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Resource partitioning in polymorphic fish species is expected to be altered by human exploitation, as individual specialization is density dependent in many vertebrates. We tested this hypothesis using the ballan wrasseLabrus bergyltaas a model species. We compared the isotope niches of the plain and spotted morphs of the species in a marine protected area (MPA) and in adjoining areas open to fishing, both off Galicia (NW Spain). Underwater visual census confirmed a 3-fold increase in the biomass of ballan wrasse off the Cíes Islands compared to areas open to recreational fishing, thus demonstrating that populations outside MPAs are well below carrying capacity. The stable isotope ratio of C revealed differences in the resource use patterns of plain and spotted ballan wrasses both in areas open and closed to fishing, as plain wrasses were always depleted in13C compared to sympatric spotted ones. The stable isotope ratio of N showed that plain ballan wrasses foraged consistently at a higher trophic level than spotted ones in areas of high population density closed to recreational fishing, whereas differences did not exist or were reversed in areas open to fishing. These results demonstrate that the pattern of trophic resource partitioning between 2 morphs of the ballan wrasse is density dependent and that plain and spotted ballan wrasses likely had different ecological niches in pristine ecosystems.

Explaining the mechanisms driving niche partitioning among species is of great importance in ecology. Unlike the fundamental niche, a species' realised niche can only be measured in situ, as a result of biotic and abiotic interactions defining its size. Following current methodology, the realised niche of a species is often influenced by the rare and divergent individuals of the community sampled. In this study, using fish on coral and temperate reefs as an example, behavioural empirical data were collected to estimate realised niche sizes and niche overlaps. Niche measurements were made using the total area of the convex hull (TA), but as an alternative, a metric not as strongly influenced by sample size, standard ellipse area (SEA), was also used. A comprehensive description is given, and contextdependent pros and cons of using both metrics are discussed. Additionally, an alternative sample size correction was presented for both metrics. The analyses re vealed large differences in the sizes of realised niches and their overlaps between species depending on the measurement metric used. Regardless of the species, niche size and overlap were always larger for TA than SEA. Increasing sample size re - duced niche size variability for both TA and SEA, but the variation was always smaller for SEA than TA. We successfully adapted the SEA metric for analysis of behavioural niche components and demonstrated that measuring niche sizes using the 2 metrics, each with their own strengths and weaknesses, can produce contradictory results, the ecological consequences of which are likely to be important.

Fishes and invertebrates were collected from the continental slope southwest of the Balearic Islands, Mediterranean, in October 1996 and May 1998 from 3 faunally-distinct depth zones: 200 to 800 m (upper slope), 800 to 1425 m (middle slope) and 1425 to 1800 m (lower slope). Differ- ences in δ 15 N and δ 13 C between October 1996 and May 1998 were observed only in the crustaceans Aristeus antennatus and Nematocarcinus exilis. There were few differences among depths in δ 15 N or δ 13 C of benthic fishes or crustaceans: the shrimps A. antennatus and Acanthephyra eximia showed higher δ 15 N values at greater depth, and δ 13 C values of the fishes Alepocephalus rostratus (Ale- pocephalidae) and Coelorhynchus coelorhynchus (Macrouridae) also tended to increase with in- creasing depth. Differences in plankton and suprabenthos were also observed among zones (at depth, greater δ 15 N values of gelatinous plankton, calanoid copepods and peracarids were recorded, along with smaller δ 13 C values of gelatinous plankton and peracarids). These are attributed to a microbially-mediated increase in δ 15 N and a reduction in δ 13 C of the particulate material remaining after preferential utilization of labile organic matter in the stable and relatively high (13°C) tem- perature of the Mediterranean water column. Strong positive correlations between δ 15 N and δ 13 C data for plankton, fishes and crustaceans indicated a single primary source material for these com- munities, which is attributed to marine snow.