Arbacia lixula and Paracentrotus lividus are the dominant sea urchins in the Mediterranean sublittoral, where they are key structuring species due to their grazing activity. It has been commonly accepted that competition between both species is minimized by specializing in different algal foods. A. lixula is considered to feed mainly on encrusting coralline algae, while P. lividus prefers fleshy macroalgae. We used stable isotope analysis to test if these species occupy different trophic positions at 3 locations in the western Mediterranean and one in Macaronesia. Our results show unambiguously that A. lixula always occupies a higher trophic level than P. lividus, with a δ 15 N comparable in some locations to strict carnivores such as Actinia schmidti or Marthasterias glacialis. A temporal monitoring at one locality showed that this signature of a higher trophic level is consistent throughout the year. These results are incompatible with the current belief of an herbivorous diet for A. lixula and suggest that it must be considered an omnivore tending to carnivory in Mediterranean ecosystems, feeding at least partially on sessile animals such as Cirripedia, Hydrozoa or Bryozoa. A parallel analysis of gut contents showed a predominance of vegetal items in both species, although A. lixula consistently had a higher abundance of animal components than P. lividus. Our results challenge the validity of using gut content observations alone for characterizing the trophic behaviour of omnivorous marine invertebrates that feed on a variety of food sources with different digestibility.
KEY WORDS: Arbacia lixula · Paracentrotus lividus · Trophic relationships · Benthic community · Stable isotope analysis.
Resale or republication not permitted without written consent of the publisherMar Ecol Prog Ser 441: [117][118][119][120][121][122][123][124][125][126][127][128] 2011 This herbivorous behaviour described in Arbacia lixula is, however, in sharp contrast with other species in the genus Arbacia, where omnivorous or unambiguously carnivorous diets have been reported. North American A. punctulata feeds on sessile invertebrate species, sand dollars and other Arbacia individuals, as well as some algae (Harvey 1956, Karlson 1978, Cobb & Lawrence 2005. The diet of South Atlantic A. dufresnei is mainly carnivorous (Penchaszadeh 1979, Penchaszadeh & Lawrence 1999. The Pacific A. spatuligera showed preference for animal food over common species of algae from its habitat (Silva et al. 2004). Moreover, some observations indicate omnivorous or carnivorous behaviour of A. lixula outside the Mediterranean (Marques 1984, Oliveira 1991, Tavares & Borzone 2005.The crucial importance of sea urchins in shaping benthic ecosystems (Lawrence 1975) has been demonstrated by many ecological experiments along the Mediterranean coasts (e.g. Benedetti-Cecchi & Cinelli 1995, Sala & Zabala 1996, Benedetti-Cecchi et al. 1998, Palacín et al. 1998a, Bulleri et al. 1999, Guidetti et al. 2004, Bonaviri et al. 2011. The underlying premise in these ...