The significance of hatch date for the growth and survival of the sandeel,Ammodytes marinus, was investigated using otolith microstructure. Hatch dates of 2 to 6 mo-old juvenileA. marinus caught near Shetland were compared between 1990 and 1992, during which period year-class strength varied by more than an order of magnitude. The hatch-date distribution of juveniles in the 1992 year-class was compared with that estimated directly from the abundance of newly emerged larvae on the spawning grounds. The extent of larval hatching periods in 1990 and 1991 was also estimated from continuous plankton-recorder data. There were significant differences in hatching periods between all three years, hatching in 1990 and 1992 being markedly earlier than the long-term mean peak in hatching indicated from archival data. Most individuals from the 1991 year-class attained a larger size by July than those in other year-classes, despite hatching later. Variation in individual growth rates both within and between year-classes indicated that there was a seasonal cycle of growth opportunity in all years investigated. The study suggests that the degree of coupling between hatching and the onset of spring secondary production may be an important contributory factor to year-class variability in this species.

Timing of hatching inAmmodytes marinus from Shetland waters and its significance to early growth and survivorship

Pacific sand lance Ammodytes hexapterus in Kachemak Bay, Alaska, showed no sexual dimorphism in length-to-weight (gonad-free) ratio or length-at-age relationship. Most matured in their second year, males earlier in the season than females, but females (31%) attained a higher gonadosomatic index than males (21%). Sand lance spawned intertidally once each year in late September and October on fine gravel or sandy beaches soon after the seasonal peak in water temperatures. Sand lance in Cook Inlet and Prince William Sound displayed similar maturation schedules. Schools were dominated 2: 1 by males as they approached the intertidal zone at a site where spawning has taken place for decades. Sand lance spawned vigorously in dense formations, leaving scoured pits in beach sediments. Fecundity of females (93–199 mm) was proportional to length, ranging from 1468 to 16 081 ova per female. About half of the overall spawning school fecundity was derived from age group 1 females (55% of the school by number). Spawned eggs were 1·02 mm in diameter, demersal, slightly adhesive, and deposited in the intertidal just below the waterline. Sand lance embryos developed over 67 days through periods of intertidal exposure and sub-freezing air temperatures.

Maturation, fecundity, and intertidal spawning of Pacific sand lance in the northern Gulf of Alaska

Fish were sampled with beach seines and small-meshed beam trawls in nearshore (<1 km) and shallow (<25 m) habitats on the southern coast of Kachemak Bay, Cook Inlet, Alaska, from June to August, 1996–1998. Fish distributions among habitats were analysed for species composition, catch-per-unit-effort (CPUE) and frequency of occurrence. Two oceanographically distinct areas of Kachemak Bay were sampled and compared: the Outer Bay and the Inner Bay. Outer Kachemak Bay is exposed and receives oceanic, upwelled water from the Gulf of Alaska, whereas the Inner Bay is more estuarine. Thermohaline properties of bottom water in the Outer and Inner Bay were essentially the same, whereas the Inner Bay water-column was stratified with warmer, less saline waters near the surface.

Distribution and abundance of pelagic schooling fish corresponded with area differences in stratification, temperature and salinity. The Inner Bay supported more species and higher densities of schooling and demersal fish than the Outer Bay. Schooling fish communities sampled by beach seine differed between the Outer and Inner Bays. Juvenile and adult Pacific sand lance (Ammodytes hexapterus), Pacific herring (Clupea harengus pallasi), osmerids (Osmeridae) and sculpins (Cottidae) were all more abundant in the Inner Bay. Gadids (Gadidae) were the only schooling fish taxa more abundant in the Outer Bay. Thermohaline characteristics of bottom water were similar throughout Kachemak Bay. Correspondingly, bottom fish communities were similar in all areas. Relative abundances (CPUE) were not significantly different between areas for any of the five demersal fish groups: flatfishes (Pleuronectidae), ronquils (Bathymasteridae), sculpins (Cottidae), gadids (Gadidae) and pricklebacks (Stichaeidae).

Nearshore Fish Distributions in an Alaskan Estuary in Relation to Stratification, Temperature and Salinity

Parallel structure among environmental gradients and three trophic levels in a subarctic estuary

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, 229 437 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, her- ring) 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 essen- tially 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 sig- nificantly 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.

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Oceanographic conditions structure forage fishes into lipid-rich and lipid-poor communities in lower Cook Inlet, Alaska, USA

Three waves of spawning Pacific sand lance (Ammo-dytes hexapterus) entered the Port Moller estuary from mid-January to late May 1990 Each wave laid its eggs on sand in lower Moller Bay with the center of egg distribution about 14 km inside the estuary After incubation for 45 to 94 d, each cohort of eggs hatched out over a 41- to 63-d period Larvae moved at a rate of 021 knvd“1 toward a deep fjordlike basin at the head of Herendeen Bay inside the estuary about 20 km southwest of the center of hatch The basin has the lowest flushing rate of the estuary, and unlike the rest of Port Moller, it is vertically stratified, which allows the development of a spring-summer zooplankton community with greater biomass than any other location in the estuary Larvae may have moved to the basin to enhance growth or to avoid offshore transport to areas of low food abundance, but we cannot demonstrate a direct link between growth and habitat We conclude that the Port Moller sand lance stock has an estuarine early life history that evolved in response to the unique physical conditions of the Port Moller estuary–a shallow, well-mixed site with sandy substrate that is suitable for incubation of demersal eggs next to a deep, vertically stratified fjord with a rich zooplankton community that is suitable for rearing of larvae

H. David Warburton

Papers

Pacific sand lance of the Port Moller estuary, southeastern Bering Sea: an estuarine^dependent early life history