Northeast Arctic cod population persistence in the Lofoten-Barents Sea system under fishing.

Population growth, and hence the population's persistence, is affected by several factors such as climate, species interaction, and harvesting pressure. Proper resource management requires an understanding of these factors. We apply techniques based upon age-structured population matrices to analyze estimated stock sizes derived from annual bottom trawl sampling in the winter feeding area of northeast Arctic cod (Gadus morhua L.) from 1981 to 2003. We run generalized additive models to explain population growth rate by different explanatory variables. Cod population growth was found to be positively related to the abundance of capelin (Mallotus villosus Miller), negatively related to the number of cannibalistic cod with a two-year lag, and marginally positively related to the winter North Atlantic Oscillation index (NAO). This model remains true independently from the population status (i.e., fished or non-fished). Capelin abundance is the main variable that to some degree can be adjusted in order to maintain the population size at a given level of cod harvesting. Our results point to the importance of managing conjointly cod and capelin stocks.     

Spatial anatomy of species survival: effects of predation and climate-driven environmental variability

The majority of survival analyses focus on temporal scales. Consequently, there is a limited understanding of how species survival varies over space and, ultimately, how spatial variability in the environment affects the temporal dynamics of species abundance. Using data from the Barents Sea, we study the spatiotemporal variability of the juvenile Atlantic cod (Gadus morhua) survival. We develop an index of spatial survival based on changes of juvenile cod distribution through their first winter of life (from age-0 to age-1) and study its variability in relation to biotic and abiotic factors. Over the 25 years analyzed (1980-2004), we found that, once the effect of passive drift due to dominant currents is accounted for, the area where age-0 cod survival was lowest coincided with the area of highest abundance of older cod. Within this critical region, the survival of age-0 cod was negatively affected by its own abundance, by that of older cod, and by bottom depth. Furthermore, during cold years, age-0 cod survival increased in the eastern and coldest portion of the examined area, which was typically avoided by older conspecifics. Based on these results we propose that within the examined area top-down mechanisms and predation-driven density dependence can strongly affect the spatial pattern of age-0 cod survival. Climate-related variables can also influence the spatial survival of age-0 cod by affecting their distribution and that of their predators. Results from these and similar studies, focusing on the spatial variability of survival rates, can be used to characterize species habitat quality of marine renewable resources.     

Diet of 0-group stages of capelin (<Emphasis Type="Italic">Mallotus villosus</Emphasis>), herring (<Emphasis Type="Italic">Clupea harengus</Emphasis>) and cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) during spring and summer in the Barents Sea

Recruitment of capelin in the Barents Sea fail when juvenile herring and cod are abundant and the potential for feeding competition of wild sympatric capelin and herring larvae and small cod juveniles were investigated. The frequency of gut evacuation after capture of capelin larvae were also studied in mesocosms. Small capelin larvae (<35 mm length) fed on small prey including phytoplankton, invertebrate eggs and nauplii, bivalves, other invertebrate larvae and small copepods. Calanus copepodites were only observed in large capelin larvae (>26 mm length). Calanus copepodites were the major food sources for contemporary herring larvae (25–35 mm length) and Calanus and euphausiids were the major prey for small juvenile herring (37–60 mm length) and cod (18–40 mm length). Capelin larvae reared in mesocosms evacuated the guts shortly after capture. Capelin larvae had a smaller mouth and fed on smaller prey than herring and cod of the same length. This implies that the small capelin larvae, in contrast to sympatric small herring and cod, are not tightly linked to the food chain involving Calanus and euphausiids. Thus, exploitative competition between capelin larvae and planktivorous fish that rely on Calanus and euphausiids in the Barents Sea may be relaxed.     

Distribution of marine predator hotspots explained by persistent areas of prey

Investigations of distributional and density patterns of marine predators often reveal areas where high abundances of one or many species overlap in space and time (‘biological hotspots’); however, mechanisms underlying hotspot formation are often unclear, leading to difficulties determining spatial and temporal boundaries of protected areas. On the northeast Newfoundland coast, I previously described annually persistent aggregations of a key forage fish species, capelin (Mallotus villosus): (1) two pre- and post-spawning staging areas in deep (>150 m) bathymetric channels, (2) a cluster of four persistently used demersal spawning sites (17–40 m), and (3) a coastal migratory route (<50 m). Through at-sea surveys repeated over 8 years (2000–2003, 2007, and 2009–2011), I show that the majority of predator hotspots identified were spatially associated with (i.e., within 10 km) these persistent capelin areas for breeding seabirds (common murre: 85.2 ± 4.6 %; northern gannet: 66.9 ± 6.6 %), overwintering seabirds (great and sooty shearwaters: 88.0 ± 6.9 %), and baleen whales (humpback, minke, and fin whales: 86.8 ± 8.6 %). Most predator hotspots were closer to the spawning (3.8–14.0 km) relative to the staging areas (13.1–27.6 km), especially for murres and shearwaters. Interspecific differences were attributed to variation in maximum dive depths and dietary preferences. Predators only aggregated within the spawning area, while capelin were spawning, suggesting that interannual variation in association with predator and capelin hotspots was attributed to variation in survey timing relative to capelin spawning. As these areas of persistent capelin are bound by static bathymetric and large-scale oceanographic features and can be delimited in time based on the capelin spawning period, they may be important candidate areas for protection.     

Interactions between cod, herring and sprat in the changing environment of the Baltic Sea: A dynamic model analysis

The interactions between cod (Gadus morhua), herring (Clupea harengus) and sprat (Sprattus sprattus) in the Central Baltic Sea were examined with a simple dynamic model, an alternative to more complicated and data-demanding multispecies and ecosystem models. The main aims of the study were to compare the effect of alternative structures on the model output and examine the control relationships in the fish assemblage under different environmental conditions. The effect of environmental conditions was modelled using a stock-recruitment equation for cod incorporating an environmental index. The model output was especially sensitive to the functional response in predation by cod on herring and sprat. The type II functional response led to a collapse of the clupeid stocks when cod was abundant, while the type III response produced more realistic stock dynamics. According to the simulations, an abundant cod stock was able to keep the sprat stock at a low level, while the herring stock was less affected and benefited from the decreased density of sprat. Simulation of different fishing scenarios indicated that reducing fishing mortality to the level currently advised by ICES would allow the recovery of the cod stock even in unfavourable environmental conditions.     

Influence of increased cod abundance and temperature on recruitment of northern shrimp (Pandalus borealis)

Despite an increase in northern shrimp (Pandalus borealis) female biomass in the past years, the recruitment of the offshore population north and northeast of Iceland has remained very low. In this study, the influence of abiotic and biotic factors was studied in relation to shrimp recruitment. Two factors, cod (Gadus morhua) abundance and summer sea surface temperature (SST), were found to have a negative effect on offshore shrimp recruitment, explaining 71 % of the observed variation. Both cod abundance and temperature on the offshore shrimp grounds have increased in the past years, while recruitment has decreased and been at historically low levels since 2005. No significant relationship was found between recruitment and spawning biomass, indicating that recruitment variability is mainly driven by other factors. Cod abundance and summer SST are likely to affect different life stages of shrimp, as SST influences shrimp during its planktonic phase while cod abundance influences the demersal stage.     

Environmental variability and fledging body mass of Common Guillemot Uria aalge chicks

To gain a better understanding of population processes and in the light of the critically endangered status of the Common Guillemot Uria aalge in Norway, we investigate which environmental factors might affect the fitness of guillemot chicks as they depart from the nest site over a 16-year period on a colony in NE Norway. Although prey composition did not seem to influence the fledging body mass of the chicks, there were significant relationships between the yearly variations in chick body mass and abundance of two important prey species (1-group herring Clupea harengus that is an important chick food item and 0-group cod Gadus morhua that is an important adult food item), population size and the sea surface temperature around the colony. The positive influence of young herring and cod on Common Guillemot chick mass occurred during a period of warming in the Barents Sea such that future recruitment into the population will depend partly on the long-term changes in ocean climate in the region.     

Strength of evidence for density dependence in abundance time series of 1198 species

Population limitation is a fundamental tenet of ecology, but the relative roles of exogenous and endogenous mechanisms remain unquantified for most species. Here we used multi-model inference (MMI), a form of model averaging, based on information theory (Akaike's Information Criterion) to evaluate the relative strength of evidence for density-dependent and density-independent population dynamical models in long-term abundance time series of 1198 species. We also compared the MMI results to more classic methods for detecting density dependence: Neyman-Pearson hypothesis-testing and best-model selection using the Bayesian Information Criterion or cross-validation. Using MMI on our large database, we show that density dependence is a pervasive feature of population dynamics (median MMI support for density dependence = 74.7-92.2%), and that this holds across widely different taxa. The weight of evidence for density dependence varied among species but increased consistently, with the number of generations monitored. Best-model selection methods yielded similar results to MMI (a density-dependent model was favored in 66.2-93.9% of species time series), while the hypothesis-testing methods detected density dependence less frequently (32.6-49.8%). There were no obvious differences in the prevalence of density dependence across major taxonomic groups under any of the statistical methods used. These results underscore the value of using multiple modes of analysis to quantify the relative empirical support for a set of working hypotheses that encompass a range of realistic population dynamical behaviors.     

A simulation model to explore the relative value of stock enhancement versus harvest regulations for fishery sustainability

Harvest restrictions and stock enhancement are commonly proposed management responses for sustaining degraded fisheries, but comparisons of their relative effectiveness have seldom been considered prior to making policy choices. We built a population model that incorporated both size-dependent harvest restrictions and stock enhancement contributions to explore trade-offs between minimum length limits and stock enhancement for improving population sustainability and fishery metrics (e.g., catch). We used a Murray cod Maccullochella peelii peelii population as a test case, and the model incorporated density-dependent recruitment processes for both hatchery and wild fish. We estimated the spawning potential ratio (SPR) and fishery metrics (e.g., angler catch) across a range of minimum length limits and stocking rates. Model estimates showed that increased minimum length limits were much more effective than stock enhancement for increasing SPR and angler catches in exploited populations, but length limits resulted in reduced harvest. Stocking was predicted to significantly increase total recruitment, population sustainability, and fishery metrics only in systems where natural reproduction had been greatly reduced via habitat loss, fishing mortality was high, or both. If angler fishing effort increased with increased fish abundance from stocking efforts, fishing mortality was predicted to increase and reduce the benefits realized from stocking. The model also indicated that benefits from stock enhancement would be reduced if reproductive efficiency of hatchery-origin fish was compromised. The simulations indicated that stock enhancement was a less effective method to improve fishery sustainability than measures designed to reduce fishing mortality (e.g., length limits).     

Contrasting foraging tactics by northern gannets (<Emphasis Type="Italic">Sula bassana</Emphasis>) breeding in different oceanographic domains with different prey fields

In order to forage and to provision offspring effectively, seabirds negotiate a complex of behavioural, energetic, environmental and social constraints. In first tests of GPS loggers with seabirds in North America, we investigated the foraging tactics of free-ranging northern gannets (Sula bassana) at a large and a medium-sized colony that differed in oceanography, coastal position and prey fields. Gannets at Low Arctic colony (Funk Island) 50 km off the northeast coast of Newfoundland, Canada provisioned chicks almost entirely with small forage fish (capelin Mallotus villosus, 89%), while at boreal colony (Bonaventure Island) 3 km from shore in the Gulf of St. Lawrence, Quebec, Canada, large pelagic fish dominated parental prey loads (Atlantic mackerel Scomber scombrus 50%, Atlantic herring Clupea harengus 33%). Mean foraging range and the total distance travelled per foraging trip were significantly greater at the larger inshore colony (Bonaventure) than at the smaller offshore colony (Funk Island; 138 and 452 km vs. 64 and 196 km, respectively). Gannets from Funk Island consistently travelled inshore to forage on reproductive capelin shoals near the coast, whereas foraging flights of birds from Bonaventure were much more variable in direction and destination. Birds from the Low Arctic colony foraged in colder sea surface water than did birds from the boreal colony, and dive characteristics differed between colonies, which is concordent with the difference in prey base. Differences between the colonies reflect oceanographic and colony-size influences on prey fields that shape individual foraging tactics and in turn generate higher level colony-specific foraging “strategies”.     

Movements of farmed and wild Atlantic cod (Gadus morhua) released in Trinity Bay,Newfoundland

Fishery scientists and managers are investigating the feasibility of enhancing annual recruitment to the northern Atlantic cod (Gadus morhua L.) stock complex off Labrador and northeastern Newfoundland through the release of farmed fish back into the sea. Release of newly matured fish and adults with farm-advanced fecundity would increase the spawning biomass. Enhancement efforts might be measurably successful in major bays that are year-round habitats for cod. To determine if farmed cod would remain and spawn in Trinity Bay, 14 fish with surgically implanted transmitters were released in November 1992. Sonic tracking confirmed that farmed cod released on the western side of Trinity Bay overwintered within the bay, and integrated with wild cod approaching spawning condition in April 1993. Blood plasma antifreeze levels confirmed that these wild cod had overwintered inshore in subzero waters. A spawning aggregation was found in July 1993, providing evidence that northern cod reproduce in Newfoundland bays. These findings suggest that it may be possible to increase the number of cod spawning inshore through the release of farmed fish.     

Energy content and fecundity of capelin (<Emphasis Type="Italic">Mallotus villosus</Emphasis>) along a 1,500-km latitudinal gradient

Capelin is a planktivorous key fish species in many subarctic ecosystems representing the link between lower trophic levels and apex predators. Little is known, however, of capelin in Greenland waters, but it has been suggested that size and growth increases due to difference in food availability and temperature conditions along a 1,500 km south–north gradient on the west coast. It is presently unknown how the qualitative state of capelin energy content is affected along this gradient. Based on 2007 and 2008 samples, we show that energy content increases with capelin length in both spawning and non-spawning fish and that it varies with latitude in spawning fish along West Greenland (60–71°N). Combining our results on energy content with information on capelin growth along the same latitudinal climate gradient demonstrates that less and lower-quality food is available to capelin predators in the south than in the north.     

Ecological commonalities among pelagic fishes: comparison of freshwater ciscoes and marine herring and sprat

Systematic comparisons of the ecology between functionally similar fish species from freshwater and marine aquatic systems are surprisingly rare. Here, we discuss commonalities and differences in evolutionary history, population genetics, reproduction and life history, ecological interactions, behavioural ecology and physiological ecology of temperate and Arctic freshwater coregonids (vendace and ciscoes, Coregonus spp.) and marine clupeids (herring, Clupea harengus, and sprat, Sprattus sprattus). We further elucidate potential effects of climate warming on these groups of fish based on the ecological features of coregonids and clupeids documented in the previous parts of the review. These freshwater and marine fishes share a surprisingly high number of similarities. Both groups are relatively short-lived, pelagic planktivorous fishes. The genetic differentiation of local populations is weak and seems to be in part correlated to an astonishing variability of spawning times. The discrete thermal window of each species influences habitat use, diel vertical migrations and supposedly also life history variations. Complex life cycles and preference for cool or cold water make all species vulnerable to the effects of global warming. It is suggested that future research on the functional interdependence between spawning time, life history characteristics, thermal windows and genetic differentiation may profit from a systematic comparison of the patterns found in either coregonids or clupeids.     

Egg buoyancy of Baltic cod (Gadus morhua) and its implications for cod stock fluctuations in the Baltic

Successful spawning of Baltic cod (Gadus morhua L.) is restricted to the deep basins of the Baltic Sea and is dependent on abiotic conditions such as salinity and water oxygen content. Due to irregular inflows of saline water from the North Sea, the conditions for spawning fluctuate and consequently so does the cod stock. In May and June 1990 eggs were obtained from spawning cod caught off northern Gotland, Sweden. Our investigation revealed neutral egg buoyancy for Baltic cod at a salinity of 14.4±1.1 ppt, with a slight decrease some days before hatching. Today salinities of this magnitude occur only in the Bornholm basin, whereas in the Gdansk and Gotland basins the eggs sink toward the bottom and are exposed to lethal oxygen conditions.     

Effects of salinity on spermatozoa motility,percentage of fertilized eggs and egg development of Baltic cod (Gadus morhua), and implications for cod stock fluctuations in the Baltic

Variation in cod stock strength in the Baltic Sea is considered to be dependent on abiotic conditions such as salinity and water oxygen content in the spawning areas (the Baltic deep basins). Spawning cod were caught off northern Gotland, Sweden, from April to June in 1989 and 1990. Our investigation revealed a reduction in spermatozoan motility and a subsequent reduction in the percentage of fertilized eggs in salinities of 10 to 12. Normal egg development required a minimum salinity of 11. This coincides with the osmolality of the seminal plasma and egg yolk; i.e., the Baltic cod is adapted to hyperosmotic conditions for spawning and is thus totally dependent on periodical inflows of saline water from the North Sea.     

Correlation between the seasonal distribution of harbour porpoises and their prey in the Sound,Baltic Sea

Low densities of harbour porpoises in winter (November–March) and high densities in summer (April–October) were found in the Sound, connecting the Baltic Sea and Kattegat. Due to their high energy requirements, it is hypothesized that the density of harbour porpoises is related to local prey abundance. This was tested by examining the stomach content of 53 harbour porpoises collected between 1987 and 2010 in the Sound (high season, 34 porpoises; low season, 19 porpoises). A total of 1,442 individual fish specimens from thirteen species were identified. Twelve of these were present in the high–porpoise density season and seven in the low-density season. The distribution of occurrence and the distribution of number of fish species were different between seasons, indicating a shift in prey intake between seasons. Furthermore, during the high-density season, the mean and total prey weight per stomach as well as the prey species diversity was higher. However, no difference was found in the number of prey species between the two seasons, indicating a higher quality of prey in the high-density season. Atlantic cod was found to be the main prey species in terms of weight in the high-density season while Atlantic herring and Atlantic cod were equally important during the low-density season. Prey availability and predictability are suggested as the main drivers for harbour porpoise distribution, and this could be caused by the formation of frontal zones in spring in the northern part of the Sound, leading to prey concentrations in predictable areas.     

Neighborhood and community interactions determine the spatial pattern of tropical tree seedling survival

Factors affecting survival and recruitment of 3531 individually mapped seedlings of Myristicaceae were examined over three years in a highly diverse neotropical rain forest, at spatial scales of 1-9 m and 25 ha. We found convincing evidence of a community compensatory trend (CCT) in seedling survival (i.e., more abundant species had higher seedling mortality at the 25-ha scale), which suggests that density-dependent mortality may contribute to the spatial dynamics of seedling recruitment. Unlike previous studies, we demonstrate that the CCT was not caused by differences in microhabitat preferences or life history strategy among the study species. In local neighborhood analyses, the spatial autocorrelation of seedling survival was important at small spatial scales (1-5 m) but decayed rapidly with increasing distance. Relative seedling height had the greatest effect on seedling survival. Conspecific seedling density had a more negative effect on survival than heterospecific seedling density and was stronger and extended farther in rare species than in common species. Taken together, the CCT and neighborhood analyses suggest that seedling mortality is coupled more strongly to the landscape-scale abundance of conspecific large trees in common species and the local density of conspecific seedlings in rare species. We conclude that negative density dependence could promote species coexistence in this rain forest community but that the scale dependence of interactions differs between rare and common species.     

Concentrations of iron-55 in commercial fish species from the North Atlantic

Concentrations of the weapon-test, fallout radionuclide iron-55 have been reported for migratory species, such as the Pacific salmon, which suggest a marked decrease in concentration with decrease in latitude of the point of capture. This situation has been examined for North Atlantic cod stocks in order to provide data for a species representative of a large geographical area, but divided into more or less distinct stocks each with a restricted territory, in order to provide further data on the importance of latitude in determining iron-55 concentrations. The data obtained show that there is a marked dependence upon latitude, Arctic cod averaging 90 pCi ⁵⁵Fe/mg Fe compared with 15 pCi ⁵⁵Fe/mg Fe for cod from middle latitudes. The mechanisms that could lead to such distribution are examined, and it is concluded that deposition of this radionuclide was exceptionally high over the sea in northern latitudes compared with its deposition on land, and that this pattern of deposition, taken together with the very high biological availability of the material at the time of deposition, has led to the very high specific activities found in northern latitudes and recorded for Arctic cod stocks in particular.     

Tolerance of soil pathogens co-varies with shade tolerance across species of tropical tree seedlings

A negative feedback between local abundance and natural enemies could contribute to maintaining tree species diversity by constraining population growth of common species. Soil pathogens could be an important mechanism of such noncompetitive distance and density-dependent (NCDD) mortality, but susceptibility to local pathogens may be ameliorated by a life history strategy that favors survivorship. In a shade-house experiment (1% full sun), we tested seedling life span, growth, and mass allocation responses to microbial extract filtered from conspecific-cultured soil in 21 tree species that varied in abundance and shade tolerance in a wet tropical forest (La Selva Biological Station, Costa Rica). Forty-three percent of the species had significant reductions, and 10% of the species had significant increases in life span, growth, root length, or root surface area when inoculated with microbial extract; 10% of the species experienced opposing reductions and increases in these characteristics. Contrary to expectation, species' local abundance was not related to species-specific responses to microbial extracts from cultured soils. Across species, seedling shade tolerance (survival at 1% full sun) was negatively correlated with susceptibility to the microbial. treatment for both survival and total mass accumulation, thereby exaggerating shade tolerance differences among species. Thus, soil pathogens may contribute to species coexistence through heightening niche differentiation rather than through negative density dependence in common species.     

Widespread density-dependent seedling mortality promotes species coexistence in a highly diverse Amazonian rain forest

Negative density-dependent mortality can promote species coexistence through a spacing mechanism that prevents species from becoming too locally abundant. Negative density-dependent seedling mortality can be caused by interactions among seedlings or between seedlings and neighboring adults if the density of neighbors affects the strength of competition or facilitates the attack of natural enemies. We investigated the effects of seedling and adult neighborhoods on the survival of newly recruited seedlings for multiple cohorts of known age from 163 species in Yasuni National Park, Ecuador, an ever-wet, hyper-diverse lowland Amazonian rain forest. At local scales, we found a strong negative impact on first-year survival of conspecific seedling densities and adult abundance in multiple neighborhood sizes and a beneficial effect of a local tree neighborhood that is distantly related to the focal seedling. Once seedlings have survived their first year, they also benefit from a more phylogenetically dispersed seedling neighborhood. Across species, we did not find evidence that rare species have an advantage relative to more common species, or a community compensatory trend. These results suggest that the local biotic neighborhood is a strong influence on early seedling survival for species that range widely in their abundance and life history. These patterns in seedling survival demonstrate the role of density-dependent seedling dynamics in promoting and maintaining diversity in understory seedling assemblages. The assemblage-wide impacts of species abundance distributions may multiply with repeated cycles of recruitment and density-dependent seedling mortality and impact forest diversity or the abundance of individual species over longer time scales.