Low-level copper exposures increase visibility and vulnerability of juvenile coho salmon to cutthroat trout predators

Copper contamination in surface waters is common in watersheds with mining activities or agricultural, industrial, commercial, and residential human land uses. This widespread pollutant is neurotoxic to the chemosensory systems of fish and other aquatic species. Among Pacific salmonids (Oncorhynchus spp.), copper-induced olfactory impairment has previously been shown to disrupt behaviors reliant on a functioning sense of smell. For juvenile coho salmon (O. kisutch), this includes predator avoidance behaviors triggered by a chemical alarm cue (conspecific skin extract). However, the survival consequences of this sublethal neurobehavioral toxicity have not been explored. In the present study juvenile coho were exposed to low levels of dissolved copper (5-20 microg/L for 3 h) and then presented with cues signaling the proximity of a predator. Unexposed coho showed a sharp reduction in swimming activity in response to both conspecific skin extract and the upstream presence of a cutthroat trout predator (O. clarki clarki) previously fed juvenile coho. This alarm response was absent in prey fish that were exposed to copper. Moreover, cutthroat trout were more effective predators on copper-exposed coho during predation trials, as measured by attack latency, survival time, and capture success rate. The shift in predator-prey dynamics was similar when predators and prey were co-exposed to copper. Overall, we show that copper-exposed coho are unresponsive to their chemosensory environment, unprepared to evade nearby predators, and significantly less likely to survive an attack sequence. Our findings contribute to a growing understanding of how common environmental contaminants alter the chemical ecology of aquatic communities.     

Pacific harbor seals (<Emphasis Type="Italic">Phoca vitulina</Emphasis>) and salmon: genetics presents hard numbers for elucidating predator-prey dynamics

We used a PCR-based species test and microsatellite genotyping to determine salmonid species composition and quantity of Chinook among Pacific harbor seal (Phoca vitulina) prey. Tests were applied to DNA extracted from all salmonid bones recovered from scat (fecal) samples. Condition of bone samples and quantity of DNA varied substantially. Although DNA extraction and species identification was only successful for just over half of the bone samples, 93% of all scat samples had bones that were identified to species. Most often only a single salmonid prey species was evident within a scat sample (39% exclusively coho and 46% exclusively Chinook) but 13% contained bones from more than one species of salmonid. For scat samples that contained Chinook bones, 68% had skeletal remains from the same fish and 32% had bones from more than one individual Chinook, varying in number from two to four individuals per scat. In one case, bones from one individual Chinook were recovered in three different scat samples.     

Ecosystem-based management of predator-prey relationships: piscivorous birds and salmonids.

Predator-prey relationships are often altered as a result of human activities. Where prey are legally protected, conservation action may include lethal predator control. In the Columbia River basin (Pacific Northwest, USA and Canada), piscivorous predators have been implicated in contributing to a lack of recovery of several endangered anadromous salmonids (Oncorhynchus spp.), and lethal and nonlethal control programs have been instituted against both piscine and avian species. To determine the consequences of avian predation, we used a bioenergetics approach to estimate the consumption of salmonid smolts by waterbirds (Common Merganser, California and Ring-billed Gull, Caspian Tern, Double-crested Cormorant) found in the mid-Columbia River from April through August, 2002-2004. We used our model to explore several predator-prey scenarios, including the impact of historical bird abundance, and the effect of preserving vs. removing birds, on smolt abundance. Each year, <1% of the estimated available salmonid smolts (interannual range: 44,830-109,209; 95% CI = 38,000-137,000) were consumed, 85-98% away from dams. Current diet data combined with historical gull abundance at dams suggests that past smolt consumption may have been 1.5-3 times current numbers, depending on the assumed distribution of gulls along the reaches. After the majority (80%) of salmonid smolts have left the study area, birds switch their diet to predominantly juvenile northern pikeminnow (Ptychocheilus oregonensis), which as adults are significant native salmonid predators in the Columbia River. Our models suggest that one consequence of removing birds from the system may be increased pikeminnow abundance, which--even assuming 80% compensatory mortality in juvenile pikeminnow survival--would theoretically result in an annual average savings of just over 180,000 smolts, calculated over a decade. Practically, this suggests that smolt survival could be maximized by deterring birds from the river when smolts are present, allowing bird presence after the diet switch to act as a tool for salmonid-predator control, and conducting adult-pikeminnow control throughout. Our analysis demonstrates that identifying the strength of ecosystem interactions represents a top priority when attempting to manage the abundance of a particular ecosystem constituent, and that the consequences of a single-species view may be counterintuitive, and potentially counterproductive.     

Salmon, wildlife, and wine: marine-derived nutrients in human-dominated ecosystems of central California.

Pacific salmon transfer large quantities of marine-derived nutrients to adjacent forest ecosystems with profound effects on plant and wildlife production. We investigated this process for two highly modified California wine country rivers, one with consistent salmon runs (Mokelumne River) and one without (Calaveras River). Mokelumne River Chinook salmon transported biomass and N comparable to Pacific Northwest salmon streams. Calaveras River levels were much less. Scavenger numbers correlated with salmon carcass counts over time on the Mokelumne River but not the Calaveras River. Likewise, salmon carcasses were consumed significantly faster on the Mokelumne River. Native riparian vegetation as well as cultivated wine grapes adjacent to Mokelumne River spawning sites received 18-25% of foliar N from marine sources, significantly higher than vegetation along the Calaveras River. These data suggest that robust salmon runs continue to provide important ecological services with high economic value, even in impaired watersheds. Loss of Pacific salmon can not only negatively affect stream and riparian ecosystem function, but can also affect local economies where agriculture and salmon streams coexist.     

Modelling predation by transient leopard seals for an ecosystem-based management of Southern Ocean fisheries

Correctly quantifying the impacts of rare apex marine predators is essential to ecosystem-based approaches to fisheries management, where harvesting must be sustainable for targeted species and their dependent predators. This requires modelling the uncertainty in such processes as predator life history, seasonal abundance and movement, size-based predation, energetic requirements, and prey vulnerability. We combined these uncertainties to evaluate the predatory impact of transient leopard seals on a community of mesopredators (seals and penguins) and their prey at South Georgia, and assess the implications for an ecosystem-based management. The mesopredators are highly dependent on Antarctic krill and icefish, which are targeted by regional fisheries. We used a state-space formulation to combine (1) a mark-recapture open-population model and individual identification data to assess seasonally variable leopard seal arrival and departure dates, numbers, and residency times; (2) a size-based bioenergetic model; and (3) a size-based prey choice model from a diet analysis. Our models indicated that prey choice and consumption reflected seasonal changes in leopard seal population size and structure, size-selective predation and prey vulnerability. A population of 104 (90–125) leopard seals, of which 64% were juveniles, consumed less than 2% of the Antarctic fur seal pup production of the area (50% of total ingested energy, IE), but ca. 12–16% of the local gentoo penguin population (20% IE). Antarctic krill (28% IE) were the only observed food of leopard seal pups and supplemented the diet of older individuals. Direct impacts on krill and fish were negligible, but the “escapement” due to leopard seal predation on fur seal pups and penguins could be significant for the mackerel icefish fishery at South Georgia. These results suggest that: (1) rare apex predators like leopard seals may control, and may depend on, populations of mesopredators dependent on prey species targeted by fisheries; and (2) predatory impacts and community control may vary throughout the predator's geographic range, and differ across ecosystems and management areas, depending on the seasonal abundance of the prey and the predator's dispersal movements. This understanding is important to integrate the predator needs as natural mortality of its prey in models to set prey catch limits for fisheries. Reliable estimates of the variability of these needs are essential for a precautionary interpretation in the context of an ecosystem-based management.     

Distribution of salmon-habitat potential relative to landscape characteristics and implications for conservation.

The geographic distribution of stream reaches with potential to support high-quality habitat for salmonids has bearing on the actual status of habitats and populations over broad spatial extents. As part of the Coastal Landscape Analysis and Modeling Study (CLAMS), we examined how salmon-habitat potential was distributed relative to current and future (+100 years) landscape characteristics in the Coastal Province of Oregon, USA. The intrinsic potential to provide high-quality rearing habitat was modeled for juvenile coho salmon (Oncorhynchus kisutch) and juvenile steelhead (O. mykiss) based on stream flow, valley constraint, and stream gradient. Land ownership, use, and cover were summarized for 100-m analysis buffers on either side of stream reaches with high intrinsic potential and in the overall area encompassing the buffers. Past management seems to have concentrated nonindustrial private ownership, agriculture, and developed uses adjacent to reaches with high intrinsic potential for coho salmon. Thus, of the area in coho salmon buffers, 45% is either nonforested or recently logged, but only 10% is in larger-diameter forests. For the area in steelhead buffers, 21% is either non-forested or recently logged while 20% is in larger-diameter forests. Older forests are most extensive on federal lands but are rare on private lands, highlighting the critical role for public lands in near-term salmon conservation. Agriculture and development are projected to remain focused near high-intrinsic-potential reaches for coho salmon, increasing the importance of effectively addressing nonpoint source pollution from these uses. Percentages of larger-diameter forests are expected to increase throughout the province, but the increase will be only half as much in coho salmon buffers as in steelhead buffers. Most of the increase is projected for public lands, where policies emphasize biodiversity protection. Results suggest that widespread recovery of coho salmon is unlikely unless habitat can be improved in high-intrinsic-potential reaches on private lands. Knowing where high-intrinsic-potential stream reaches occur relative to landscape characteristics can help in evaluating the current and future condition of freshwater habitat, explaining differences between species in population status and risk, and assessing the need for and feasibility of restoration.     

Incorporating covariates into fisheries stock assessment models with application to Pacific herring.

We present a framework for evaluating the cause of fishery declines by integrating covariates into a fisheries stock assessment model. This allows the evaluation of fisheries' effects vs. natural and other human impacts. The analyses presented are based on integrating ecological science and statistics and form the basis for environmental decision-making advice. Hypothesis tests are described to rank hypotheses and determine the size of a multiple covariate model. We extend recent developments in integrated analysis and use novel methods to produce effect size estimates that are relevant to policy makers and include estimates of uncertainty. Results can be directly applied to evaluate trade-offs among alternative management decisions. The methods and results are also broadly applicable outside fisheries stock assessment. We show that multiple factors influence populations and that analysis of factors in isolation can be misleading. We illustrate the framework by applying it to Pacific herring of Prince William Sound, Alaska (USA). The Pacific herring stock that spawns in Prince William Sound is a stock that has collapsed, but there are several competing or alternative hypotheses to account for the initial collapse and subsequent lack of recovery. Factors failing the initial screening tests for statistical significance included indicators of the 1989 Exxon Valdez oil spill, coho salmon predation, sea lion predation, Pacific Decadal Oscillation, Northern Oscillation Index, and effects of containment in the herring egg-on-kelp pound fishery. The overall results indicate that the most statistically significant factors related to the lack of recovery of the herring stock involve competition or predation by juvenile hatchery pink salmon on herring juveniles. Secondary factors identified in the analysis were poor nutrition in the winter, ocean (Gulf of Alaska) temperature in the winter, the viral hemorrhagic septicemia virus, and the pathogen Ichthyophonus hoferi. The implication of this result to fisheries management in Prince William Sound is that it may well be difficult to simultaneously increase the production of pink salmon and maintain a viable Pacific herring fishery. The impact can be extended to other commercially important fisheries, and a whole ecosystem approach may be needed to evaluate the costs and benefits of salmon hatcheries.     

Recherches sur la situation trophique d'un groupe d'organismes pélagiques (Euphausiacea). IV. Relations avec les autres éléments du micronecton

The predation of euphausiids by micronektonic migrant or deep-living fishes has been studied by examination of 1825 stomach contents selected from 282 samples collected with a 10-foot Isaacs-Kidd midwater trawl (IKMT) in the central equatorial Pacific Ocean. On the whole, euphausiids account for 8% of the total food ingested; this percentage rises to 21% when the genus Cyclothone is excepted. The species consumed belong to the genera Euphausia (45%), Stylocheiron (40%) and Nematoscelis (13%). 90% of the specimens are smaller than 18 mm and weigh less than 37 mg (wet weight); in most cases, there exists an almost linear relationship between predator size and the occurrence of euphausiids in the stomachs and also with the sizes of the preys. It is shown that the predation of euphausiids by micronektonic migrant and deep-living fishes is less during light hours and rises in the late afternoon and night time. Moreover, the species consumed are not the same at different times of the day; Euphausia spp. are ingested by night (in subsurface layers) as well as during the daytime (in deeper layers), small Stylocheiron mainly by night in subsurface layers, Nematoscelis spp. and Stylocheiron abbreviatum chiefly at the time of vertical migrations (sunset and sunrise).     

Evidence of Natural Reproduction of Aquaculture-Escaped Atlantic Salmon in a Coastal British Columbia River

Abstract: We present evidence of the first successful natural spawning of Atlantic salmon (   Salmo salar ) documented on the Pacific coast of North America. Twelve juvenile Atlantic salmon composed of two year classes were captured in the Tsitika River, British Columbia. We analyzed restriction-length polymorphisms of PCR-amplified 5S rDNA and mtDNA to confirm that these individuals were Atlantic salmon. Scale analysis strongly suggested they were the products of natural spawning by feral adults. The gut contents, size, and condition of these individuals suggest that Atlantic salmon are successfully maturing in the Tsitika River, British Columbia. This event has raised concerns that the presence and possible establishment of feral Atlantic salmon may further jeopardize the continued persistence of already fragile native Pacific salmonids through competition for resources and occupation of niches that are currently underutilized.     

Continued decline of an Atlantic cod population: how important is gray seal predation?

The continental shelf ecosystem on the Eastern Scotian Shelf (ESS) has experienced drastic changes. Once common top predators are a small fraction of their historical abundance, and much of the current community structure is now dominated by pelagic fishes and invertebrates. Embedded within this food web, Atlantic cod and gray seal populations have recently exhibited nearly opposite trends. Since 1984, cod populations have decreased exponentially at a rate averaging 17% per year, whereas gray seals have continued to increase exponentially at a rate of 12%. We reexamined the impact of gray seals on Atlantic cod dynamics using more than 30 years of data on the population trends of cod and gray seals while incorporating new information on seal diet and seasonal distribution. The closure of the cod fishery over 10 years ago allowed for a better estimation of natural mortality rates. We quantified the impact of seals on ESS cod by (1) estimating trends in seal and cod abundance, (2) estimating the total energy needed for seal growth and maintenance from an energetics model, (3) using estimates of the percentage of cod in the total diet derived from quantitative fatty acid signature analysis (QFASA) and of the size-specific selectivity of cod consumed (derived from otoliths collected from fecal samples), and (4) assuming a gray seal functional response. Uncertainties of the model estimates were calculated using the Hessian approximation of the variance-covariance matrix. Between 1993 and 2000, cod comprised, on average, < 5% of a gray seal's diet. Our model shows that, since the closure of the fishery, gray seals have imposed a significant level of instantaneous mortality (0.21), and along with other unknown sources of natural mortality (0.62), are contributing to the failure of this cod stock to recover.     

Prioritizing Pacific Salmon Stocks for Conservation

Over 300 native stocks of Pacific salmon, steelhead, and coastal cutthroat trout (Oncorhynchus spp.) are at risk of extinction in the Pacific Northwest. With only limited resources available for conservation and recovery, prioritization of these stocks may become necessary if meaningful measures are to be implemented. We propose criteria by which prioritization may be guided. First, we rank stocks for risk of extinction, either by population viability analysis or by a set of surrogate measures. Then we rank stocks for biological consequences of extinction, using sets of questions designed to establish the genetic and evolutionary consequences and the ecological consequences if a stock were to become extinct. Together, these rankings allow stocks to be prioritized for a range of possible actions, with those stocks at highest risk and bearing the greatest biological consequences of extinction receiving attention first. Application of the prioritization process to 20 Pacific anadromous salmonid stocks worked as intended, although data limitations are considerable. The process is most likely to work successfully when applied to many stocks on which data exist, when several experts carry out the prioritization, and when the results are peer reviewed.     

Feeding and aggressive behaviours in juvenile coho salmon (Oncorhynchus kisutch) under chemically-mediated risk of predation

Summary Juvenile coho salmon (Oncorhynchus kisutch) spend the first year of their lives in their natal streams, where they may often hold feeding territories. They also face significant risk of predation by birds and fish, and should alter their behaviour to reduce risk of mortality when these predators are present. Although there is laboratory evidence that coho react to predator visual stimuli, chemoreception of avian predator presence has not previously been reported. We tested the influence of chemical stimuli of common merganser (Mergus merganser), preying on juvenile coho, on two aspects of coho territorial behaviour, foraging and aggression, in flow-through aquaria. After a mixture of merganser- and coho-conditioned water was introduced into the system, juvenile coho significantly reduced their attack distance on drifting prey. The fish also significantly decreased their aggressive behaviour directed towards mirrors (total number of acts, intensity of acts and time spent) when the same odour was present. They did not change their behaviour in either experiment after control introductions of water treated with fish alone. These results are interpreted within the framework of a trade-off between juvenile growth and mortality.     

Techno-Arrogance and Halfway Technologies: Salmon Hatcheries on the Pacific Coast of North America

Humankind has adopted an arrogant and ultimately self-defeating attitude toward nature that places technological mastery over nature at the forefront of our approach to many environmental problems. This "techno-arrogance" fails to recognize limitations on, and ramifications of, attempted control of nature. An example of techno-arrogance is the flawed attempt to recover Pacific salmonid fisheries through technological application in the form of hatcheries. Countless salmon stocks have declined precipitously over the last century as a result of overfishing and widespread habitat destruction. A central feature of recovery efforts has been to build many hatcheries to produce large quantities of fish to restock streams. This approach addresses the symptoms but not the causes of the declines (an example of a halfway technology), because the habitats remain largely unsuitable for salmon. There are at least six reasons why the hatchery approach will ultimately fail: (1) data demonstrate that hatcheries are not solving the problem—salmon continue to decline despite decades of hatchery production; (2) hatcheries are costly to run, and divert resources from other efforts, such as habitat restoration; (3) hatcheries are not sustainable in the long term, requiring continual input of money and energy, (4) hatcheries are a genetically unsound approach to management that can adversely affect wild populations; (5) hatchery production leads to increased harvest of declining wild populations of salmon; and (6) hatcheries conceal from the public the truth of real salmon decline. I recommend that salmonid management turn from the symptoms to the causes of decline. Overharvest and habitat destruction must be directly addressed in a major, landscape-level effort, on a scale comparable to the hatchery program, if salmonid fisheries are to remain a part of the ecological recreational, commercial and asthetic arenas in the long term.     

Evidence of a maternal foraging cycle resembling that of otariid seals in a small phocid,the harbor seal

Summary Lactation strategies in the two largest families of seals have been characterized as a phylogenetic dichotomy, with sea lions and fur seals (Otariidae) exhibiting foraging cycles and true seals (Phocidae) a strategy of fasting. We show that a lactating phocid, the harbor seal, Phoca vitulina, has a foraging cycle similar to that of otariids. Time-depth recorders attached to lactating harbor seal mothers revealed that 9 of 11 females began bouts of diving, averaging 12–40 m, by mid-lactation (12 days). During the remainder of lactation, females made an average of seven diving trips, lasting about 7 h. They returned to the rookery during the interval between successive bouts to nurse their pups. Diving was more frequent during daylight than at night and diving bouts increased in duration as lactation progressed. The diving behavior of females that had weaned their pups and previously collected data from stomach lavage, suggest that the bouts of diving represent successful foraging. We propose that the lactation strategy of the harbor seal is intermediate to that of the otariids and other phocids studied. The harbor seal has a foraging cycle like the otariids, but typically resembles other phocids in length of lactation, rate of mass gain in pups, and in milk fat content. As harbor seals are among the smallest phocids, and only slightly larger than most otariids, it seems likely that maternal size constrains the amount of stored energy harbor seal females can bring to the rookery, forcing them to start feeding during the lactation period.Correspondence to: D.J. Boness     

Colonization of the Southern Patagonia Ocean by Exotic Chinook Salmon

Abstract:  Anadromous salmonids have been particularly successful at establishing wild populations in southern Patagonia, in contrast to their limited success elsewhere outside their native ranges. The most recent such discovery is a spawning population of Chinook salmon in the Santa Cruz River, which flows into the Atlantic Ocean from Argentina. We used mitochondrial DNA analysis to discriminate between alternative potential sources of this population and were able to discard in situ introductions of fish imported directly from California in the early twentieth century. Our results showed that the fish most likely came from Puget Sound, Washington, imported into southern Chile for salmon-ranching experiments in the 1980s. This finding provides concrete evidence of colonization of Atlantic rivers from Pacific locations. The southern Pacific and Atlantic oceans provide a favorable marine environment for the success of invading salmon. In particular, the waters associated with fjords, southern channels, and the inshore portion of the Patagonian shelf provide a rather bounded, continuous waterway for exotic anadromous salmonids, rich in diverse forage species.     

Fatty acid profiles of juvenile salmon indicate prey selection strategies in coastal marine waters

Juvenile salmon exhibit high growth rates upon their arrival into the marine environment. Dietary changes from freshwater and estuarine habitats to those derived from the marine environment may play an important role in ultimate adult survival. We measured the total lipid and fatty acid (FA) composition of juvenile Chinook salmon (Oncorhynchus tshawytscha), coho salmon (O. kisutch), and 18 of their potential prey items sampled from coastal waters during their first few months at sea. Coho salmon had significant reductions in their lipid content (% wet weight) between May and June, likely due to early marine growth. We did not find a significant drop between May and June Chinook salmon lipid content, which may indicate an earlier ontogenetic selection to marine prey that are higher in lipids and essential fatty acids (EFAs). Juvenile salmon ate prey of both high and low lipids. Significant FA compositional changes occurred for both coho and Chinook salmon between May and June. In May, the FA profile of juvenile salmon, especially coho salmon, did not resemble their prey items; however, in June, there was a strong correlation between salmon and their common fish prey as determined by gut content analysis. Significant increases in the level of EFAs, especially docosahexaenoic acid (DHA, 22:6n-3) accounted for the majority of the monthly differences in salmon tissue FA composition. In order for juvenile salmon to adequately meet their physiological requirements, they may have adapted to select advantageous prey with higher levels of EFAs, especially DHA, in order to rapidly increase their growth and ultimate survival.     

Reconstruction of Pacific salmon abundance from riparian tree-ring growth.

We use relationships between modern Pacific salmon (Oncorhynchus spp.) escapement (migrating adults counted at weirs or dams) and riparian tree-ring growth to reconstruct the abundance of stream-spawning salmon over 150-350 years. After examining nine sites, we produced reconstructions for five mid-order rivers and four salmon species over a large geographic range in the Pacific Northwest: chinook (O. tschwatcha) in the Umpqua River, Oregon, USA; sockeye (O. nerka) in Drinkwater Creek, British Columbia, Canada; pink (O. gorbuscha) in Sashin Creek, southeastern Alaska, USA; chum (O. keta) in Disappearance Creek, southeastern Alaska, USA; and pink and chum in the Kadashan River, southeastern Alaska, USA. We first derived stand-level, non-climatic growth chronologies from riparian trees using standard dendroecology methods and differencing. When the chronologies were compared to 18-55 years of adult salmon escapement we detected positive, significant correlations at five of the nine sites. Regression models relating escapement to tree-ring growth at the five sites were applied to the differenced chronologies to reconstruct salmon abundance. Each reconstruction contains unique patterns characteristic of the site and salmon species. Reconstructions were validated by comparison to local histories (e.g., construction of dams and salmon canneries) and regional fisheries data such as salmon landings and aerial surveys and the Pacific Decadal Oscillation climate index. The reconstructions capture lower-frequency cycles better than extremes and are most useful for determination and comparison of relative abundance, cycles, and the effects of interventions. Reconstructions show lower population cycle maxima in both Umpqua River chinook and Sashin Creek pink salmon in recent decades. The Drinkwater Creek reconstruction suggests that sockeye abundance since the mid-1990s has been 15-25% higher than at any time since 1850, while no long-term deviations from natural cycles are detected for salmon in the Kadashan River or in Disappearance Creek. Decadal-scale cycles in salmon abundance with periods of 25-68 years were detected in all of the reconstructions. This novel approach provides river-specific, long-term perspectives on salmon abundance and cycles. Additionally, it provides a new frame of reference for maintaining and rebuilding individual stocks and for striking a balance between societal demands and the limited, always-changing salmon resource.     

Effects of brackish water incursions and diel phasing of tides on vertical excursions of the keystone predator Pisaster ochraceus

The physical factors that constrain the vertical foraging excursions of the keystone predator, the sea star Pisaster ochraceus, hold considerable interest because they indirectly shape the vivid patterns of zonation of rocky shore communities by impeding or enhancing the ability of P. ochraceus to traverse the intertidal zone. In this paper, we describe a study conducted in the Pacific Northwest of North America in which we examined, in the field and laboratory, the abiotic factors that can affect vertical excursions by P. ochraceus. Our field observations revealed that the extreme upward reach and average shore level height reached by P. ochraceus were significantly lower for daylight high tides than nocturnal high tides. Based on diver observations following a severe storm, it would also appear that these diurnal movements can be impeded by freshwater incursions into the intertidal zone; a regularly occurring event in the Pacific Northwest. As part of an experimental investigation into this phenomenon, we observed that sea stars maintained in tall cylindrical aquaria, without tidal flux, remained near the bottom during daylight and moved to the top of the column at night, suggesting that photoperiod alone can influence the cycle of vertical movement. Adding a freshwater layer to the aquaria restricted these vertical excursions. Our results suggest that on rocky coastlines susceptible to fresh water incursions, the suppression of foraging may be an important factor in the spatial and temporal variation in the intensity of predation. Furthermore, given the relative increase in frequency and intensity of freshwater incursions in the Pacific Northwest and the intolerance of P. ochraceus to lowered salinity, there is the long-term potential to significantly alter patterns of species zonation in this essential marine habitat.     

Migration and habitat of white sharks (<Emphasis Type="Italic">Carcharodon carcharias</Emphasis>) in the eastern Pacific Ocean

Satellite telemetry studies of 20 adult and sub-adult white sharks (360–530 cm estimated total length (TL)) in the eastern North Pacific during 1999–2005 revealed long distance seasonal migrations from the coast of California to an offshore focal area 2,500 km west of the Baja Peninsula, as well as the Hawaii Islands. Three tags were recovered allowing detailed behavioral analyses, including one shark’s migration cycle from the coast to the offshore focal area and back. While near pinniped rookeries in autumn and winter, sharks avoided the surface and used water to 50 m depth, consistent with a silhouette-based hunting strategy. Offshore migrations were initiated during November–March and followed periods of decreasing pinniped abundance. Migrations were highly directed, taking 23 ± 5 days to reach the offshore focal area along similar paths among sharks and years, defining a migration corridor. Sharks exhibited a broad depth distribution (0–644 m) in the offshore focal area, and remained there for up to 167 days during spring and summer, though primary productivity and fishery data suggest that forage resources are scarcer there than in other regions of the eastern North Pacific. Archival data from one shark revealed intensive oscillatory movements while in the offshore focal area, a behavior that may be related to foraging or mating. Sharks traveling to Hawaii remained near the islands up to 122 days, potentially feeding on pelagic fishes and marine mammals that concentrate around the islands. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Isotopes in teeth and a cryptic population of coastal freshwater seals

Human activities threaten the biodiversity of aquatic mammals across the globe. Conservation of these species hinges on the ability to delineate movements and foraging behaviors of animals, but gaining such insights is hampered by difficulties in tracing individuals over their lives. We determined isotope ratios in teeth (⁸⁷Sr/⁸⁶Sr, ¹³C/¹²C, and ¹⁸O/¹⁶O) to examine lifelong movement and resource-use patterns of a unique freshwater population of a wide-ranging pinniped species (harbor seal [Phoca vitulina]) that resides in Iliamna Lake, Alaska (U.S.A.). This population's potentially unique migratory behavior and use of different trophic resources are unknown. The isotope ratios we measured in teeth showed that seals were born in the lake, remained lifelong residents, and relied principally on resources produced from in the lake, even when seasonally abundant and nutrient-dense spawning anadromous fish (i.e., sockeye salmon [Oncorhynchus nerka]) were available in the lake. Our results illustrate how serial isotope records in teeth, particularly ⁸⁷Sr/⁸⁶Sr ratios, can be used to quantify how coastal mammal populations exploit both freshwater and marine ecosystems. Understanding lifelong patterns of habitat and resource use is essential information when designing effective conservation plans for threatened coastal mammals. We present the Iliamna Lake harbor seals as a unique case study into how isotope records within teeth can help reveal the cryptic ecology of such a population residing in an intact ecosystem. The results also provide critical baseline information for the Kvichak River system, which is facing an uncertain future due to proposed large-scale industrial development and a rapidly changing climate.