Basin-scale phenology and effects of climate variability on global timing of initial seaward migration of Atlantic salmon (Salmo salar).

Migrations between different habitats are key events in the lives of many organisms. Such movements involve annually recurring travel over long distances usually triggered by seasonal changes in the environment. Often, the migration is associated with travel to or from reproduction areas to regions of growth. Young anadromous Atlantic salmon (Salmo salar) emigrate from freshwater nursery areas during spring and early summer to feed and grow in the North Atlantic Ocean. The transition from the freshwater ('parr') stage to the migratory stage where they descend streams and enter salt water ('smolt') is characterized by morphological, physiological and behavioural changes where the timing of this parr-smolt transition is cued by photoperiod and water temperature. Environmental conditions in the freshwater habitat control the downstream migration and contribute to within- and among-river variation in migratory timing. Moreover, the timing of the freshwater emigration has likely evolved to meet environmental conditions in the ocean as these affect growth and survival of the post-smolts. Using generalized additive mixed-effects modelling, we analysed spatio-temporal variations in the dates of downstream smolt migration in 67 rivers throughout the North Atlantic during the last five decades and found that migrations were earlier in populations in the east than the west. After accounting for this spatial effect, the initiation of the downstream migration among rivers was positively associated with freshwater temperatures, up to about 10 °C and levelling off at higher values, and with sea-surface temperatures. Earlier migration occurred when river discharge levels were low but increasing. On average, the initiation of the smolt seaward migration has occurred 2.5 days earlier per decade throughout the basin of the North Atlantic. This shift in phenology matches changes in air, river, and ocean temperatures, suggesting that Atlantic salmon emigration is responding to the current global climate changes.

From spawner habitat selection to stock-recruitment: Implications for assessment.

The relationship between the spawning stock size and subsequent number of recruits is a central concept in fisheries ecology. The influence of habitat selection of spawning individuals on the stock-recruitment relationship is poorly known. Here we explore how each of four different spawner behaviors might influence the stock-recruitment relationship and estimates of its parameters in the two most commonly used stock-recruitment functions (Beverton-Holt and Ricker). Using simulated stock-recruitment data generated by four different spawner behaviors applied to multiple discrete habitats, we show that when spawners were distributed proportionally to local carrying capacities, there was small or no bias in estimated recruitment and stock-recruitment parameters. For an ideal free distribution of spawners, larger bias in the estimates of recruitment and stock-recruitment parameters was obtained, whereas a random and a stepwise spawner behavior introduced the largest bias. Using stock-recruitment data corresponding to a "realistic" range of population densities and adding measurement error (20%-60%) to the simulated stock-recruitment data generated larger variation in the estimation bias than what was introduced by the spawner behavior. Thus, for exploited stocks at low population density and where spawning stock size and recruitment cannot be observed perfectly, partial observation of the possible spawner abundance range and measurement error might be of higher concern for management.

Bioturbation-driven release of buried PCBs and PBDEs from different depths in contaminated sediments.

Bioturbation can remobilize previously buried contaminants, leading to an increased exposure of aquatic biota. The remobilization of buried polychlorinated biphenyls (PCBs) and polybrominated diphenyl ethers (PBDEs) from three different sediment depth layers (2.0-2.5 cm, 5.0-5.5 cm, and 10.0-10.5 cm) was studied in a laboratory experiment with two benthic macrofauna species, the amphipod Monoporeia affinis and the polychaete Marenzelleria spp. Remobilization of PCBs and PBDEs was significantly higher in the presence of Marenzelleria spp. than in M. affinis treatments and controls (without macrofauna). The highest remobilization occurred from the most shallow layers (2.0-2.5 cm > 5.0-5.5 cm > 10.0-10.5 cm), but contaminants were remobilized due to bioturbation from layers down to at least 10 cm. Congeners with lower hydrophobicity were remobilized to a higher extent than more hydrophobic congeners. The contaminant distribution between the particulate and the dissolved phase in the water column depended on hydrophobicity and burial depth of the contaminant, with congeners from deeper layers displaying an increased distribution to the particulate phase. Release fluxes and sediment-to-water mass transfer coefficients (MTCs) show that bioturbation by the polychaete Marenzelleria spp. can lead to a significant remobilization of buried contaminants from Baltic Sea sediments.

Scaling up the functional response for spatially heterogeneous systems.

Scale transition theory is a framework for predicting regional population dynamics from local process functions and estimates of spatial heterogeneity. Using this framework, we estimated regional scale functional responses for a benthic predator-prey system in the Baltic Sea. Functional responses were based on laboratory experiments or field observations of stomach contents, and prey densities measured at a local scale (0.1 m(2)) or a regional scale (300 km(2)). Laboratory data overestimated consumption at high prey densities, whereas predictions based on local scale data tallied closely with consumption observed at the regional scale. The predicted regional functional response was different for increasing and decreasing prey densities, reflecting that predator and prey densities, as well as the covariance between them, exhibit oscillatory dynamics. We conclude that it is important to validate laboratory data with small-scale field observations and that scale transition is a powerful tool for scaling-up process functions in heterogeneous systems.

Plugging space into predator-prey models: an empirical approach.

Extrapolating ecological processes from small-scale experimental systems to scales of natural populations usually entails a considerable increase in spatial heterogeneity, which may affect process rates and, ultimately, population dynamics. We demonstrate how information on the heterogeneity of natural populations can be taken into account when scaling up laboratory-derived process functions, using the technique of moment approximation. We apply moment approximation to a benthic crustacean predator-prey system, where a laboratory-derived functional response is made spatial by including correction terms for the variance in prey density and the covariance between prey and predator densities observed in the field. We also show how moment approximation may be used to incorporate spatial information into a dynamic model of the system. While the nonspatial model predicts stable dynamics, its spatial equivalent also produces bounded fluctuations, in agreement with observed dynamics. A detailed analysis shows that predator-prey covariance, but not prey variance, destabilizes the dynamics. We conclude that second-order moment approximation may provide a useful technique for including spatial information in population models. The main advantage of the method is its conceptual value: by providing explicit estimates of variance and covariance effects, it offers the possibility of understanding how heterogeneity affects ecological processes.

Reducing spatial variation in environmental assessment of marine benthic fauna.

The Benthic Quality Index, BQI, is widely used for benthic quality assessment. Here, we investigated if spatial variation in the BQI can be reduced by accounting for the environmental factors instead of having different boundaries for different salinity regimes between status classes in the EU Water Framework Directive and Marine Strategy Framework Directive. For this purpose we tested salinity, sediment structure, and depth in a regression model to test their contribution to variations in BQI. The spatial variation in BQI was better explained by depth than by salinity or sediment structure. The proposed assessment method uses the residuals from the regression model between BQI and depth. With this method the variance in BQI between samples was reduced by 50% to 75% in the majority of situations. A method to establish the boundary between good and moderate status and how to derive EQR-values according to the WFD is presented.

Calculation of species sensitivity values and their precision in marine benthic faunal quality indices.

A challenging aspect of benthic quality indices used for assessing the marine environment has been to compile reliable measures of the species' sensitivity to disturbances. Sensitivity values and their uncertainties can be calculated, but a problem to cope with is that the results may depend on the actual proportion of samples from disturbed and undisturbed environments. Here we calculated sensitivity values for each species along an artificial disturbance gradient created by bootstrapping varying numbers of samples from disturbed and undisturbed environments. The values were increasing, decreasing, or more or less constant along this gradient. The lowest value with the lowest uncertainty was adopted as the species sensitivity value. Analyses of the uncertainties indicated that the accuracy rather than the precision might be a concern. We suggest a method to exclude species for which the uncertainty is outside predefined limits as a precaution to reduce bias in the environmental status classification.

Multiple density dependence in two sub-populations of the amphipod Monoporeia affinis: a potential for alternative equilibria.

Possible mechanisms for differences in population densities and dynamics were investigated in the amphipod Monoporeia affinis at two deep sites in the northern Bothnian Sea. The two sites were sampled yearly for 10 years. Average sizes, growth and mortality of the different age-classes were estimated from the cohort structure of the two populations. Laboratory experiments also investigated the ability of the common predatory isopod Saduria entomon to cause densitydependent (DD) mortality of the prey M. affinis. At site A, 43 m depth, the average density of M. affinis was twice as high as at site B, 81 m depth. The fluctuations in density were asynchronous between the two sites. Recruitment and subadult sizes of Monoporeia affinis were density dependent at both sites. The main functional difference between the two populations seemed to be the DD mortality of the 1 + cohort that occurred only at the low-density site B. A corresponding DD mortality was found in the predation experiments at densities of 1 + m. affinis corresponding to those found at site B. The potential importance of the predator was also indicated by a significant negative correlation between the biomass of S. entomon and the rate of change in M. affinis density in the field. The similarities in the abiotic factors between the two sites suggested that differences in carrying capacity should be small. The results could be explained by the predation regulation hypothesis for the low-density population at site B, while at site A M. affinis seemed to be regulated by intra-specific competition and limited by predation. It is suggested that in this simple predator-prey system there is potential for the existence of alternative equilibria.

Effects of large Saduria entomon (Isopoda) on spatial distribution of their small S. entomon and Monoporeia affinis (Amphipoda) prey.

We performed laboratory experiments to investigate the effects of predator avoidance and numerical effects of predation on spatial distribution of small Saduria entomon (Isopoda) and Monoporeia affinis (Amphipoda), with large S. entomon as predators. The horizontal distribution and mortality of the prey species, separately and together, were studied in aquaria with a spatial horizontal refuge. We also estimated effects of refuge on mortality of small S. entomon and M. affinis by experiments without the refuge net. In addition, we investigated whether predation risk from large S. entomon influenced the swimming activity of M. affinis, to clarify the mechanisms behind the spatial distribution. Both small S. entomon and M. affinis avoided large S. entomon. The avoidance behaviour of M. fffinis contributed about 10 times more to the high proportion in the refuge than numerical effects of predation. Due to the low mortality of small S. entomon the avoidance behaviour of this species was even more important for the spatial distribution. The combined effect of avoidance behaviour and predation in both species was aggregation, producting a positive correlation between the species in density. M. affinis showed two types of avoidance behaviour. In the activity experiments they reduced activity by 36% and buried themselves in the sediment. In the refuge experiments we also observed avoidance behaviour with the emigration rate from the predator compartment being twice the immigration rate. The refuge did not lower predation mortality in M. affinis, probably due to the small scale of the experimental units in relation to the mobility of the species. Predation mortality in small S. entomon was higher in absence of a refuge and especially high in absence of M. affinis.

Direct and indirect effects of predation by Saduria entomon (Isopoda) on the size-structure of Monoporeia affinis (Amphipoda).

We performed a 6-month laboratory experiment to investigate the direct and indirect effects of predation by the benthic invertebrate predator Saduria entomon on the growth and survival of Monopreia affinis prey individuals in different age-cohorts at low and high prey densities. The experimental results were compared with changes of growth and abundance in corresponding age-cohorts of M. affinis at sites with different S. entomon and M. affinis densities in the deep sublittoral zone of the Bothnian Sea during the same year. In the experiment, the presence of S. entomon reduced growth rate of M. affinis in the 1-year and 2-year age-cohorts at low amphipod density. Increased refuge use by M. affinis, expressed as a decrease in swimming activity in the presence of S. entomon, is suggested to have reduced feeding rate and therefore growth of the amphipods. The recruitment of M. affinis offspring was reduced in presence of S. entomon. In the field, the growth rate of amphipods in the 1-year cohort increased with increasing S. entomon density at low amphipod density. We found no corresponding increase of M. affinis growth in the 2-year cohort. The positive effect on 1-year amphipod growth indicated that predation reduced intra-cohort competition of M. affinis and increased growth of surviving prey at high predator density. In both the experiment and the field data we found indications of size-selective predation on smaller M. affinis specimens. This was because of the changed ratio between number of individuals in the juvenile age-cohorts and lower recruitment of amphipod offspring connected to S. entomon density. The experimental results and field data suggest that predation by S. entomon may have both direct and indirect effects on the size-structure of M. affinis populations.

Partial migration and transient coexistence of migrants and residents in animal populations.

Partial migration, whereby a proportion of the population migrates, is common across the animal kingdom. Much of the focus in the literature has been on trying to explain the underlying mechanisms for the coexistence of migrants and residents. In addition, there has been an increasing number of reports on the prevalence and frequency of partially migratory populations. One possible explanation for the occurrence of partial migration, which has received no attention in the literature, is that of 'transient coexistence' during the invasion phase of a superior behaviour. In this study we develop a theoretical basis for explaining partial migration as a transient coexistence and derive a method to predict the frequency of residents and migrants in partially migrating populations. This method is useful to predict the frequencies of migrants and residents in a small set of populations as a complementing hypothesis to 'an Evolutionary Stable Strategy (ESS)'. We use the logistic growth equation to derive a formula for predicting the frequencies of residents and migrants. We also use simulations and empirical data from white perch (Morone americana), moose (Alces alces) and red deer (Cervus elaphus) to demonstrate our approach. We show that the probability of detecting partial migration due to transient coexistence depends upon a minimum number of tracked or marked individuals for a given number of populations. Our approach provides a starting point in searching for explanations to the observed frequencies, by contrasting the observed pattern with both the predicted transient and the uniform random pattern. Aggregating such information on observed patterns (proportions of migrants and residents) may eventually lead to the development of a quantitative theory for the equilibrium (ESS) populations as well.

Influence of contaminant burial depth on the bioaccumulation of PCBs and PBDEs by two benthic invertebrates (Monoporeia affinis and Marenzelleria spp.).

The bioaccumulation of buried polychlorinated biphenyls (PCBs) and polybrominated diphenyl ethers (PBDEs) added to specific depths in sediment (2.0-2.5, 5.0-5.5 and 10.0-10.5cm) was studied in two infaunal species with similar feeding habits (surface deposit-feeders) but different bioturbation modes. The deep-burrowing polychaetes Marenzelleria spp. (Mz) displayed up to 36 times higher tissue concentrations of buried (spiked) contaminants than the surface-dwelling biodiffusing amphipod Monoporeia affinis. The differences in bioaccumulation were most pronounced for less hydrophobic contaminants due to the bioirrigating activity of Mz. Contaminants buried at shallow depths displayed higher accumulation than more deeply buried contaminants. In contrast, the bioaccumulation of unspiked (native) contaminants with a uniform vertical distribution in the sediment was similar between the species. For Mz, the BSAFs increased with increased K(OW) for the uniformly distributed contaminants, but decreased for the buried contaminants, which indicates that the dominant uptake routes of the buried contaminants can differ from the uniformly distributed contaminants. The surface sediment concentration of buried contaminants increased in Mz treatments, showing that Mz bioturbation can remobilize historically buried contaminants to the biologically active surface layer and increase the exposure for surface-dwelling species.

Theoretical and practical aspects on benthic quality assessment according to the EU-Water Framework Directive--examples from Swedish waters.

A previously presented objective method to calculate each species sensitivity to disturbance is here slightly modified and implemented in the Benthic Quality Index (BQI) for marine benthic invertebrates. A framework for assessment of water bodies based on multi-site BQI-values is also presented, where a certain variation of BQI-values is allowed to cover the heterogeneity within each water body. The 20th percentile, using bootstrapping, from the available sites' BQI-values is compared with the status boundaries for quality assessment. The reliability of the assessment depends on the background information available for the boundary setting as well as the number of sampling sites included in the assessment. Agreement between time series of quality assessments in areas with known changes in anthropogenic disturbances is encouraging. Problems associated with water body assessment based on few or no samples, as well as multiple sampling occasions during the 6-yr WFD cycle are discussed.

Prediction of mecA-positive coagulase-negative staphylococci: assessment of different phenotypic methods, breakpoints, culture media and culture conditions.

Sensitivity for the detection of mecA-positive coagulase-negative staphylococci (CNS) was evaluated for different breakpoints of resistance for oxacillin using three different susceptibility tests, either on Mueller-Hinton agar supplemented with 2% NaCl (MH-NaCl agar) or on paper disc method agar supplemented with 5% defibrinated blood (PDM-blood agar). The Etest, multipoint inoculation test and disc diffusion test showed comparable sensitivity (0.96, 0.96 and 0.95, respectively) using an oxacillin breakpoint of > or = 0.5 mg/L or < or = 17 mm for the disc test, after incubation at 35 degrees C for 24 h on MH-NaCl agar. The sensitivity decreased for breakpoints > or = 1 mg/L and when PDM-blood agar was used instead of MH-NaCl agar.