The duration of sea and freshwater residence of the sea trout,Salmo trutta,from the Vardnes River in northern Norway

Synopsis The sea trout,Salmo trutta, from the Vardnes River generally stays in freshwater during winter. The results from 11 years of tagging experiments showed that the mean annual duration of the feeding migration in the sea was 68 ± 21 days (yearly means ranging between 54 and 88 days). A pronounced variation was found with size, sex and time of migration of the fish and between the different years. The males stayed for a shorter time in the sea (mean 66 days) than the females (mean 69 days) and they usually descended later and ascended earlier. Those fish that descended first stayed for the longest period in the sea. The mean annual duration of stay in the sea for first and second time migrants to the sea was positively correlated with the mean sea temperature in May (r² > 0.5). A positive correlation was also found between the annual duration of stay in the sea for the sea trout above 33 cm and the mean temperature in the river in July (r² > 0. 6). Prolonged mean yearly sea sojourn was generally correlated with higher sea temperatures. Water temperature-salinity effects are discussed.     

Migratory timing, marine survival and growth of anadromous brown trout Salmo trutta in the River Imsa, Norway

The aim of the paper was to study sea migration, growth and survival of brown trout Salmo trutta of the River Imsa, 1976–2005. The migratory S. trutta were individually tagged and fish leaving or entering the river were monitored daily in traps located near the river mouth. The mean annual duration of the sea sojourn was 6–9 months for first-time migrants moving to sea between January and June. It was 8–18 months for those migrating to sea between July and December. Veteran migrants stayed 12 months or less at sea and most returned to the river in August. Early ascending fish stayed the longest in fresh water because most returned to sea in April to May. The day number of 50% cumulative smolt descent correlated negatively with mean water temperature in February to March and the February North Atlantic Oscillation index (NAOI). Mean annual sea growth during the first 2 years after smolting was higher for S. trutta spending the winter at sea than those wintering in the River Imsa. First year's sea growth was lower for S. trutta descending in spring than autumn. For first-time migrants, it correlated negatively with the February NAOI of the smolt year. Sea survival was higher for spring than autumn descending first-time migratory S. trutta with a maximum in May (14·9%). Number of anadromous S. trutta returning to the river increased linearly with the size of the cohort moving to sea, with no evidence of density-dependent sea mortality. Sea survival of S. trutta smolts moving to sea between January and June correlated positively both with the annual number of Atlantic Salmo salar smolts, the specific growth rate at sea, and time of seaward migration in spring. This is the first study indicating how environmental factors at the time of seaward migration influence the sea survival of S. trutta .     

The influence of precocious sexual maturation on survival to adulthood of river stocked Baltic salmon, Salmo salar, smolts

During three consequtive years, 1975–1977, Individually tagged Baltic salmon Salmo salar smolts of sexually immature male and female fish (n = 35027, mean size: 15.2 cm) and precocious males (n = 6518, mean size: 14.2 cm) were released into Umeälven (Ume river), northern Sweden. Rate of survival (% captured adults) based on 3714 recoveries was significantly higher (p < 0.01) for smolts from immature fish (10.2%) than those from smolts of early maturing males, i.e. precocious males (2.2%). corresponding to an average yield of 474 and 85 kg per KHX) smolts released, respectively. Gain in survival was on average 2.5% and 1.4% per cm increase in smolt size for immature smolts and smolts from precocious males, respectively. The poor survival among smolts of precocious males is suggested to he related to an interaction between sexual maturation and smolting linked to incompletely resorbed gonads leading to a non migratory behaviour. These non migratory males are then suggested to suffer heavily by predation in the river.
The two smolt categories had a similar growth pattern in sea. Smolts from precocious males did not mature early in sea indicating no relation to grisling, i.e. sexually maturing fish returning after first winter in sea. Adult weight of fish returning the fourth summer after release was related to smolt size (P < 0.05). Our Response Surface Model (RSA) predicted that large smolts (19.0 cm) had a higher specific growth rate over their life-span compared to small smolts (<15.0 cm), 0.86% d⁻¹ and 0.46% d⁻¹, respectively. Large smolts (19.0 cm) attained a size of 3.0 kg during their second winter in sea about six months earlier than small smolts (13.0 cm). The paper discusses alternative release strategies that can be employed if the ultimate goal of salmon stocking is maximizing yield.     

The timing, sex and age composition of the wild and reared Atlantic salmon ascending the Simojoki River, northern Finland

In the Simojoki River in the northern Gulf of Bothnia, reared salmon stocked as smolts produced considerable numbers of ascending one‐sea‐winter (1 SW) males, whereas the proportion of male 1 SW salmon was low among spawning migrants of wild or reared parr origin. The sex ratio among ascending wild fish and reared salmon stocked as parr was similar, with females predominating, while reared salmon stocked as smolts were mainly males. The multi‐sea‐winter (MSW) salmon entered the river annually within a fairly short time period from the beginning of the migration season, independent of their sex or origin. 1 SW males migrated into the river significantly later in the season than MSW males. The results indicate that the delayed opening of the fishing season in the Gulf of Bothnia is effective in reducing the harvest of MSW salmon at sea. However, as the timing of the ascent may vary by several weeks from year to year, the effect of this regulation bound to certain calendar days may also vary considerably from year to year.     

Seasonal patterns of growth, lipid deposition and lipid depletion in anadromous Arctic charr

Seasonal patterns of growth, lipid deposition and lipid depletion were studied in anadromous Arctic charr from a north Norwegian population. Samples were collected in late May when fish were migrating between fresh water and the sea, and in mid-July when the fish re-entered fresh water. A sample of maturing fish captured in mid-July was held in captivity until late September to assess lipid mobilization linked to the final stages of maturation. The fish increased substantially in weight during their 40–50-day summer residence in sea water (immatures from c . 300 to 600 g; maturing fish from c . 500 to 800 g), and body lipid stores were increased approximately fivefold. The carcass (head, skeleton and skin) was the major lipid depot, accounting for c . 50% of the total lipid content when the fish re-entered fresh water from the sea. The muscle of the ascending charr contained 35–40% of the body lipids, whereas the gut and liver each held 4–5%. Body lipid decreased 30–40% during the period between the re-entry of the fish to fresh water and spawning; although lipids were depleted from all depots the carcass and muscle were quantitatively the most important. By the time of spawning, the gonads of the females held almost 25% of the body lipid, whereas in the males the gonads accounted for <3% of the total lipids. Females lost c . 80% of their body lipid during spawning and overwintering, and, consequently, the lipid depots were severely depleted by the early spring. By contrast, lipid depletion in the males amounted to 50–55% of total lipid in the same period. These data suggest that the combination of spawning and overwintering in fresh water imposes a greater load on the females than on the males. It may be that the severity of the depletion prevents females from spawning in successive years.     

Effect of origin, sex and sea age of Atlantic salmon on their recapture rate after river ascent

The recapture rate of Atlantic salmon (Salmo salar L.) after river ascent was examined by the trapping and tagging of ascending spawners in the lower reaches of the Simojoki River, which flows into the northern Baltic Sea. In 1997 and 1998, altogether 825 Carlin‐tagged salmon were released to continue their upstream migration. Of these, 800 could be sexed and categorized as reared (91%) or wild (9%) salmon. In 1997, most of the ascending salmon were multi‐sea‐winter (MSW) fish, whereas in 1998 almost all were one‐sea‐winter (1SW) male grilse due to the late trapping season. About 10% of all tagged fish were recaptured, two‐thirds of which were caught in the river before their descent to the sea. There was no difference in the recapture rate between salmon of wild (8.5%) or reared (9.5%) origin, or between females (11.6%) and males (9.3%). Generalized linear models for data from 1997 showed that the recapture rate increased with length and age of females, but that the opposite was true for males. River fishing did not seem to remove proportionally more early ascending salmon than fish that ascended later.     

Life history and habitat use of Norwegian brown trout (Salmo trutta)

SUMMARY. 1. Brown trout ( Salmo trutta) life history and habitat use were studied in two Norwegian rivers: the Vosso river system, western Norway, and the Søre Osa, eastern Norway.
2. Age-groups were partly segregated in feeding habitats, the youngest fish living mainly in running water and in the littoral zone of lakes, the older fish also exploiting pelagic waters and deeper epibenthic habitats. In a population with free access to and from the sea, some individuals smoltified and became sea-run migrants, performing yearly migrations to the coastal sea, whereas others stayed as freshwater residents throughout their entire life span.
3. Within local populations, females were larger and less variable in size than males. This was partly because females matured at an older age than males, partly because the sexes tended to exploit feeding habitats with different food and growth Conditions. Within age-groups, females were more pelagic and migrated more than males, whereas males were more confined to running water and epibenthic areas than females. In the pelagic zone, males were more abundant in near-surface water, and females more abundant in deeper areas. When exploiting the same feeding areas, the two sexes grew at the same rate. There therefore appears to be a connection between feeding habitat and the reproductive ecology of brown trout.     

Maturation in chinook salmon, Oncorhynchus tshawytscha (Walbaum): early identification based on the development of a bimodal weight-frequency distribution

Male chinook salmon, orhynchus tshawytscha (Walbaum), are known to mature sexually after one summer in sea water, one year earlier than any of the females. Two year classes of first generation domestic chinook salmon stock reared to sexual maturity on a commercial salmon farm in British Columbia, Canada were sampled for wet weight at various times during the spring after their first winter in the sea. Frequency distributions of wet weight were developed for both year classes. The weight-frequency distributions for both cohorts became statistically bimodal in May of their first year in the sea. The fish from the two modal groups for both cohorts were sorted and kept separate over the summer. The difference in the mean weight for the two groups increased throughout the summer in both years. The higher weight group in each cohort proved to be >95% early maturing males, and represented 64–46% of all the early maturing males evident in the September following the sort. The potential use of this information for commercial salmon growers and fisheries scientists interested in sexual maturation of the chinook salmon is discussed.     

Fluctuating recruitment and variable life history of migratory brown trout, Salmo trutta L., in a small, unstable stream

The recruitment dynamics and life history of migratory brown trout, Sulmo trutta L., were investigated in a small Baltic coast stream subject to recurring drought. Spawning males consisted of both mature male parr (101–206 mm t.l. ) and migrant males (205–780 mm t.l. ). Spawning females were all migrants which delayed maturity until reaching a significantly greater size on average (424–805 mm t.l. ) than migrant males. Male: female ratios were very high in spawning aggregations (9–12 males: 1 female) with males representing up to five year-classes or more. Gametes from several generations of males per spawning event may be important for maintaining the genetic viability of this population with few female spawners per year. The amount of spawning was dependent on precipitation just prior to and during the spawning period since migrants could not enter the stream under drought conditions. Migrants did not overwinter in the stream.
Drought also caused variable fry mortality following emergence in early summer. Recruitment of 0+ parr in autumn varied from c . 175 to 3000 during 3 years. Smolts were relatively young (ages 1–2) and small (≥8 cm), and were significantly longer on average than sibling parr. Yet emigration of 1-year-olds was not related to 0+ parr size the previous autumn because of overlapping growth rates.
Persistence of the migratory brown trout in this unstable environment may be the consequence of (i) life history adaptation (e.g. short freshwater residence of both juveniles and spawners), and (ii) a complementary set of individual life histories where variation in age of migrant spawners and the occurrence of mature male parr result in a stable spawner population despite inconsistent recruitment of migrants to the sea.     

Winter feeding: an adaptive strategy broadening the niche of the central mudminnow,Umbra limi

Synopsis Mudminnows, Umbra limi, were euryphagic carnivores during the ice-free period consuming invertebrates of terrestrial and aquatic origins. In winter, diversity and amount of food eaten were reduced. Age 0 fish consumed aquatic invertebrates as did males aged 1 and 2+ but females aged 1 and 2+ consumed mainly fishes. In the laboratory mudminnows were able to capture small fish at temperatures of 12.6° C and less. Their rates of digestion of fish prey varied directly with temperature and estimates of time for 100%, gastric evacuation ranged from 161 h at 1.1° C to 26 h at 12.6° C. The ability to forage actively during winter and to digest food relatively rapidly at cold temperatures are seen as mechanisms to broaden the niche along dimensions of resources utilized and time, thereby increasing the chances of survival and reproduction in an environment with high temporal heterogeneity.     

Incubation and brooding performance of the Antarctic Petrel Thalassoica antarctica at Svarthamaren, Dronning Maud Land

Antarctic Petrel Thalassoica antarctica incubation and brooding effort was studied at Svarthamaren, Dronning Maud Land, during the austral summer of 1991–1992. The females probably left the nest site shortly after egg laying. The duration of incubation and brooding shifts as well as the daily weight loss (absolute and proportionate) were comparable with those of other similar-sized procellariform species. Males spent more time incubating and brooding than did females, suggesting higher female energy stress due to egg laying. Incubating birds which were below average weight were likely to desert the nests before their mates returned from feeding trips. Both males and females lost approximately one-fifth of their body-weight during their first incubation shifts. Nevertheless, they increased their initial weights from egg laying to hatching and had their highest initial weights when they returned to start the shift during which the egg hatched. No factors related to adult body-weight explained the duration of the incubation shifts. Both males and females gained weight at a higher rate when at sea than they lost it during incubation, and it is suggested that factors unrelated to food availability or individual feeding skills may be important in regulating the duration of the incubation shifts and the stay at sea.     

Partial migration: niche shift versus sexual maturation in fishes

Summary Many fish populations have both resident and migratory individuals. Migrants usually grow larger and have higher reproductive potential but lower survival than resident conspecifics. The decision about migration versus residence probably depends on the individual growth rate, or a physiological process like metabolic rate which is correlated with growth rate. Fish usually mature as their somatic growth levels off, where energetic costs of maintenance approach energetic intake. After maturation, growth also stagnates because of resource allocation to reproduction. Instead of maturation, however, fish may move to an alternative feeding habitat and their fitness may thereby be increased. When doing so, maturity is usually delayed, either to the new asymptotic length, or sooner, if that gives higher expected fitness. Females often dominate among migrants and males among residents. The reason is probably that females maximize their fitness by growing larger, because their reproductive success generally increases exponentially with body size. Males, on the other hand, may maximize fitness by alternative life histories, e.g. fighting versus sneaking, as in many salmonid species where small residents are the sneakers and large migrants the fighters. Partial migration appears to be partly developmental, depending on environmental conditions, and partly genetic, inherited as a quantitative trait influenced by a number of genes.     

Contemporary ocean warming and freshwater conditions are related to later sea age at maturity in Atlantic salmon spawning in Norwegian rivers

Atlantic salmon populations are reported to be declining throughout its range, raising major management concerns. Variation in adult fish abundance may be due to variation in survival, growth, and timing of life history decisions. Given the complex life history, utilizing highly divergent habitats, the reasons for declines may be multiple and difficult to disentangle. Using recreational angling data of two sea age groups, one‐sea‐winter (1SW) and two‐sea‐winter (2SW) fish originated from the same smolt year class, we show that sea age at maturity of the returns has increased in 59 Norwegian rivers over the cohorts 1991–2005. By means of linear mixed‐effects models we found that the proportion of 1SW fish spawning in Norway has decreased concomitant with the increasing sea surface temperature experienced by the fish in autumn during their first year at sea. Furthermore, the decrease in the proportion of 1SW fish was influenced by freshwater conditions as measured by water discharge during summer months 1 year ahead of seaward migration. These results suggest that part of the variability in age at maturity can be explained by the large‐scale changes occurring in the north‐eastern Atlantic pelagic food web affecting postsmolt growth, and by differences in river conditions influencing presmolt growth rate and later upstream migration.     

Variation in the degree of silvering of wild coho salmon, Oncorhynchus kisutch, smolts migrating seaward from Carnation Creek, British Columbia

Degree of silvering (colouration) is often used as an index of the degree of smoltification in salmonids. To judge the reliability of silvering as an indicator of migratory readiness, we examined the silvering of wild coho salmon, Oncorhynchus kisutch , smolts upon entry into the Carnation Creek, British Columbia, estuary. Silvering of migrants was greater in larger fish and increased over the course of the migratory period. Photoperiod appeared to account for most of the observed increase in silvering over time; increased silvering was not correlated with temperature or lunar phase. Overall, 50% of seaward migrants were completely silvered, 45% were in a transitional phase, and 5% still retained colouration characteristics of parr. Colouration thus appears to have limited utility as a reliable indicator of migratory readiness. Our findings suggest that estuarine residence is important for completion of parr-smolt transformation and that acclimation of hatchery smolts in brackish water prior to sea water entry may enhance their survival.     

Mixed-Effects Modelling of Scale Growth Profiles Predicts the Occurrence of Early and Late Fish Migrants

Fish growth is commonly used as a proxy for fitness but this is only valid if individual growth variation can be interpreted in relation to conspecifics'' performance. Unfortunately, assessing individual variation in growth rates is problematic under natural conditions because subjects typically need to be marked, repeated measurements of body size are difficult to obtain in the field, and recaptures may be limited to a few time events which will generally vary among individuals. The analysis of consecutive growth rings (circuli) found on scales and other hard structures offers an alternative to mark and recapture for examining individual growth variation in fish and other aquatic vertebrates where growth rings can be visualized, but accounting for autocorrelations and seasonal growth stanzas has proved challenging. Here we show how mixed-effects modelling of scale growth increments (inter-circuli spacing) can be used to reconstruct the growth trajectories of sea trout (Salmo trutta) and correctly classify 89% of individuals into early or late seaward migrants (smolts). Early migrants grew faster than late migrants during their first year of life in freshwater in two natural populations, suggesting that migration into the sea was triggered by ontogenetic (intrinsic) drivers, rather than by competition with conspecifics. Our study highlights the profound effects that early growth can have on age at migration of a paradigmatic fish migrant and illustrates how the analysis of inter-circuli spacing can be used to reconstruct the detailed growth of individuals when these cannot be marked or are only caught once.     

Timing and pattern of annual silver eel migration in two European watersheds are determined by similar cues

Many animals perform long‐distance migrations in order to maximize lifetime reproductive success. The European eel migrates several thousand kilometers between their feeding habitats in continental waters (fresh‐, brackish, and sea water) and their spawning area in the Sargasso Sea. Eels residing in freshwaters usually initiate their spawning migration as silver eels during autumn, triggered by diverse environmental cues. We analyzed the time series of silver eel downstream migration in Burrishoole, Ireland (1971–2015), and Imsa, Norway (1975–2015), to examine factors regulating the silver eel migration from freshwater to the sea. The migration season (90% of the run) generally lasted from 1 August to 30 November. Environmental factors acting in the months before migration impacted timing and duration of migration, likely through influencing the internal processes preparing the fish for migration. Once the migration had started, environmental factors impacted the day‐to‐day variation in number of migrants, apparently stimulating migration among those eels ready for migration. Both the day‐to‐day variation in the number of migrants and the onset of migration were described by nearly identical models in the two rivers. Variables explaining day‐to‐day variation were all associated with conditions that may minimize predation risk; number of migrants was reduced under a strong moon and short nights and increased during high and increasing water levels. Presence of other migrants stimulated migration, which further indicates that silver eel migration has evolved to minimize predation risk. The onset of migration was explained mainly by water levels in August. The models for duration of the migration season were less similar between the sites. Thus, the overall migration season seems governed by the need to reach the spawning areas in a synchronized manner, while during the actual seaward migration, antipredator behavior seems of overriding importance.     

The post‐spawning movements of migratory brown trout Salmo trutta L.

The post spawning behaviour of sea trout Salmo trutta was studied over a 2 year period in the river and estuary of the River Fowey, south‐west England. Forty‐five sea trout kelts were trapped immediately after spawning in December and intraperitoneally tagged with miniature acoustic transmitters. The subsequent emigration into coastal waters was monitored using acoustic receivers deployed throughout the river catchment. The levels of gill Na⁺K⁺ATPase activity in sea trout kelts sampled at the same time as the tagged fish were within the range of 2·5 to 4·5 μmol Pi per mg protein per h indicating that the post‐spawning fish were not physiologically adapted to salt water. The tagged kelts were resident in fresh water between 4 and 70 days before entering the estuary. Sixty two per cent of the tagged kelts subsequently migrated successfully into coastal waters, with a higher success rate for male fish (75%) than females (58%). There was a significant size related difference in the run‐timing of the kelts with the larger fish moving more quickly into coastal waters after spawning than smaller fish. Seaward migration within fresh water was predominantly nocturnal and generally occurred in conjunction with increasing river discharge and rising water temperature. Migration through the estuary continued to be predominantly nocturnal and occurred during an ebbing tide. Residency within the estuary varied amongst individuals although it was invariably short, with most fish moving out into coastal waters within one to two tidal cycles. Five tagged kelts returned from the coastal zone and re‐entered fresh water during April and June. Marine residence time varied between 89 and 145 days (mean 118 days) and the minimum estimated marine survival was c. 18%. One of these sea trout was subsequently recaptured after successfully spawning in the vicinity where it had been previously tagged demonstrating a degree of spawning site fidelity.     

Feeding dynamics of Palaemon peringueyi (Decapoda,Caridea) in the temporarily open/closed Kasouga Estuary,South Africa

The feeding ecology of the caridean shrimp Palaemon peringueyi in the temporarily open/closed Kasouga Estuary, Eastern Cape, was examined during summer and winter employing stomach fullness index (SFI) analysis. Mean SFI values for male and female shrimp during summer ranged from 0.85% to 2.36% and from 0.56% and 2.61% body dry weight, respectively. During winter, SFI values for males and females were lower, and varied between 0.23% and 1.56% and 0.36% and 1.84% body dry weight, respectively. There were no significant differences in the SFI values between males and females during the two seasons. A peak in the feeding activity for both sexes was observed during the daytime during both summer and winter. For both males and females, non-linear regression analysis best explained the change in SFI over time. The gut evacuation rate constants (k) of males and females during summer were estimated at 0.43 h^?1 and 0.30 h^?1, respectively. These values correspond to a daily food intake equivalent to 16.2% body dwt for males and 11.1% body dwt for females. During winter, the gut evacuation rates of the shrimp were estimated at 0.35 h^?1 for males and 0.28 h^?1 for females. The daily ration for adult male and female shrimp during winter was estimated at 6.1% and 6.8% body dwt, respectively. The lower estimates obtained during winter appear largely to reflect the influence of the reduced water temperatures on the metabolic activities of the shrimp.     

Effects of seawater-acclimatization and release sites on survival of hatchery–reared brown trout Salmo trutta

To test whether seawater–acclimatization of hatchery–reared anadromous and freshwater resident brown trout before release increased the survival of adults, smolts were retained 0, 2, 4 and 8 weeks in sea water before release. Total recapture rate increased for smolts retained 4 and 8 weeks in sea water before release relative to the controls. This trend was more pronounced for Freshwater resident than for anadromous stocks, Offspring of anadromous fish stayed longer at sea than offspring of freshwater resident fish. Recapture rates in fresh water were higher for brown trout released in the river than in the fjord in the R, Drammen area, but not in the R. Imsa. In both cases, most fish were recaptured in the sea. Moving into the R. Imsa (relative to other rivers) appeared higher for fish released at the mouth of the river (93%) than in the fjord (47%). Judged from the recapture rates, sea survival appeared to be the same whether released in the fjord or at the mouth of the river.     

Sex differences in contaminant concentrations of fish: a synthesis

A comparison of whole-fish polychlorinated biphenyl (PCB) and total mercury (Hg) concentrations in mature males with those in mature females may provide insights into sex differences in behavior, metabolism, and other physiological processes. In eight species of fish, we observed that males exceeded females in whole-fish PCB concentration by 17 to 43 %. Based on results from hypothesis testing, we concluded that these sex differences were most likely primarily driven by a higher rate of energy expenditure, stemming from higher resting metabolic rate (or standard metabolic rate (SMR)) and higher swimming activity, in males compared with females. A higher rate of energy expenditure led to a higher rate of food consumption, which, in turn, resulted in a higher rate of PCB accumulation. For two fish species, the growth dilution effect also made a substantial contribution to the sex difference in PCB concentrations, although the higher energy expenditure rate for males was still the primary driver. Hg concentration data were available for five of the eight species. For four of these five species, the ratio of PCB concentration in males to PCB concentration in females was substantially greater than the ratio of Hg concentration in males to Hg concentration in females. In sea lamprey (Petromyzon marinus), a very primitive fish, the two ratios were nearly identical. The most plausible explanation for this pattern was that certain androgens, such as testosterone and 11-ketotestosterone, enhanced Hg-elimination rate in males. In contrast, long-term elimination of PCBs is negligible for both sexes. According to this explanation, males not only ingest Hg at a higher rate than females but also eliminate Hg at a higher rate than females, in fish species other than sea lamprey. Male sea lamprey do not possess either of the above-specified androgens. These apparent sex differences in SMRs, activities, and Hg-elimination rates in teleost fishes may also apply, to some degree, to higher vertebrates including humans. Our synthesis findings will be useful in (1) developing sex-specific bioenergetics models for fish, (2) developing sex-specific risk assessment models for exposure of humans and wildlife to contaminants, and (3) refining Hg mass balance models for fish and higher vertebrates.