Oceanographic conditions structure forage fishes into lipid-rich and lipid-poor communities in lower Cook Inlet, Alaska, USA
Forage fishes were sampled with a mid-water trawl in lower Cook Inlet, Alaska, USA, from late July to early August 1996 to 1999. We sampled 3 oceanographically distinct areas of lower Cook Inlet: waters adjacent to Chisik Island, in Kachemak Bay, and near the Barren Islands. In 163 tows using a mid-water trawl, 229437 fishes with fork length <200 mm were captured. More than 39 species were captured in lower Cook Inlet, but Pacific sand lance Ammodytes hexapterus, juvenile Pacific herring Clupea pallasi, and juvenile walleye pollock Theragra chalcogramma comprised 97.5% of the total individuals. Both species richness and species diversity were highest in warm, low-salinity, weakly stratified waters near Chisik Island. Kachemak Bay, which had thermohaline values between those found near Chisik Island and the Barren Islands, had an intermediate value of species richness. Species richness was lowest at the Barren Islands, an exposed region that regularly receives oceanic, upwelled water from the Gulf of Alaska. Non-metric multidimensional scaling (NMDS) was used to compute axes of species composition based on an ordination of pairwise site dissimilarities. Each axis was strongly rank-correlated with unique groups of species and examined separately as a function of environmental parameters (temperature, salinity, depth), area, and year. Oceanographic parameters accounted for 41 and 12% of the variability among forage fishes indicated by Axis 1 and Axis 2, respectively. Axis 1 also captured the spatial variability in the upwelled area of lower Cook Inlet and essentially contrasted the distribution of species among shallow, nearshore (sand lance, herring) and deep, offshore (walleye pollock) habitats. Axis 2 captured the spatial variability in forage fish communities from the north (Chisik Island) to the south (Barren Islands) of lower Cook Inlet and essentially contrasted a highly diverse community dominated by salmonids and osmerids (warmer, less saline) with a fish community dominated by Pacific sand lance (colder, more saline). Axis 3 reflected the negative spatial association of capelin Mallotus villosus and Pacific cod Gadus macrocephalus. Correlations of year with Axes 1 and 3 indicate that from 1996 to 1999 the forage fish community significantly decreased in lipid-poor gadids (walleye pollock and Pacific cod), and significantly increased in lipid-rich species such as Pacific sand lance, Pacific herring, and capelin.