Temporal variability in marine feeding of sympatric Arctic charr and sea trout

The marine feeding pattern of anadromous brown trout (sea trout) Salmo trutta and Arctic charr Salvelinus alpinus was studied during June to August in 1992–1993 and 2000–2004 in a fjord in northern Norway. In general sea trout fed proportionally more on fishes than on crustaceans and insects (81, 1 and 18% by mass, respectively) by comparison with Arctic charr (52, 25 and 22% by mass, respectively). Herring Clupea harengus dominated the total fish diet of both species, but the Arctic charr also fed significantly on gadoids and sandlance Ammodytes spp. While sea trout became virtually all piscivorous at fork lengths ( L _F) ≥250 mm, the Arctic charr was ≥400 mm L _F before shifting totally to a fish diet. Despite annual variation in diet and forage ratios, there was a clear shift in diet from 1992–1993 to 2000–2004. Sandlance and different crustaceans constituted most of the diet during the initial period with a shift towards gadoids and especially herring during the latter period. This shift seemed to be associated with a high abundance of herring larvae during the latter sampling period, indicating a preferential selection on herring when present, particularly by sea trout. Furthermore, an index indicated dietary overlap in years with intensive feeding on herring of both species, and usually differences in the trophic ecology during years feeding mostly on other prey species. In combination, it was hypothesized that the two species reflect the type of marine prey present within a fjord system over time, and therefore provide an index of variation in the production and biological diversity of their potential prey within fjords.     

Ontogenetic changes in temperature preference of Atlantic cod

Final thermal preferendum ( T ) experiments were conducted in a horizontal thermal gradient tank from the beginning of August 2001 to mid‐November 2001 using Atlantic cod Gadus morhua from 6·5 to 79·0 cm fork length ( L _F). The value of T varied significantly ( P  < 0·005) with L _F( T  = 7·23–0·054 L _F), with smaller (younger) fish choosing higher temperatures than larger (older) fish. The preferendum varied from 6·9° C for fish of 6·5 cm to 3·0° C for those of 79·0 cm. Experiments comparing fish positions in the gradient tank between thermal gradients of 0·5–11·0 and 4·5–14·5° C demonstrated that fish positions were determined by temperature selection instead of undesirable tank effects. This study is the first to demonstrate the effect of ontogeny on temperature preferences of a marine fish species.     

Reproductive biology of gag in the southern Gulf of Mexico

Aspects of the reproductive biology of gag Mycteroperca microlepis in the southern Gulf of Mexico were studied by following seasonal variations in the gonado‐somatic index and through histological examination of gonads. Gag were collected from inshore and offshore waters of the Campeche Bank, Yucatan, Mexico, between April 1996 and December 2001. This species is a protogynous hermaphrodite, and appeared to be depth‐size distributed. The smallest gag (9–49 cm L _F) collected were all juvenile females, and were caught in inshore waters (1–10 m depth), while the largest (49–116 cm L _F), mainly adult females, males and transitionals, were captured in offshore waters (33–167 m depth). Overall the offshore male to female ratio was female‐biased (1 : 3·3) and differed significantly from unity. The species spawns at depths of c . 50–53 m, from early winter to mid‐spring, with peak spawning activity occurring between January and March. Fifty per cent of females reached first maturity at 72·1 cm L _F. At 103 cm L _F, 50% of sampled females had changed into males. Gag can be considered a monandric species, and sexual transition for this grouper seemed to occur in fish distributed within a narrow size range (85–111 cm L _F). The results are compared with those of other authors for gag stocks from the south‐east Atlantic coast of the U.S.A. and the north‐east Gulf of Mexico.     

Diet of brown trout in relation to variation in abundance and size of pelagic fish prey

The seasonal diet of a predator, brown trout Salmo trutta [total length ( L _T) 17–69 cm] and simultaneous density and size‐structure of prey populations, vendace Coregonus albula and smelt Osmerus eperlanus (4–16 cm L _T), in a large boreal lake were analysed and compared in 2001 and 2002. The upper L _T limit for consumed prey was c . 40% of the predator L _T. All brown trout, however, preferred small (<10 cm L _T) and avoided large (≥10 cm L _T) prey. The results also suggested that equal densities of similar‐sized (4–10 cm L _T) fish of the two prey species led to random foraging on these species by brown trout, but if either one of the prey species predominated (>50%) in the lake, brown trout shifted to foraging on this species almost exclusively. Brown trout diets thus reflected the density dynamics of the two alternative prey species.     

Geographical differences in marine feeding of Atlantic salmon post‐smolts in Norwegian fjords

Stomach content analyses were conducted on Atlantic salmon Salmo salar post‐smolt (average size, 119–154 mm fork length, L _F) caught in eight large Norwegian fjord systems along a north–south geographical axis during 1998–2001. In general, post‐smolts from southern Norway showed low feeding intensity in the fjords, whereas extensive feeding was observed in fjords in the northern and middle parts of Norway. The marine diet mainly included different crustaceans and in particular marine pelagic fish larvae (sand-eels Ammodytes spp., herring Clupea harengus and gadoids), but with a substantial spatial and annual variation in prey diversity and feeding intensity. Insects were most frequently taken in the estuary, although fishes often made a large contribution in mass. In contrast, fishes, and to some extent various crustaceans (particularly Hyperiidae, Gammaridae, Euphausiacea and Copepoda) dominated the diet in the middle and outer parts of the fjords, where post‐smolts also fed more extensively than in the inner part. The results indicate that extensive feeding immediately after sea entrance may be more common for post‐smolts in the northern and middle parts of Norway, than in the southern fjords. The observed differences in post‐smolt feeding may be due to spatial and temporal differences in prey availability within and between the different types of fjord systems, and this might influence post‐smolt growth and survival.     

Variation in the diet of Genypterus blacodes(Ophidiidae) around the Falkland Islands

The diet change with size, season and area was investigated using the stomachs of 496 kingclip Genypterus blacodes collected around the Falkland Islands (south‐west Atlantic) between August 2001 and September 2002. The key prey species were rockcod Patagonotothen spp., benthic isopods and Patagonian grenadier Macruronus magellanicus . Kingclip <50 cm total length ( L _T) fed mainly on crustaceans and small fishes. With size the diet shifted away from crustaceans towards Patagonotothen spp. in kingclip 50–100 cm L _T, and finally towards larger fishes such as M. magellanicus and Micromesistius australis australis in kingclip >100 cm L _T. The niche breadth was highest in fish >100 cm L _T and the lowest in fish <50 cm L _T. The larger kingclip generally selected larger individuals of the same prey species, with the exception of the Patagonian squid Loligo gahi , where all ingested squid were of similar size, regardless of the predator length. The importance of the main prey species varied substantially between five consequent seasons studied, and appeared to follow the seasonal abundance and availability of prey. The spatial variability in the diet was found in kingclip caught in regions occupied by transformed temperate and sub‐Antarctic waters. The rockcod, which is available throughout the year around the Falkland Islands, was the most important prey in the kingclip diet. Kingclip takes advantage of other seasonally abundant prey species during their seasonal migrations ( e.g . L. gahi ) and also scavenge on discards from fishing vessels when available.     

Feeding ecology of silver hake larvae on the Western Bank, Scotian Shelf, and comparison with Atlantic Cod

The feeding ecology of the larvae of silver hake Merluccius bilinearis was examined during two time periods (October 1998 and December 1992) on the Western Bank, Scotian Shelf, north‐west Atlantic, and compared with the feeding ecology of Atlantic cod Gadus morhua larvae collected in the same samples in December 1992. During both time periods silver hake exhibited strong selection for late stage copepodids and adult copepods at a small size (>3·5 mm total length, L _T). The niche width measured as the diet breadth index ( I _DB) of silver hake declined rapidly as they increased in size and remained relatively constant from 3 to 11 mm L _T, during each time period. Atlantic cod larvae exhibited a broader niche width that was curvilinear over the same L _T. Atlantic cod were also less selective than silver hake, incorporating both naupliar and early stage copepodids in their diets throughout the length classes examined. Simple isometric relationships did not explain the differences in diet, as Atlantic cod larvae continued to feed on early stages of copepods at large size, while silver hake larvae quickly switched to large prey items. The strong selection and narrow I _DB observed for silver hake probably reflects adaptation to spawning during the periods between major secondary production peaks in temperate waters.     

Seasonal, diel and ontogenetic variation in feeding patterns of small yellow croaker in the central Yellow Sea

Stomach contents of 1603 small yellow croaker Pseudosciaena polyactis , sampled from seasonal bottom trawl surveys in the central Yellow Sea between March 2001 and January 2002, were examined. The results showed that small yellow croaker was a carnivorous predator and >30 prey species were identified from stomach contents analysis. Crustaceans (mainly euphausiids and decapods) were the most important prey, occurring in 93·1% of the stomachs containing food, and accounting for 77·6% of the total food by mass. Feeding activity was highest in autumn and lowest in spring and winter. Decapods were more important in summer, whereas euphausiids were more important during other seasons. Ontogenetic differences were found in the diet composition and feeding activity within the range of size (standard length, L _S) studied. The importance of fishes and decapods increased with L _S, whereas euphausiids, copepods and amphipods decreased in importance with L _S. Dietary breadth increased markedly for adults. A positive relationship was found between L _S and prey size. In each season the maximum diel feeding activity occurred at 0800 and 2400 hours, indicating that there was crepuscular and nocturnal feeding by small yellow croaker.     

Age, growth and reproduction of the barrelfish Hyperoglyphe perciformis (Mitchill) in the western North Atlantic

Otoliths ( n = 847) and gonads ( n = 817) were collected from barrelfish Hyperoglyphe perciformis that were captured by commercial fishermen in the waters off South Carolina and Georgia in 1995, 1997 and 2001–2006. Of the otoliths collected, 97% were aged successfully, and specimens sampled ranged from 5 to 85 years, with a median age of 12 years. The von Bertalanffy growth parameters yielded the equation: L_t = 857·8{1 − e^{−0·0985[ t −(−8·95)]}}, where L_t is fork length ( L _F) at time t . Through histological examination, 94% of the gonads assessed were assigned to a sex and reproductive class. Females spawned from September to May with a peak from November to January. Males spawned year round, but had a peak from September to April. The sex ratio (M:F) for this population was 1:1·34. The smallest mature female was 605 mm L _F and the youngest immature female was 697 mm L _F. Estimates of L _F and age at 50% maturity ( L ₅₀ and A ₅₀) for females were 660 mm L _F (95% CI = 633–667 mm L _F) and 6·08 years (95% CI = 3·50–7·27 years), respectively. The youngest mature male was 575 mm L _F and the oldest immature male was 762 mm L _F, and no estimates of L ₅₀ or A ₅₀ were made for males. It was determined that barrelfish exhibit the typical characteristics of long life span, slow growth and high age at maturity seen in other deepwater fishes, and that care should be taken to manage this species accordingly.     

Functional response of juvenile pink and chum salmon: effects of consumer size and two types of zooplankton prey

Feeding rate experiments were conducted for pink salmon Oncorhynchus gorbuscha fry [mean fork length ( L _F) 39 mm], juveniles (103–104 mm L _F) and juvenile chum salmon Oncorhynchus keta (106–107 mm L _F). Fishes were presented with small copepod ( Tisbi sp.) or larger mysid shrimp ( Mysidopsis bahia ) prey at varying densities ranging from 1 to 235 prey l⁻¹ in feeding rate experiments conducted at water temperatures ranging from 10·5 to 12·0° C under high light levels and low turbidity conditions. Juvenile pink and chum salmon demonstrated a type II functional response to mysid and copepod prey. Mysid prey was readily selected by both species whereas the smaller bodied copepod prey was not. When offered copepods, pink salmon fry fed at a higher maximum consumption rate (2·5 copepods min⁻¹) than larger juvenile pink salmon (0·4 copepods min⁻¹), whereas larger juvenile chum salmon exhibited the highest feeding rate (3·8 copepods min⁻¹). When feeding on mysids, the maximum feeding rate for larger juvenile pink (12·3 mysids min⁻¹) and chum (11·5 mysids min⁻¹) salmon were similar in magnitude, and higher than feeding rates on copepods. Functional response models parameterized for specific sizes of juvenile salmon and zooplankton prey provide an important tool for linking feeding rates to ambient foraging conditions in marine environments, and can enable mechanistic predictions for how feeding and growth should respond to spatial-temporal variability in biological and physical conditions during early marine life stages.     

Seasonal growth of Atlantic cod: effects of temperature, feeding and reproduction

Growth of 2659 Atlantic cod Gadus morhua aged 4 to 9 years examined in Placentia Bay, Newfoundland, peaked in most cases in June and was at a minimum in October or November. Water temperature, partial fullness index ( I _P) and gonado‐somatic index ( I _G) explained between 31 and 52% of the monthly variability in growth. Temperature and I _P of capelin Mallotus villosus had significant effects on growth of all age groups and explained most of the variance for ages 6–8 and 4–5 years, respectively. The I _P of large invertebrates (ages 4 to 7 years), sandlance ( Ammodytes sp. age 6 years) and demersal fishes (age 9 years) had age‐specific effects in the model. Overall, amphipods, decapods and echinoderms dominated the Atlantic cod diet in most seasons, but fish consumption by Atlantic cod was high in June and July, particularly on capelin. The rapid increase in somatic mass during June and July occurred despite cold water temperatures ( < 3° C at 50 m) and moderate to high gonado‐somatic index. The findings of this study suggest that when food was not a limiting factor, growth tended to increase even when Atlantic cod occupied colder waters, but when food was limiting, the opposite may have occured.     

Lacustrine growth of juvenile pink salmon and a comparison with sympatric sockeye salmon

The growth rates of naturally sympatric juvenile pink Oncorhynchus gorbuscha and sockeye Oncorhynchus nerka salmon were compared in a common lacustrine environment in south‐west Alsaka, an unusual opportunity given the normal disparity in freshwater residence time of these two species. Fork length ( L _F) frequency distributions of juvenile pink salmon caught in the lake during the summer in 1991 and 1999–2003 indicated a growth rate of 0·54 mm day⁻¹, 54% greater than the estimated growth rate of juvenile sockeye salmon sampled from 1958 to 2003 (0·35 mm day⁻¹). Examination of daily growth rings on otoliths indicated that pink salmon in Lake Aleknagik grew an average of 1·34 mm day⁻¹ in 2003 but sockeye salmon grew only 0·63 mm day⁻¹(average specific growth rates were 3·0 and 1·8% day⁻¹, respectively). Pink salmon increased from c . 32 mm L _F and 0·2 g at emergence to 78 mm L _F and 3·0 g within 3–4 weeks. After experiencing these rapid growth rates, the pink salmon appeared to leave the lake by late July in most years. The diets of pink and sockeye salmon in the littoral zone of the lake were very similar; >80% of the stomach contents consisted of adult and pupal insects and the remainder was zooplankton. This high degree of diet overlap suggested that the observed differences in growth rate were not attributable to variation in prey composition.     

Growth of silver hake within the U.S. continental shelf ecosystem of the northwest Atlantic Ocean

Von Bertalanffy growth parameters were estimated for silver hake Merluccius bilinearis within four regions of the United States continental shelf ecosystem of the northwest Atlantic Ocean during 1975–1980, 1982–1987 and 1988–1992. Differences in silver hake growth were found between the Middle Atlantic and southern Georges Bank regions ( P <0–01) over all three time periods. Growth also differed between northern Georges Bank and the Gulf of Maine ( P <0–01) during 1975–1980 and 1982–1987, and significant differences between southern Georges Bank and northern Georges Bank were found only during 1982–1987. Silver hake in the Gulf of Maine exhibited larger asymptotic sizes, but attained their asymptotic size at slower rates (L₀₀= 47 cm and K =0–33) in comparison to silver hake in the Middle Atlantic ( L₀₀= 43 cm and K =0–51), while growth parameter estimates for silver hake on Georges Bank were intermediate between these. Although significant differences in growth of silver hake between the various regions are consistent with separate stocks in the U.S. continental shelf ecosystem, the growth characteristics from the different regions are inconsistent with the presently assigned stock boundaries. (c) 1996 The Fisheries Society of the British Isles     

Histological investigation on the ovarian cycle of the bluefin tuna in the western and central Mediterranean

The histological analysis of eastern Atlantic bluefin tuna Thunnus thynnus ovaries caught from February to September 1999–2000, made it possible to distinguish the presence of seven oocyte developmental stages and allowed the characterization of six time-dependent ovary maturity stages. The ovaries of mature (fork length, L _F ≥ 110 cm) bluefin tuna were non-active from August (spent period) to March (quiescent period) when they contained only perinucleolar-stage oocytes. Ovary development started in April to early May (recrudescent period) with the appearance of oocytes at the lipid stage. Vitellogenesis appeared in mid-May (ripening period) and post-vitellogenesis occurred in late May to mid-June (pre-spawning period). In late June to early July, hydrated oocytes, a sign of imminent spawning, were found only in specimens caught in Balearic waters. Females ranging between 100 and 110 cm L _F, captured during the recrudescent and ripening periods, had the largest oocytes at the lipid stage, most of which were degenerating. An extensive vitellogenic atresia was observed in the ovaries of five females caught during the spawning period in non-spawning areas.     

Reproductive biology of the threatened golden galaxias Galaxias auratus Johnston and the influence of lake hydrology

Golden galaxias Galaxias auratus (31–235 mm fork length, L _F) were collected monthly from littoral habitats in Lakes Crescent and Sorell, Tasmania, Australia, between July 2000 and December 2002. Spawning habitats were identified and monitored in both lakes, and surveyed in Lake Crescent. Trends in gonado-somatic indices and reproductive stages of development indicated that gonad development in both sexes begins in midsummer and peaks in late autumn to early winter. Males mature at smaller sizes (50% at 52 mm L _F) than females (50% at 76 mm L _F), larger individuals are predominately females (95% of fish ≥138 mm L _F), and overall male to female ratios are female biased ( c . 1:2). Spawning occurs late autumn to early spring (water temperatures = 1·4–9·7° C) with peaks in spawning activity in winter (mean water temperatures <5° C). Demersal adhesive eggs ( c. 1·5 mm diameter) were found on cobble substrata ( c. 20–250 mm diameter) in littoral areas ( c. 0·2–0·6 m deep) and fecundity of fish 71–181 mm L _F ranged from 619 to 14 478 eggs. The rate of change in water level over the 20 days prior to monthly sampling was important in explaining the occurrence of spent fish and this accounted for temporal differences in spawning between the populations. Lake hydrology influences the reproductive cycle of G. auratus by possibly providing a stimulus for spawning and it controls the availability of spawning habitat in Lake Crescent. Seasonal hydrological cycles ( i.e. rises during late autumn to winter) and a minimum water level of 802·20 m Australian Height Datum in Lake Crescent during autumn (above which littoral areas of cobble substratum are inundated) are critical to G. auratus populations.     

The effects of body mass and feeding on metabolic rate in small juvenile Atlantic cod

Apparent specific dynamic action (SDA) amplitude in young juvenile Atlantic cod Gadus morhua (1 to 8 g wet mass), fed a standardized meal and then exercised in a circular swimming respirometer at a constant swimming speed of 0·5 ± 0·3 body lengths s^-1, occurred within l h after feeding in all juveniles. SDA amplitude were 1·4 to 1·8 times higher in fed fish compared to unfed fish, and rates of oxygen consumption decreased as body mass increased. SDA duration had a tendency to decrease with increasing body mass and was shortest, at 6 h, in the smallest fish (1–1·5 g), but increased to 10–11 h in the largest fish. Apparent SDA in fed fish ( R _r) scaled with a mass exponent of 0·89, while maximum metabolic rate ( R _max) determined by chasing fish to exhaustion and then measuring oxygen consumption for 12 h, and unfed routine metabolic rate (R_r) scaled with a mass exponent of 0·79 and 0·76 respectively. Relative aerobic scope ( R _max– unfed R _r) did not appear to vary over the 1 to 8 g increase in wet mass. These results show that as body mass increased in young juvenile Atlantic cod: (1) apparent SDA ( R _f) increased more rapidly than R _max, and (2) apparent SDA took up >98% of the relative aerobic scope and that young Atlantic cod allocated most of the energy to growth, and left little for other metabolic activities.     

Spatially structured interactions between a migratory pelagic predator, the Norwegian spring-spawning herring Clupea harengus L., and its zooplankton prey

Spring-spawning herring Clupea harengus was patchily distributed over large parts of the Norwegian Sea in May 1995–2005, during the early phase of the annual feeding migration. Overall, herring tended to be found in areas with intermediate biomasses of zooplankton prey, intermediate water temperatures and relatively high salinities. Herring had more food in their stomachs in areas of relatively low water temperature and high herring abundance. Hydrographical conditions revealed that herring was feeding mainly within Atlantic water masses, and more intensely in western and northern regions of the Norwegian Sea. Zooplankton biomass was patchily distributed, and was generally higher towards the western parts of the Norwegian Sea. Here, zooplankton biomass in year _{i +1} was also negatively associated with herring spawning stock biomass in year _i , while there was no evidence for such an association in the eastern region; indicating that herring may have a geographically structured 'top–down' effect on the recruitment of its zooplankton prey. The fact that herring was not typically associated with the areas containing the greatest zooplankton biomasses may reflect that the fish had not yet reached the most profitable feeding grounds or alternatively that herring was depleting zooplankton biomass.     

Thermal tolerance of a northern population of striped bass Morone saxatilis

Thermal tolerance of age 0+ year Shubenacadie River (Nova Scotia, Canada) striped bass Morone saxatilis juveniles (mean ± s . e . fork length, L _F, 19·2 ± 0·2 cm) acclimated in fresh water to six temperatures from 5 to 30° C was measured by both the incipient lethal technique (72 h assay), and the critical thermal method ( C _m). The lower incipient lethal temperature ranged from 2·4 to 11·3° C, and the upper incipient lethal temperature ( I _U) from 24·4 to 33·9° C. The area of thermal tolerance was 618° C². In a separate experiment, the I _U of large age 2+ year fish (34·4 ± 0·5 cm L _F) was 1·2 and 0·6° C lower ( P < 0·01) than smaller age 1+ year fish (21·8 ± 0·5 cm L _F) at acclimation temperatures of 16 and 23° C. Using the C _m, loss of equilibrium occurred at 27·4–37·7° C, loss of righting response at 28·1–38·4° C and onset of spasms at 28·5–38·8° C, depending on acclimation temperature. The linear regression slopes for these three responses were statistically similar (0·41; P > 0·05), but the intercepts differed (25·3, 26·0 and 26·5° C; P < 0·01). The thermal tolerance of this northern population appears to be broader than southern populations.     

Size variability of juvenile Atlantic salmon: links to environmental conditions

Climate change models predict a 2 to 6° C increase in air temperature within the next 100 years in the Maritime Provinces of eastern Canada. Higher air temperatures are expected to contribute to increased water temperatures, alterations in stream flow conditions, and ultimately reductions in fish growth. Mean annual size-at-age of juvenile Atlantic salmon Salmo salar decreased in the Northwest Miramichi and Southwest Miramichi Rivers between 1971–1999. Lengths-at-age of juveniles were significantly correlated between the two rivers. For Atlantic salmon parr, stronger associations between inter-cohort fork length ( L _F) than intra-cohort L _F were observed, suggesting that environmental conditions in the current year of growth have the more significant effects on size of age 2 year parr than conditions encountered the previous year by age 1 year parr of the same cohort. Fork lengths of parr were significantly and negatively associated with spring air and water temperatures. In the Miramichi River, increases in air and water temperature as predicted from climate change models may adversely affect growth of juvenile Atlantic salmon parr, reducing the overall productivity of the Atlantic salmon populations in this region.