidad mediante diferentes índices cuantitativos. También se analizó la variación intraespecífica de la dieta con la talla y la profundidad en las especies más abundantes. Las presas más frecuentes en especies de la plataforma continental fueron los peces y los crustáceos natantia y reptantia. En aguas del talud las presas principales de S. canicula y G. melastomus fueron los eufausiáceos, mientras que E. spinax consumió preferente- mente cefalópodos. Las especies con la dieta más especializada y más generalista fueron G. melastomus del talud superior y S. canicula de la plataforma costera, respectivamente. En aguas de la plataforma se detectó solapamiento de dieta entre todas las especies de batoideos, así como entre S. canicula y R. clavata; el solapamiento fue significativo también entre S. cani- cula y G. melastomus del talud. Se observaron cambios ontogénicos en la dieta de S. canicula, G. melastomus y R. clavata, mientras que la única especie que mostró cambios de dieta con la profundidad fue S. canicula. Palabras clave: elasmobranquio, batoideo, tiburón, estrategia alimentaria, solapamiento de dieta, amplitud de nicho, Medi- terráneo. INTRODUCTION the marine ecosystems with a top-down control on the size and dynamics of many species (Wetherbee and Trophic relationships are fundamental to under- Cortes, 2004). Globally, there is increasing evidence standing biological interactions in animal communities that elasmobranchs are more affected than teleosts by and how they respond to human exploitation. Elasmo- fishing exploitation (Stevens et al., 2000). Insular ar- branchs are top predators playing an important role in eas from the western and central Mediterranean sustain 634 • M. VALLS et al. elasmobranch assemblages with higher diversity and abundance than adjacent mainland areas (Massutí and Moranta, 2003). Despite the importance of feeding relationships to understand the food structure and dynamics of marine ecosystems, little is known about the feeding ecology of most elasmobranchs (Heithaus, 2004; Wetherbee and Cortes, 2004). This is especially true for batoids, which have received considerably less attention than sharks at a worldwide level (Ishihara, 1990; Motta, 2004). Research on trophic relationships among elasmobranch sympatric species is also scarce in the western Mediterranean, where selachian feeding habits have been studied exclusively with regard to single species and groups of two or three species with similar ecological roles or habitat distribution (Macpherson, 1980, Carrason et al., 1992; Saidi et al., 2009). Further- more, few studies have focussed on the feeding ecol- ogy of batoids individual species in the Mediterranean Sea (Jardas et al., 2004; Romanelli et al., 2007; Saglam and Bascinar, 2008). In the present paper we analyze the diet, feeding habits and trophic interactions for three selachians (Etmopterus spinax, Scyliorhinus canicula and Galeus melastomus) and five batoids (Myliobatis aquila, Leu- Fig. 1. – Map of sampling sites of the eight elasmobranch species coraja naevus, Raja polystigma, R. miraletus and R. taken in the Balearic Islands (western Mediterranean). The 100, clavata) from waters off the Balearic Islands (western 200, 500 and 800 m isobaths are shown. Mediterranean). 1 1 MATERIALS AND METHODS Bi = −1 , n −1 p2 ∑ j ij The elasmobranch species were taken from the fishery-independent MEDITS bottom trawl surveys where pij is the proportion of diet of predator i that is conducted around the Balearic Islands (Fig. 1) in early made up of prey j and n is the number of prey catego- summer from 2007 to 2009. Both sampling scheme and ries. This index ranges from 0 to 1, low values indi- gear (GOC-73) were the same as those used through- cating diets dominated by a few prey items (specialist out the Mediterranean in the MEDITS programme predators) and higher values indicating generalist diets (Bertrand, 2002). A total of 141 hauls were performed (Krebs, 1999); and 6) species diversity in both prey during daylight hours between 45 and 755 m depth. number (H’n) and prey volume (H’v) calculated using Whenever possible, ten individuals of each elasmo- the Shannon-Wiener index. branch species from each haul were separated and their To standardize data and to facilitate diet compari- total length, weight and sex were noted. Fresh stomach sons and analyses, the following twelve major prey contents were identified on board to the lowest possible categories were established: Polychaeta, Sipuncula, taxonomic level and then all prey items were counted Gastropoda, Bivalvia, Cephalopoda, Euphausiacea, and their volume was measured with a calibrated de- Amphipoda, Isopoda, Mysidacea, Crustacea Reptantia, vice used in previous studies (Olaso et al., 1998). Crustacea Natantia and Teleostea. Prey categories with The diet of each elasmobranch species was quanti- frequency of occurrence lower than 3% and unidentifi- fied using the following indices: 1) frequency of oc- able remains were excluded from these analyses. Diet currence (%F), percentage of stomachs with a specific overlap between species was calculated using the Sch- type of prey in relation to the total number of stomachs oener index (Hurlbert, 1978), which ranges from 0 (no containing food; 2) numerical (%N) and volumetric overlap) to 1 (complete overlap), with values greater (%V) composition, expressed as the percentage con- than 0.6 representing a significant overlap (Sala and tribution of each prey to the whole content, in number Ballesteros, 1997; Wallace, 1981). In order to compare or volume respectively; 3) index of relative importance exclusively coexisting species, all indexes characteriz- (IRI=%F(%N+%V), which was standardized follow- ing the diet were analyzed separately for the following ing %IRI=(IRI/∑IRI)×100 (Cortes, 1997); 4) vacuity four bathymetric strata described in the study area (Or- index (v), the percentage of empty stomachs; and 5) dines et al., 2011): 1) continental shelf (CS: 45-180 m); diet breadth, which was calculated using the Levin’s 2) shelf-break (SB: 180-330 m); 3) upper slope (US: standardized index: 330-495 m); and 4) middle slope (MS: 495-750 m). SCI. MAR., 75(4), December 2011, 633-639. ISSN 0214-8358 doi: 10.3989/scimar.2011.75n4633 FEEDING ECOLOGY OF MEDITERRANEAN ELASMOBRANCHS • 635 Table 1. – General information and diet composition (standardized index of relative importance, %IRI) of eight elasmobranch species caught at different bathymetric strata (CS, continental shelf; SB, shelf-break; US, upper slope; MS, middle slope) in the Balearic Sea (western Mediterranean). N, sample size; TL, total length; WR, weight range; DR, depth range; v, vacuity index; H’n, H’v: Shannon-Wiener diversity in number and volume respectively; and Bi, Levin’s niche breadth. Values in brackets are the number of taxa in the twelve major taxonomic groups of prey (see Supplementary material Appendix 1). R.pol., R. polystigma; M.aqu., M. aquila; L.nae., L. naevus; R.mir., R. miraletus. R.pol. M.aqu. L.nae. R.mir. R. clavata S. canicula G. melastomus E. spinax CS CS CS CS CS SB US CS SB US US MS MS N 15 23 27 31 266 48 6 766 66 60 170 167 46 TL (cm) 28-45 51-116 22-52 24-43 18-88 18-77
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