Spatially explicit estimates of length and biomass of Clupea harengus (Norwegian spring spawning herring) from acoustic and trawl surveys in the North-East Atlantic in May 2011

Acoustic estimates of herring and blue whiting abundance were obtained during the surveys using the Simrad ER60 scientific echosounder. The allocation of NASC-values to herring, blue whiting and other acoustic targets were based on the composition of the trawl catches and the appearance of echo recordings. To estimate the abundance, the allocated NASC -values were averaged for ICES-squares (0.5° latitude by 1° longitude). For each statistical square, the unit area density of fish (rA) in number per square nautical mile (N*nm-2) was calculated using standard equations (Foote et al., 1987; Toresen et al., 1998). To estimate the total abundance of fish, the unit area abundance for each statistical square was multiplied by the number of square nautical miles in each statistical square and then summed for all the statistical squares within defined subareas and over the total area. Biomass estimation was calculated by multiplying abundance in numbers by the average weight of the fish in each statistical square then summing all squares within defined subareas and over the total area. The Norwegian BEAM soft-ware (Totland and Godø 2001) was used to make estimates of total biomass.

Spatially explicit estimates of length and biomass of Clupea harengus (Norwegian spring spawning herring) from acoustic and trawl surveys in the North-East Atlantic in May 2007

Acoustic estimates of herring and blue whiting abundance were obtained during the surveys using the Simrad ER60 scientific echosounder. The allocation of NASC-values to herring, blue whiting and other acoustic targets were based on the composition of the trawl catches and the appearance of echo recordings. To estimate the abundance, the allocated NASC -values were averaged for ICES-squares (0.5° latitude by 1° longitude). For each statistical square, the unit area density of fish (rA) in number per square nautical mile (N*nm-2) was calculated using standard equations (Foote et al., 1987; Toresen et al., 1998). To estimate the total abundance of fish, the unit area abundance for each statistical square was multiplied by the number of square nautical miles in each statistical square and then summed for all the statistical squares within defined subareas and over the total area. Biomass estimation was calculated by multiplying abundance in numbers by the average weight of the fish in each statistical square then summing all squares within defined subareas and over the total area. The Norwegian BEAM soft-ware (Totland and Godø 2001) was used to make estimates of total biomass.

The North Sea autumn spawning Herring (Clupea harengus L.) Spawning Component Abundance Index (SCAI)

The North Sea autumn-spawning herring (Clupea harengus) stock consists of a set of different spawning components. The dynamics of the entire stock have been well characterized, but although time-series of larval abundance indices are available for the individual components, study of the dynamics at the component level has historically been hampered by missing observations and high sampling noise. A simple state-space statistical model is developed that is robust to these problems, gives a good fit to the data, and proves capable of both handling and predicting missing observations well. Furthermore, the sum of the fitted abundance indices across all components proves an excellent proxy for the biomass of the total stock, even though the model utilizes information at the individual-component level. The Orkney-Shetland component appears to have recovered faster from historic depletion events than the other components, whereas the Downs component has been the slowest. These differences give rise to changes in stock composition, which are shown to vary widely within a relatively short time. The modelling framework provides a valuable tool for studying and monitoring the dynamics of the individual components of the North Sea herring stock.

Age determination methods for northwest Atlantic species

The successful application of techniques to enhance detection of age marks in biological specimens is of vital importance in fisheries research. This manual documents age determination techniques used by staff at the Woods Hole Laboratory, National Marine Fisheries Service. General information on procedures for preparing anatomical structures is described, together with criteria used to interpret growth patterns and assign ages. Annotated photographs of age structures are provided to illustrate criteria. Detailed procedures are given for the following species: Atlantic herring (Clupea harengus), haddock (Melanogrammus aeglefinus), Atlantic cod (Gadus morhua), pollock (Pollachius virens), silver hake (Merluccius bilinearis), red hake (Urophycis chuss), black sea bass (Centropristis striata), weakfish (Cynoscion regalis), Atlantic mackerel (Scomber scombrus), butterfish (Peprilus triacanthus), redfish (Sebastes fasciatus), summer flounder (Paralichthys dentatus), winter flounder (Pseudopleuronectes americanus), witch flounder (Glyptocephalus cynoglossus), American plaice (Hippoglossoides platessoides), yellowtail flounder (Limanda ferruginea), surf clam (Spisula solidissima), and ocean quahog (Arctica islandica). (PDF file contains 142 pages.)

From the river to the open sea - a critical life phase of young Atlantic salmon migrating from the Simojoki river

Long-term monitoring data collected from wild smolts of Atlantic salmon (Salmo salar) in the Simojoki river, northern Finland, were used in studying the relationships between the smolt size and age, smolt and postsmolt migration, environmental conditions and postsmolt survival. The onset of the smolt run was significantly dependent on the rising water temperature and decreasing discharge of the river in the spring. The mean length of smolts migrating early in the season was commonly higher and the mean age always older than among smolts migrating later. Many of the smolts migrating early in the season and almost all smolts migrating later had started their new growth in spring in the river before their sea entry. Among postsmolts, the time required for emigration from the estuary was dependent on the sea surface temperature (SST) off the river, being significantly shorter in years with warm than cold sea temperatures. After leaving the estuary, the postsmolts migrated southwards along the eastern coast of the northern Gulf of Bothnia, the geographical distribution of the tag recoveries coinciding with the warm thermal zone in spring in the coastal area. After arriving in the southern Gulf of Bothnia in late summer the postsmolts mostly migrated near the western coast, reaching the Baltic Main Basin in late autumn. Until the early 1990s there was only a weak positive association between smolt length and postsmolt survival. However, following a subsequent decrease in the mean smolt size, a significant positive dependence was observed between smolt size and the reported recapture rate of tagged salmon. The differences in recapture rates between smolts tagged during the first and second half of the annual migration season were insignificant, indicating that the seasonal variation in smolt size and age seem to be too small to affect survival. Among the climatic factors examined, the summer SST in the Gulf of Bothnia was most clearly related to the survival of the wild postsmolts. Postsmolt survival appeared to be highest in years when the SST in June in the Bothnian Bay varied between 9 and 12 ºC. In addition, the survival of wild postsmolts showed a significant positive dependence on the SST in July in the Bothnian Sea, but not on the abundance of the prey fish (0+ herring, Clupea harengus and sprat, Sprattus sprattus) in the Bothnian Sea and in the Baltic Main Basin. The results suggest, that if the incidence of extreme weather conditions were to increase due to climatic changes, it would probably reduce the postsmolt survival of wild salmon populations. For improving the performance of hatchery-reared smolts, it could be useful to examine opportunities to produce smolts that are in their smolt traits and abilities more similar to the wild smolts described in this thesis.

Differences in diet of Atlantic bluefin tuna (Thunnus thynnus) at five seasonal feeding grounds on the New England continental shelf

The stomachs of 819 Atlantic bluefin tuna (Thunnus thynnus) sampled from 1988 to 1992 were analyzed to compare dietary differences among five feeding grounds on the New England continental shelf (Jeffreys Ledge, Stellwagen Bank, Cape Cod Bay, Great South Channel, and South of Martha’s Vineyard) where a majority of the U.S. Atlantic commercial catch occurs. Spatial variation in prey was expected to be a primary influence on bluefin tuna distribution during seasonal feeding migrations. Sand lance (Ammodytes spp.), Atlantic herring (Clupea harengus), Atlantic mackerel (Scomber scombrus), squid (Cephalopoda), and bluefish (Pomatomus saltatrix) were the top prey in terms of frequency of occurrence and percent prey weight for all areas combined. Prey composition was uncorrelated between study areas, with the exception of a significant association between Stellwagen Bank and Great South Channel, where sand lance and Atlantic herring occurred most frequently. Mean stomach-contents biomass varied significantly for all study areas, except for Great South Channel and Cape Cod Bay. Jeffreys Ledge had the highest mean stomach-contents biomass (2.0 kg) among the four Gulf of Maine areas and Cape Cod Bay had the lowest (0.4 kg). Diet at four of the five areas was dominated by one or two small pelagic prey and several other pelagic prey made minor contributions. In contrast, half of the prey species found in the Cape Cod Bay diet were demersal species, including the frequent occurrence of the sessile fig sponge (Suberites ficus). Prey size selection was consistent over a wide range of bluefin length. Age 2–4 sand lance and Atlantic herring and age 0–1 squid and Atlantic mackerel were common prey for all sizes of bluefin tuna. This is the first study to compare diet composition of western Atlantic bluefin tuna among discrete feeding grounds during their seasonal migration to the New England continental shelf and to evaluate predator-prey size relationships. Previous studies have not found a common occurrence of demersal species or a pre-dominance of Atlantic herring in the diet of bluefin tuna.

Environmental factors and uncertainty in fisheries management in the northern Baltic Sea

The Baltic Sea is a geologically young, large brackish water basin, and few of the species living there have fully adapted to its special conditions. Many of the species live on the edge of their distribution range in terms of one or more environmental variables such as salinity or temperature. Environmental fluctuations are know to cause fluctuations in populations abundance, and this effect is especially strong near the edges of the distribution range, where even small changes in an environmental variable can be critical to the success of a species. This thesis examines which environmental factors are the most important in relation to the success of various commercially exploited fish species in the northern Baltic Sea. It also examines the uncertainties related to fish stocks current and potential status as well as to their relationship with their environment. The aim is to quantify the uncertainties related to fisheries and environmental management, to find potential management strategies that can be used to reduce uncertainty in management results and to develop methodology related to uncertainty estimation in natural resources management. Bayesian statistical methods are utilized due to their ability to treat uncertainty explicitly in all parts of the statistical model. The results show that uncertainty about important parameters of even the most intensively studied fish species such as salmon (Salmo salar L.) and Baltic herring (Clupea harengus membras L.) is large. On the other hand, management approaches that reduce uncertainty can be found. These include utilising information about ecological similarity of fish stocks and species, and using management variables that are directly related to stock parameters that can be measured easily and without extrapolations or assumptions.

Antipredator behaviour of Baltic planktivores

Predation is an important source of mortality for most aquatic animals. Thus, the ability to avoid being eaten brings substantial fitness benefits to individuals. Predator detection abilities and antipredator behaviour were examined in various planktivores, i.e. the littoral mysids Neomysis integer and Praunus flexuosus, three-spined stickleback Gasterosteus aculeatus larvae, pelagic mysids Mysis mixta and M. relicta, and the predatory cladoceran Cercopagis pengoi, with cues from their respective predators European perch Perca fluviatilis and Baltic herring Clupea harengus membras. The use of different aquatic macrophytes as predation refuges by the littoral planktivores was also examined. All pelagic planktivores and stickleback larvae were able to detect the presence of their predator by chemical cues alone. The littoral mysids N. integer and P. flexuosus responded only when chemical and visual predator cues were combined. The responses of stickleback larvae were stronger to the combined cues than the chemical cue alone. A common antipredator behaviour in all of the planktivores studied was decreased ingestion rate in response to predator cues. N. integer and stickleback larvae also decreased their swimming activity. Pelagic mysids and C. pengoi altered their prey selectivity patterns in response to predator cues. The effects of predator cues on the swarming behaviour of N. integer were examined. Swarming brings clear antipredator advantages to N. integer, since when they feed in a swarm, they do not significantly decrease their feeding rate. However, the swarming behaviour of N. integer was not affected by predation risk, but was instead a fixed strategy. Despite the presence or absence of predator cues, N. integer individuals attempted to associate with a swarm and preferred larger to smaller swarms. In studies with aquatic macrophytes, stickleback larvae and P. flexuosus utilized vegetation as a predation refuge, spending more time within vegetation when under predation threat. The two macroalgal species studied, bladderwrack Fucus vesiculosus and stonewort Chara tomentosa, were preferred by P. flexuosus, whereas Eurasian watermilfoil Myriophyllum spicatum was strongly avoided by N. integer and stickleback larvae. In fact, when in dense patches in aquaria, M. spicatum caused acute and high mortality (> 70%) in littoral mysids, but not in sticklebacks, whereas C. tomentosa and northern watermilfoil M. sibiricum did not. In contrast, only 2-4% mortality in N. integer was observed with intact and broken stems of M. spicatum in field experiments. The distribution of littoral mysids in different vegetations, however, suggests that N. integer avoids areas vegetated by M. spicatum.

Molecular Tuning of Rhodopsins for High Visual Sensitivity in Different Light Environments : Variation in Absorbance Spectrum and Opsin Sequence within and between Species

Visual pigments of different animal species must have evolved at some stage to match the prevailing light environments, since all visual functions depend on their ability to absorb available photons and transduce the event into a reliable neural signal. There is a large literature on correlation between the light environment and spectral sensitivity between different fish species. However, little work has been done on evolutionary adaptation between separated populations within species. More generally, little is known about the rate of evolutionary adaptation to changing spectral environments. The objective of this thesis is to illuminate the constraints under which the evolutionary tuning of visual pigments works as evident in: scope, tempo, available molecular routes, and signal/noise trade-offs. Aquatic environments offer Nature s own laboratories for research on visual pigment properties, as naturally occurring light environments offer an enormous range of variation in both spectral composition and intensity. The present thesis focuses on the visual pigments that serve dim-light vision in two groups of model species, teleost fishes and mysid crustaceans. The geographical emphasis is in the brackish Baltic Sea area with its well-known postglacial isolation history and its aquatic fauna of both marine and fresh-water origin. The absorbance spectrum of the (single) dim-light visual pigment were recorded by microspectrophotometry (MSP) in single rods of 26 fish species and single rhabdoms of 8 opossum shrimp populations of the genus Mysis inhabiting marine, brackish or freshwater environments. Additionally, spectral sensitivity was determined from six Mysis populations by electroretinogram (ERG) recording. The rod opsin gene was sequenced in individuals of four allopatric populations of the sand goby (Pomatoschistus minutus). Rod opsins of two other goby species were investigated as outgroups for comparison. Rod absorbance spectra of the Baltic subspecies or populations of the primarily marine species herring (Clupea harengus membras), sand goby (P. minutus), and flounder (Platichthys flesus) were long-wavelength-shifted compared to their marine populations. The spectral shifts are consistent with adaptation for improved quantum catch (QC) as well as improved signal-to-noise ratio (SNR) of vision in the Baltic light environment. Since the chromophore of the pigment was pure A1 in all cases, this has apparently been achieved by evolutionary tuning of the opsin visual pigment. By contrast, no opsin-based differences were evident between lake and sea populations of species of fresh-water origin, which can tune their pigment by varying chromophore ratios. A more detailed analysis of differences in absorbance spectra and opsin sequence between and within populations was conducted using the sand goby as model species. Four allopatric populations from the Baltic Sea (B), Swedish west coast (S), English Channel (E), and Adriatic Sea (A) were examined. Rod absorbance spectra, characterized by the wavelength of maximum absorbance (λmax), differed between populations and correlated with differences in the spectral light transmission of the respective water bodies. The greatest λmax shift as well as the greatest opsin sequence difference was between the Baltic and the Adriatic populations. The significant within-population variation of the Baltic λmax values (506-511 nm) was analyzed on the level of individuals and was shown to correlate well with opsin sequence substitutions. The sequences of individuals with λmax at shorter wavelengths were identical to that of the Swedish population, whereas those with λmax at longer wavelengths additionally had substitution F261F/Y in the sixth transmembrane helix of the protein. This substitution (Y261) was also present in the Baltic common gobies and is known to redshift spectra. The tuning mechanism of the long-wavelength type Baltic sand gobies is assumed to be the co-expression of F261 and Y261 in all rods to produce ≈ 5 nm redshift. The polymorphism of the Baltic sand goby population possibly indicates ambiguous selection pressures in the Baltic Sea. The visual pigments of all lake populations of the opossum shrimp (Mysis relicta) were red-shifted by 25 nm compared with all Baltic Sea populations. This is calculated to confer a significant advantage in both QC and SNR in many humus-rich lakes with reddish water. Since only A2 chromophore was present, the differences obviously reflect evolutionary tuning of the visual protein, the opsin. The changes have occurred within the ca. 9000 years that the lakes have been isolated from the Sea after the most recent glaciation. At present, it seems that the mechanism explaining the spectral differences between lake and sea populations is not an amino acid substitution at any other conventional tuning site, but the mechanism is yet to be found.

Food habits and dietary variability of pelagic nekton off Oregon and Washington, 1979-1984

The food habits of 20 species of pelagic nekton were investigated from collections made with small-mesh purse seines from 1979-84 off Washington and Oregon. Four species (spiny dogfish, Squalus acanthias; soupfin shark, Galeorhinus zyopterus; blue shark, Prionace glauca; and cutthroat trout, Salmo clarki) were mainly piscivorous. Six species (coho salmon, Oncorhynchus kisutch; chinook salmon, O. tshawytscha; black rockfish, Sebastes melanops; yellowtail rockfish, S. f1avidus; sablefish, Anoplopoma fimbria; and jack mackerel, Trachurus symmetricus) consumed both nektonic and planktonic organisms. The remaining species (market squid, Loligo opalescens; American shad, Alosa sapidissima; Pacific herring, Clupea harengus pallasi; northern anchovy, Engraulis mordax; pink salmon, O. gorbuscha; surf smelt, Hypomesus pretiosus; Pacific hake, Merluccius productus; Pacific saury, Cololabis saira; Pacific mackerel, Scomber japonicus; and medusafish, Icichthys lockingtom) were primarily planktonic feeders. There were substantial interannual, seasonal, and geographic variations in the diets of several species due primarily to changes in prey availability. Juvenile salmonids were not commonly consumed by this assemblage of fishes (PDF file contains 36 pages.)

Food habits of Atlantic white-sided dolphins (Lagenorhynchus acutus) off the coast of New England

Although the Atlantic white-sided dolphin (Lagenorhynchus acutus) is one of the most common dolphins off New England, little has been documented about its diet in the western North Atlantic Ocean. Current federal protection of marine mammals limits the supply of animals for investigation to those incidentally caught in the nets of commercial fishermen with observers aboard. Stomachs of 62 L. acutus were examined; of these 62 individuals, 28 of them were caught by net and 34 were animals stranded on Cape Cod. Most of the net-caught L. acutus were from the deeper waters of the Gulf of Maine. A single stomach was from the continental slope south of Georges Bank. At least twenty-six fish species and three cephalopod species were eaten. The predominant prey were silver hake (Merluccius bilinearis), spoonarm octopus (Bathypolypus bairdii), and haddock (Melanogrammus aeglefinus). The stomach from a net-caught L. acutus on the continental slope contained 7750 otoliths of the Madeira lanternfish (Ceratoscopelus maderensis). Sand lances (Ammodytes spp.) were the most abundant (541 otoliths) species in the stomachs of stranded L. acutus. Seasonal variation in diet was indicated; pelagic Atlantic herring (Clupea harengus) was the most important prey in summer, but was rare in winter. The average length of fish prey was approximately 200 mm, and the average mantle length of cephalopod prey was approximately 50 mm.

Pelagic behavior of adult Greenland halibut (Reinhardtius hippoglossoides)

It is evident from several field experiments with vertical longlines and archival tags, as well as concurrent studies of predator-prey relationships, that adult specimens of the deep-water flatfish Greenland halibut (Reinhardtius hippoglossoides) make regular excursions several hundred meters through the water column. The distribution of longline catches within the water column is confined to a well-defined depth layer overlapping with the distribution of blue whiting (Micromesistius poutassou), an important prey species, and depth recordings from archival tags overlap with Atlantic herring (Clupea harengus), the other major fish prey. The degree of pelagic use varies with fish size as well as seasons. Smaller individuals are found further off the bottom, and pelagic activity is greatest during early autumn. Interaction with pelagic prey species can influence results from bottom trawl surveys.

Decline in condition of northern bluefin tuna (Thunnus thynnus) in the Gulf of Maine

The northern bluefin tuna (Thunnus thynnus) is a highly mobile apex predator in the Gulf of Maine. Despite current stock assessments that indicate historically high abundance of its main prey, Atlantic herring (Clupea harengus), commercial fishermen have observed declines in the somatic condition of northern bluefin tuna during the last decade. We examined this claim by reviewing detailed logbooks of northern bluefin tuna condition from a local fishermen’s cooperative and applying multinomial regression, a robust tool for exploring how a categorical variable may be related to other variables of interest. The data set contained >3082 observations of condition (fat and oil content and fish shape) from fish landed between 1991 and 2004. Energy from stored lipids is used for migration and reproduction; therefore a reduction in energy acquisition on bluefin tuna feeding grounds could diminish allocations to growth and gamete production and have detrimental consequences for rebuilding the western Atlantic population. A decline in northern bluefin tuna somatic condition could indicate substantial changes in the bottom-up transfer of energy in the Gulf of Maine, shifts in their reproductive or migratory patterns, impacts of fishing pressure, or synergistic effects from multiple causes.