Genetic divergence at the synaptophysin (Syp I) locus among Norwegian coastal and north-east Arctic populations of Atlantic cod

Several nuclear RFLP loci have been discovered recently that exhibit extensive allele frequency variation among Norwegian coastal and north-east Arctic populations of Atlantic cod Gadus morhua. One of these polymorphisms was detected by hybridizing an anonymous cDNA clone (GM798) against genomic DNA digested with the restriction enzyme DraI. This cDNA clone has now been sequenced and identified as synaptophysin (Syp I), an integral synaptic vesicle membrane protein. Primers were constructed that amplify an intron of the Syp I gene that is polymorphic for the DraI site, thus making it possible to use a PCR-based assay to score the polymorphism. A total of 965 individuals sampled from the Barents Sea, coastal areas and fjords in northern Norway have been analysed for this polymorphism. The results confirm that highly significant differences exist between NE Arctic and coastal cod at the Syp I locus. A cluster analysis revealed a deep split between coastal and Arctic populations and hierarchical F-statistics indicated that about 40% of the total variation was attributable to differences between Arctic and coastal groups. The temporal stability of allele frequencies was assessed by comparing Syp I allele frequencies among samples of juveniles (0 group) captured at specific locations in fjords in consecutive years and among samples of adults and juveniles collected from the same fjord. Samples of juveniles collected in 1994 and 1995 in Malangen were genetically indistinguishable whereas juveniles sampled from Dønnesfjord and Ullsfjorden over the same 2-year period exhibited significant differences. Adults and 0-group individuals collected from the same fjord were found to be genetically indistinguishable in Malangen, but not in Balsfjorden. In addition to detecting large differences among Arctic and coastal groups, the Syp I locus suggests that genetic heterogeneity exists among resident populations of cod in different fjords and that gene flow among populations throughout northern Norway may be considerably lower than previously believed.     

Transport of North Sea cod larvae into the Skagerrak coastal populations

The Atlantic cod (Gadus morhua) is economically one of the world's most important marine species--a species presently suffering from heavy overexploitation throughout its range of distribution. Although not fully understood, the Atlantic cod is believed to be structured into populations in a rather complex manner, whereby both highly migratory and more confined ocean-spawning stocks coexist with stationary coastal populations. Owing to the complex population structure, little is presently known about how overexploitation of offshore stocks may affect other segments of the species. Here, we use microsatellite DNA analyses of coastal and offshore cod in combination with oceanographic modelling to investigate the population structure of Atlantic cod in the North Sea-Skagerrak area and evaluate the potential for larval transport into coastal populations. Our results suggest an extensive but temporally variable drift of offshore cod larvae into coastal populations. In a year (2001) with high inflow of North Sea waters into the Skagerrak we find that juvenile cod caught along the Skagerrak coast are predominantly of North Sea origin, whereas in a year (2000) with low inflow juveniles appear to be of local origin. These findings indicate that offshore cod may influence coastal cod populations over large distances.     

Pantophysin (Pan I) locus divergence between inshore v. offshore and northern v. southern populations of Atlantic cod in the north‐east Atlantic

More than 6000 cod Gadus morhua , sampled in coastal and offshore waters stretching from the Barents Sea down to the North Sea, were analysed for frequencies of alleles at the scnDNA pantophysin locus ( Pan I)[formerly called synaptophysin ( Syp I)]. The significant allele frequency difference between the two major stocks of cod in Norway, north‐east Arctic cod (NEAC) and Norwegian coastal cod (NCC), was upheld in all years of the investigation (1993 to 2001), and applied both to larval cod and post‐juveniles of various ages. On a north‐south axis, the appearance of a latitudinal cline of post‐juvenile (≥1 year) allele frequencies was exposed. The intermediate allele frequencies in coastal areas of northern Norway, seem to a large extent to be caused by intermingling of the two stocks, although the existence of populations of coastal cod with alternative Pan I frequencies could not be ruled out. The role of selection is yet unresolved. Depth of the sampling location seemed to have an effect on the allele frequencies and their temporal stability, while there was no indication of seasonal variation in the frequencies. Breeding structure was the most likely cause for upholding the extreme divergence in Pan I frequencies between NEAC and NCC.     

Morphometric Evidence for Three Juvenile Stages in Some Species of Xiphinema americanum sensu lato

One to two hundred nematodes from each of seven Xiphinema americanum-group populations were measured to determine the range of stylet and body lengths for juveniles and adults. First-stage juveniles were identified by the position of the replacement odontostyle (i.e., the tip of the replacement odontostyle overlapped the base of the odontophore). Nematodes were identified as second stage if the functional odontostyle was the same length as the replacement odontostyle of the first stage. Subsequent stages were similarly identified by establishing the range of corresponding replacement and functional odontostyle lengths. In all populations examined, this procedure created natural divisions that clearly grouped nematodes by stylet and body length. Presumably these groups identified all juvenile and adult stages. Populations of X. americanum, X. rivesi, and X. californicum from the United States had three juvenile stages, but a population of X. pachtaicum from Bulgaria had four juvenile stages.     

Ecological and genetic impact of Atlantic cod larval drift in the Skagerrak

We evaluate the hypothesis that Atlantic cod larvae are passively transported by sea currents from off-shore spawning areas to settle in coastal waters, a hypothesis which has recently gained support from genetic analysis of cod in the North Sea-Skagerrak area. Such larval transport has been suggested to be an important mechanism behind the commonly observed low spatial genetic differentiation in many marine organisms. Here, we apply an ARMAX(2,2) model for juvenile abundance and use long-term monitoring data from the Skagerrak coast, constituting 54 continuous annual series from 1945 to 1997. Analysing the model, we find that the product of the size of the North Sea breeding stock and the strength of the net inflow of North Sea waters had a significant, positive effect on the abundance of coastal juvenile cod. The peak effect occurs during the month of March, just after spawning, when eggs and larvae remain pelagic and sensitive to currents. In contrast, we find no evidence of any direct effect of the North Sea spawning stock alone. Our analyses indicate that 15-20,000 0-group larvae from the North Sea reach each fjord per year, on average. This corresponds to about 1-10% of the total 0-group population in each fjord on average. These findings clearly demonstrate a direct link between larval drift and gene flow in the marine environment.     

Long-term studies on genetic interaction between wild and ranched cod Gadus morhua by use of a genetic marked strain

Releases of farmed fish, whether accidental from commercial aquaculture facilities or intentional as part of stock enhancement/ranching activities, are considered to pose a risk to native gene pools. Stock enhancement studies of Atlantic cod, Gadus morhua , based on artificially produced juveniles, were initiated in 1984 in western Norway, and genetic aspects were incorporated. In order to investigate potential interbreeding between released and wild cod, a genetically marked cod strain was developed, being homozygotic for a rare allele ( GPI‐1*30 ) expressed in white muscle tissue. In the period from 1990 to 1994, juveniles from the genetic marked strain were released in large quantities in three locations (Masfjord, Øygarden, Heimarkspollen), giving a significant increase of the marker allele in the local wild cod populations. Recently, studies have been conducted in the same areas to estimate the extent of interbreeding between the wild and released cod. The results, however, revealed no permanent increase of the frequency of the marker allele and/or GPI‐1*30 heterozygotes as would be expected from interbreeding. The recent data are compared with comprehensive genetic data of the cod populations in the areas before the actual releases, covering the full period from 1994 to 2003. The present results are also discussed in relation to fishing pressure on coastal cod, migration information and reproductive success of released, genetically marked cod.     

Distribution and diet of 0-group cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) and haddock (<Emphasis Type="Italic">Melanogrammus aeglefinus</Emphasis>) in the Barents Sea in relation to food availability and temperature

Distribution of 0-group cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) in August–September 2005 and 2006 was mainly restricted to the Atlantic waters of the western and central areas of the Barents Sea. The main distribution of 0-group fish overlapped largely with areas of high biomass (>7 gm⁻² dry weight) of zooplankton. The copepod Calanus finmarchicus and krill Thysanoessa inermis, which are dominant zooplankton species in both Atlantic and boreal waters of the Barents Sea, were the main prey of 0-group cod and haddock. The main distribution, feeding areas and prey of 0-group cod and haddock overlapped, implying that competition for food may occur between the two species. However, though their diet coincided to a certain degree, haddock seems to prefer smaller and less mobile prey, such as Limacina and appendicularians. As 0-group fish increased in size, there seems to be a shift in diet, from small copepods and towards larger prey such as krill and fish. Overall, a largely pelagic feeding behaviour of 0-group cod and haddock was evident from this study.     

Allele frequency covariance among cohorts and its use in estimating effective size of age-structured populations

A general expression for the covariance of allele frequencies among cohorts in age-structured populations is derived. The expression is used to extend the so-called temporal method for estimating effective population size from allele frequency shifts among samples from cohorts born any number of years apart. Computer simulations are used to check on the accuracy and precision of the method, and an application to coastal Atlantic cod is presented.     

Migration patterns and recapture rates of North-east Arctic and Norwegian coastal cod reared and released under similar conditions

A total of 5000 1-group cod of Northeast Arctic and Norwegian coastal origin were reared under similar conditions, anchor tagged and released at two sites in western Norway. There were significantly more recaptures from the coastal cod. No differences in migration patterns were detected, suggesting that mortality rates were higher in North-east Arctic cod.     

Stochastic juvenile-adult models with application to a green tree frog population

We derive several stochastic models from a deterministic population model that describes the dynamics of age-structured juveniles coupled with size-structured adults. Numerical simulation results of the stochastic models are compared with the solution of the deterministic model. These models are then used to understand the effect of demographic stochasticity on the dynamics of an urban green tree frog (Hyla cinerea) population.     

Structural and functional connectivity of marine fishes within a semi-enclosed Newfoundland fjord

The interplay between structural connectivity ( i.e. habitat continuity) and functional connectivity ( i.e. dispersal probability) in marine fishes was examined in a coastal fjord (Holyrood Pond, Newfoundland, Canada) that is completely isolated from the North Atlantic Ocean for most of the year. Genetic differentiation was described in three species (rainbow smelt Osmerus mordax , white hake Urophycis tenuis and Atlantic cod Gadus morhua ) with contrasting life histories using seven to 10 microsatellite loci and a protein-coding locus, Pan I ( G. morhua ). Analysis of microsatellite differentiation indicated clear genetic differences between the fjord and coastal regions; however, the magnitude of difference was no more elevated than adjacent bays and was not enhanced by the fjord's isolation. Osmerus mordax was characterized by the highest structure overall with moderate differentiation between the fjord and St Mary's Bay ( F _ST c. 0·047). In contrast, U. tenuis and G. morhua displayed weak differentiation ( F _ST < 0·01). Nonetheless, these populations did demonstrate high rates (< 75%) of Bayesian self-assignment. Furthermore, elevated differentiation was observed at the Pan I locus in G. morhua between the fjord and other coastal locations. Interestingly, locus-specific genetic differentiation and expected heterozygosity were negatively associated in O. mordax , in contrast to the positive associations observed in U. tenuis and G. morhua . Gene flow in these species is apparently unencumbered by limited structural connectivity, yet the observed differentiation suggests that population structuring exists over small scales despite high dispersal potential.     

Derivation of stochastic partial differential equations for size- and age-structured populations

Stochastic partial differential equations (SPDEs) for size-structured and age- and size-structured populations are derived from basic principles, i.e. from the changes that occur in a small time interval. Discrete stochastic models of size-structured and age-structured populations are constructed, carefully taking into account the inherent randomness in births, deaths, and size changes. As the time interval decreases, the discrete stochastic models lead to systems of It? stochastic differential equations. As the size and age intervals decrease, SPDEs are derived for size-structured and age- and size-structured populations. Comparisons between numerical solutions of the SPDEs and independently formulated Monte Carlo calculations support the accuracy of the derivations.     

Evidence of sustained populations of a small reef fish on artificial structures. Does depth affect production on artificial reefs?

The length frequencies and age structures of resident Pseudanthias rubrizonatus (n = 407), a small protogynous serranid, were measured at four isolated artificial structures on the continental shelf of north-western Australia between June and August 2008, to determine whether these structures supported full (complete size and age-structured) populations of this species. The artificial structures were located in depths between 82 and 135 m, and growth rates of juveniles and adults, and body condition of adults, were compared among structures to determine the effect of depth on potential production. All life-history stages, including recently settled juveniles, females and terminal males, of P. rubrizonatus were caught, ranging in standard length (L(s)) from 16·9 to 96·5 mm. Presumed ages estimated from whole and sectioned otoliths ranged between 22 days and 5 years, and parameter ±s.e. estimates of the von Bertalanffy growth model were L(∞) = 152 ± 34 mm, k = 0·15(±0·05) and t(0) = -1·15(±0·15). Estimated annual growth rates were similar between shallow and deep artificial structures; however, otolith lengths and recent growth of juveniles differed among individual structures, irrespective of depth. The artificial structures therefore sustained full populations of P. rubrizonatus, from recently settled juveniles through to adults; however, confirmation of the maximum age attainable for the species is required from natural populations. Depth placement of artificial reefs may not affect the production of fish species with naturally wide depth ranges.     

Spatial and temporal analysis of juvenile blacktip reef shark (Carcharhinus melanopterus) demographies identifies critical habitats

Reef shark species have undergone sharp declines in recent decades, as they inhabit coastal areas, making them an easy target in fisheries (i.e., sharks are exploited globally for their fins, meat, and liver oil) and exposing them to other threats (e.g., being part of by-catch, pollution, and climate change). Reef sharks play a critical role in coral reef ecosystems, where they control populations of smaller predators and herbivorous fishes either directly via predation or indirectly via behavior, thus protecting biodiversity and preventing potential overgrazing of corals. The urgent need to conserve reef shark populations necessitates a multifaceted approach to policy at local, federal, and global levels. However, monitoring programmes to evaluate the efficiency of such policies are lacking due to the difficulty in repeatedly sampling free-ranging, wild shark populations. Over nine consecutive years, we monitored juveniles of the blacktip reef shark (Carcharhinus melanopterus) population around Moorea, French Polynesia, and within the largest shark sanctuary globally, to date. We investigated the roles of spatial (i.e., sampling sites) and temporal variables (i.e., sampling year, season, and month), water temperature, and interspecific competition on shark density across 10 coastal nursery areas. Juvenile C. melanopterus density was found to be stable over 9 years, which may highlight the effectiveness of local and likely federal policies. Two of the 10 nursery areas exhibited higher juvenile shark densities over time, which may have been related to changes in female reproductive behavior or changes in habitat type and resources. Water temperatures did not affect juvenile shark density over time as extreme temperatures proven lethal (i.e., 33°C) in juvenile C. melanopterus might have been tempered by daily variation. The proven efficiency of time-series datasets for reef sharks to identify critical habitats (having the highest juvenile shark densities over time) should be extended to other populations to significantly contribute to the conservation of reef shark species.     

Temporal and ontogenetic shifts in habitat use of juvenile Pacific cod (Gadus macrocephalus)

Habitat use of age-0 and age-1 juvenile Pacific cod (Gadus macrocephalus) was examined in coastal regions in Kodiak Alaska over daily, seasonal and annual scales. Catch data indicated highly variable recruitment to nursery areas, but a strong separation of distribution by depth among age groups. Age-0 cod were most abundant in the shallows (<3 m) whereas age-1 cod were typically found in depths (9.0-13.5 m). In comparison, age-1 saffron cod (Eleginus gracilis), another highly abundant gadid in the region, were found in shallower depths where age-0 cod often resided. Age-1 cod Pacific cod made diel lateral movements, moving into shallow regions at night where they co-occurred with age-0 cod to a greater extent. Laboratory light-gradient experiments indicated that age-0 cod tolerated intense lighting (~ 20-80 µE m^− 2 s^− 1) typical of shallow water regions whereas larger age-1 Pacific cod strongly avoid bright light given the choice. However, while diet data indicate age-1 cod of both species are moderately piscivorous (3% saffron cod; 16% Pacific cod), we found no direct evidence of predation on smaller conspecific cod, possibly due to the low densities of age-0 cod in the year of the diet study. Together, these data suggest that coastal regions continue to serve a nursery function beyond the 1st year of development for juvenile Pacific gadids, and that small-scale temporal and depth partitioning in these regions is a mechanism by which varying cod species and age classes co-occur.     

Influence of cruising and ambush predators on 3-dimensional habitat use in age 0 juvenile Atlantic cod Gadus morhua

Predators and prey often co-exist at high densities within the same habitat, yet the behavioural and spatial dynamics underlying this co-existence are not well known. To better understand small-scale, predator-prey co-occurrence, the spatial patterns and behaviour of age 0 juvenile cod Gadus morhua 75-88 mm SL and two of their known predators, age 2+ cod and short-horn sculpin Myoxocephalus scorpinus, were examined in two habitats (i.e., sand and eelgrass) using three-dimensional video analysis. Both habitat and predator type interacted to result in unique spatial patterns of prey. Spatial overlap between predators and prey was highest in open habitat in the presence of the cruising predator but lowest in the presence of sculpin in the same habitat. In eelgrass, age 0 cod avoided predators primarily along the vertical axis (i.e., distance off bottom). Age 0 cod stayed above eelgrass in the presence of sculpin but lowered themselves into the eelgrass while in the presence of predator cod. Anti-predator behaviour (i.e., predator-prey distance, prey cohesion and freezing) was significantly reduced over eelgrass compared to sand, suggesting eelgrass has lower ‘inherent risk’ than open habitats. However, predator consumption was similar across all treatments, suggesting that, 1) complex habitat also impairs the visual cues needed for anti-predator behaviour (e.g., schooling) and assessing the location of predators, and 2) predators change their behaviour with habitat to enhance their opportunities for finding and capturing prey.     

Stage dependent habitat use under conflicting predation pressure: An experimental test with larval and juvenile two-spotted gobies, Gobiusculus flavescens Fabricius

The effect of size, predator types and presence of multiple predators on the microhabitat use of larvae and juveniles of a sublittoral, semipelagically schooling fish, the two-spotted goby (Gobiusculus flavescens), was tested in two experiments. Larvae (15 and 25 days old, Experiment I) and juveniles (mean ± 1 S.E.: small, 15.9 mm ± 1.28; medium, 19.2 mm ± 1.43; and large, 23.4 mm ± 1.67, Experiment II) were allowed to choose between two sections of the tanks; an upper, representing a water column habitat, and a lower, artificially vegetated, representing the hyperbenthic habitat. Position of larvae or juveniles and the activity level of juveniles were recorded. Predator treatments were: (I) no predators (control), (II) a pelagic predator, the jellyfish Aurelia aurita L., (III) a hyperbenthic predator, the mysid Praunus flexuosus O.F. Müller or (VI) both predator types simultaneously. In Experiment I predators were restricted to the habitat which they were chosen to represent, while goby larvae could move freely. In Experiment II both predators and juvenile gobies were allowed to move freely between compartments.Increasing age caused larval gobies, but not juveniles to shift downwards. Only 25-day-old larvae and small juveniles avoided the mysid by shifting upwards. Larval response to A. aurita was also size dependant: 25-day-old larvae avoided medusae by shifting downwards, while 15 day olds did not. Emergent multiple predator effects were found for the vertical distribution of 15-day-old larvae and small juveniles. Larger juveniles were more active than smaller, both in the upper and the lower sections of tanks. P. flexuosus caused juvenile gobies in their vicinity (i.e. in the lower section) to increase their activity level, while small juveniles (but not medium-sized or large) increased their activity level even when further away (i.e. in the upper section). The presence of A. aurita led to a reduction in activity of small juveniles in its vicinity (i.e. in the upper section), while no response was observed among older juveniles or juveniles further away from the predator (i.e. in the lower section). Emergent multiple predator effects on the activity level of juveniles were not observed.     

Derivation of stochastic partial differential equations for size- and age-structured populations

Stochastic partial differential equations (SPDEs) for size-structured and age- and size-structured populations are derived from basic principles, i.e. from the changes that occur in a small time interval. Discrete stochastic models of size-structured and age-structured populations are constructed, carefully taking into account the inherent randomness in births, deaths, and size changes. As the time interval decreases, the discrete stochastic models lead to systems of Itô stochastic differential equations. As the size and age intervals decrease, SPDEs are derived for size-structured and age- and size-structured populations. Comparisons between numerical solutions of the SPDEs and independently formulated Monte Carlo calculations support the accuracy of the derivations.     

Variation in the quality and quantity of flavonoids in the leaves of coastal and inland Campanula punctata

Ultraviolet-B radiation is known as a noxious factor that destroys every life form. Plants that occupy the coastal area are assumed to be adaptive to UV-B as well as the other major stresses. The objective of this study was to clarify and compare (1) the flavonoid compositions of coastal and inland populations of Campanula punctata, and (2) UV-B effects on flavonoid content in plants originating from different habitats under uniform (i.e. experimental) and natural conditions. Flavonoid compositions of coastal and inland populations were shown to be identical. The UV-B exclusion experiment revealed two tendencies that were commonly observed in both coastal and inland populations: (1) the flavonoid accumulation decreased with an increase in degree of UV-B exclusion, and (2) the quantity of phenolic acids conversely became maximum under complete UV-B exclusion. Under the natural habitat conditions, significantly high accumulation of flavonoids were detected in two coastal populations even though no significant difference was found between the two other coastal populations and two inland populations. Weak correlations between UV-B intensity and flavonoid accumulation under the natural habitat conditions suggest that various micro-environmental factors may influence the production of flavonoids, and that the plants may acquire adaptive traits other than increasing flavonoids in order to inhabit the coastal environment.     

Abundance,structure, growth and origin of inshore clupeid populations of the west coast of Scotland

The abundance, structure, growth, and origin of clupeid populations in inshore waters of the west coast of Scotland were studied from April 1970 to October 1972. Clupeid populations in the area comprise mainly young fish. 0-group autumn-spawned herring, probably of Minch origin, move into the area about April and spring-spawned ones (Clyde origin) about June. The timing and the body length at which each group arrives in the area during the different years is the same. After the initial immigration, the distribution of both 0-group clupeids becomes localized.Herring populations in the sea-lochs and associated inshore waters are mainly 0-group fish, which are replaced each year by a new incoming brood. The sprat populations of the sea-lochs are dominated by the 0-group; in the more ‘open’ areas the populations comprise older individuals. Year-class distribution in the ‘open’ areas resemble that of the commercial fishery.The rate of increase of 0-group autumn-spawned herring is 3.68 mm/week and that of spring-spawned 0-group herring is 2.83 mm/week. The curves of growth of 0-group sprats of the 1970 and 1971 year-classes are different; the rate of increase in length, however, averages 3.55 mm/week for both year-classes and the differences are not significant.In sprats, after their first year of life a rapid increase in length takes place in the spring. This increase is thought to enable the majority of the population to reach the minimum size (88–90 mm) for initial gonadal maturation and thereby make them capable of reproducing in their second year of life.