Examination of the foraging habits of Pacific harbor seal (Phoca vitulina richardsi) to describe their use of the Umpqua River, Oregon, and their predation on salmonids

The increase in harbor seal (Phoca vitulina richardsi) abundance, concurrent with the decrease in salmonid (Oncorhynchus spp.) and other fish stocks, raises concerns about the potential negative impact of seals on fish populations. Although harbor seals are found in rivers and estuaries, their presence is not necessarily indicative of exclusive or predominant feeding in these systems. We examined the diet of harbor seals in the Umpqua River, Oregon, during 1997 and 1998 to indirectly assess whether or not they were feeding in the river. Fish otoliths and other skeletal structures were recovered from 651 scats and used to identify seal prey. The use of all diagnostic prey structures, rather than just otoliths, increased our estimates of the number of taxa, the minimum number of individuals and percent frequency of occurrence (%FO) of prey consumed. The %FO indicated that the most common prey were pleuronectids, Pacific hake (Merluccius productus), Pacific stag-horn sculpin (Leptocottus armatus), osmerids, and shiner surfperch (Cymatogaster aggregata). The majority (76%) of prey were fish that inhabit marine waters exclusively and fish found in marine and estuarine areas (e.g. anadromous spp.) which would indicate that seals forage predominantly at sea and use the estuary for resting and opportunistic feeding. Salmonid remains were encountered in 39 samples (6%); two samples contained identifiable otoliths, which were determined to be from chi-nook salmon (O. tshawytscha). Because of the complex salmonid composition in the Umpqua River, we used molecular genetic techniques on salmonid bones retrieved from scat to discern species that were rare from those that were abundant. Of the 37 scats with salmonid bones but no otoliths, bones were identified genetically as chinook or coho (O. kisutch) salmon, or steelhead trout (O. mykiss) in 90% of the samples.

Diet shifts of juvenile red snapper (Lutjanus campechanus) with changes in habitat and fish size

We examined the diets and habitat shift of juvenile red snapper (Lutjanus campechanus) in the northeast Gulf of Mexico. Fish were collected from open sand-mud habitat (little to no relief), and artificial reef habitat (1-m3 concrete or PVC blocks), from June 1993 through December 1994. In 1994, fish settled over open habitat from June to September, as shown by trawl collections, then began shifting to reef habitat — a shift that was almost completed by December as observed by SCUBA visual surveys. Stomachs were examined from 1639 red snapper that ranged in size from 18.0 to 280.0 mm SL. Of these, 850 fish had empty stomachs, and 346 fish from open habitat and 443 fish from reef habitat contained prey. Prey were identified to the lowest possible taxon and quantified by volumetric measurement. Specific volume of particular prey taxa were calculated by dividing prey volume by individual fish weight. Red snapper shifted diets with increasing size. Small red snapper (<60 mm SL) fed mostly on chaetognaths, copepods, shrimp, and squid. Large red snapper (60–280 mm SL) shifted feeding to fish prey, greater amounts of squid and crabs, and continued feeding on shrimp. We compared red snapper diets for overlapping size classes (70–160 mm SL) of fish that were collected from both habitats (Bray-Curtis dissimilarity index and multidimensional scaling analysis). Red snapper diets separated by habitat type rather than fish size for the size ranges that overlapped habitats. These diet shifts were attributed to feeding more on reef prey than on open-water prey. Thus, the shift in habitat shown by juvenile red snapper was reflected in their diet and suggested differential habitat values based not just on predation refuge but food resources as well.

Diet changes of Pacific cod (Gadus macrocephalus) in Alaska between 1980 and 1995

The diet of Pacific cod (Gadus macrocephalus) in the area of Pavlof Bay, Alaska, was studied in the early 1980s by Albers and Anderson (1985). They found that the dominant prey species were forage species like pandalid shrimp, capelin (Mallotus villosus), and walleye pollock (Theragra chalcogramma). The shrimp fishery in Pavlof Bay began in 1968 and closed in 1980 because of low shrimp abundance (Ruccio and Worton1). Survey data indicate that, during the period between 1972 and 1997, the abundance of forage species such as pandalid shrimp and capelin declined and higher trophic-level groundfish such as Pacific cod increased. There is a general recognition that a long-term ocean climate shift in the Gulf of Alaska has been partially responsible for the observed reorganization of the community structure (Anderson and Piatt, 1999).

A method to improve size estimates of walleye pollock (Theragra chalcogramma) Atka mackerel (Pleurogrammus monopterygius) consumed by pinnipeds: digestion correction factors applied to bones and otoliths recovered in scats

The lengths of otoliths and other skeletal structures recovered from the scats of pinnipeds, such as Steller sea lions (Eumetopias jubatus), correlate with body size and can be used to estimate the length of prey consumed. Unfortunately, otoliths are often found in too few scats or are too digested to usefully estimate prey size. Alternative diagnostic bones are frequently recovered, but few bone-size to prey-size correlations exist and bones are also reduced in size by various degrees owing to digestion. To prevent underestimates in prey sizes consumed techniques are required to account for the degree of digestion of alternative bones prior to estimating prey size. We developed a method (using defined criteria and photo-reference material) to assign the degree of digestion for key cranial structures of two prey species: walleye pollock (Theragra chalcogramma) and Atka mackerel (Pleurogrammus monopterygius). The method grades each structure into one of three condition categories; good, fair or poor. We also conducted feeding trials with captive Steller sea lions, feeding both fish species to determine the extent of erosion of each structure and to derive condition-specific digestion correction factors to reconstruct the original sizes of the structures consumed. In general, larger structures were relatively more digested than smaller ones. Mean size reduction varied between different types of structures (3.3−26.3%), but was not influenced by the size of the prey consumed. Results from the observations and experiments were combined to be able to reconstruct the size of prey consumed by sea lions and other pinnipeds. The proposed method has four steps: 1) measure the recovered structures and grade the extent of digestion by using defined criteria and photo-reference collection; 2) exclude structures graded in poor condition; 3) multiply measurements of structures in good and fair condition by their appropriate digestion correction factors to derive their original size; and 4) calculate the size of prey from allometric regressions relating corrected structure measurements to body lengths. This technique can be readily applied to piscivore dietary studies that use hard remains of fish.

Sizes of walleye pollock (Theragra chalcogramma) consumed by the eastern stock of Steller sea lions (Eumetopias jubatus) in Southeast Alaska from 1994 to 1999

Lengths of walleye pollock (Theragra chalcogramma) consumed by Steller sea lions (Eumetopias jubatus) were estimated by using allometric regressions applied to seven diagnostic cranial structures recovered from 531 scats collected in Southeast Alaska between 1994 and 1999. Only elements in good and fair condition were selected. Selected structural measurements were corrected for loss of size due to erosion by using experimentally derived condition-specific digestion correction factors. Correcting for digestion increased the estimated length of fish consumed by 23%, and the average mass of fish consumed by 88%. Mean corrected fork length (FL) of pollock consumed was 42.4 ±11.6 cm (range=10.0−78.1 cm, n=909). Adult pollock (FL>45.0 cm) occurred more frequently in scats collected from rookeries along the open ocean coastline of Southeast Alaska during June and July (74% adults, mean FL=48.4 cm) than they did in scats from haul-outs located in inside waters between October and May (51% adults, mean FL=38.4 cm). Overall, the contribution of juvenile pollock (≤20 cm) to the sea lion diet was insignificant; whereas adults contributed 44% to the diet by number and 74% by mass. On average, larger pollock were eaten in summer at rookeries throughout Southeast Alaska than at rookeries in the Gulf of Alaska and the Bering Sea. Overall it appears that Steller sea lions are capable of consuming a wide size range of pollock, and the bulk of fish fall between 20 and 60 cm. The use of cranial hard parts other than otoliths and the application of digestion correction factors are fundamental to correctly estimating the sizes of prey consumed by sea lions and determining the extent that these sizes overlap with the sizes of pollock caught by commercial fisheries.

The Hudson-Raritan Estuary as a crossroads for distribution of blue (Callinectes sapidus), lady (Ovalipes ocellatus), and Atlantic rock (Cancer irroratus) crabs

Blue (Callinectes sapidus)(Portunidae),lady (Ovalipes ocellatus)(Portunidae), and Atlantic rock (Cancer irroratus) (Cancridae) crabs inhabit estuaries on the northeast United States coast for parts or all of their life cycles. Their distributions overlap or cross during certain seasons. During a 1991–1994 monthly otter trawl survey in the Hudson-Raritan Estuary between New York and New Jersey, blue and lady crabs were collected in warmer months and Atlantic rock crabs in colder months. Sex ratios, male:female, of mature crabs were 1:2.0 for blue crabs, 1:3.1 for lady crabs, and 21.4:1 for Atlantic rock crabs. Crabs, 1286 in total, were subsampled for dietary analysis, and the dominant prey taxa for all crabs, by volume of foregut contents, were mollusks and crustaceans. The proportion of amphipods and shrimp in diets decreased as crab size increased. Trophic niche breadth was widest for blue crabs, narrower for lady crabs, and narrowest for Atlantic rock crabs. Trophic overlap was lowest between lady crabs and Atlantic rock crabs, mainly because of frequent consumption of the dwarf surfclam (Mulinia lateralis) by the former and the blue mussel (Mytilus edulis) by the latter. The result of cluster analysis showed that size class and location of capture of predators in the estuary were more influential on diet than the species or sex of the predators.

Diet of the minimal armhook squid (Berryteuthis anonychus) (Cephalopoda: Gonatidae) in the northeast Pacific during spring

The stomach contents of the minimal armhook squid (Berryteuthis anonychus) were examined for 338 specimens captured in the northeast Pacific during May 1999. The specimens were collected at seven stations between 145−165°W and 39−49°N and ranged in mantle length from 10.3 to 102.2 mm. Their diet comprised seven major prey groups (copepods, chaetognaths, amphipods, euphausiids, ostracods, unidentified fish, and unidentified gelatinous prey) and was dominated by copepods and chaetognaths. Copepod prey comprised four genera, and 86% by number of the copepods were from the genus Neocalanus. Neocalanus cristatus was the most abundant prey taxa, composing 50% by mass and 35% by number of the total diet. Parasagitta elegans (Chaetognatha) occurred in more stomachs (47%) than any other prey taxon. Amphipods occurred in 19% of the stomachs but composed only 5% by number and 3% by mass of the total prey consumed. The four remaining prey groups (euphausiids, ostracods, unidentified fish, and unidentified gelatinous prey) together composed <2% by mass and <1% by number of the diet. There was no major change in the diet through the size range of squid examined and no evidence of cannibalism or predation on other cephalopod species.

Diet composition of large striped bass (Morone saxatilis) in Chesapeake Bay

Large (>458 mm) striped bass (Morone saxatilis) are dominant predators in Chesapeake Bay. In recent years, the Chesapeake Bay stock of striped bass has increased dramatically, raising concerns about their predatory impact and their forage requirements. In response to these concerns and the need for more recent ecological studies, this investigation was conducted to characterize feeding habits of large striped bass in Chesapeake Bay. Stomach contents from 1225 striped bass from 458 to 1151 mm TL were examined in the spring and fall of 1997 and 1998. Striped bass consumed 52 different species of vertebrates and invertebrates; however, only a few species of clupeoid and sciaenid fishes dominated diets across both the seasons and size ranges of striped bass examined. Of finfish species, menhaden (Brevoortia tyrannus) was the dominant prey in most areas and gizzard shad (Dorosoma cepedianum) replaced menhaden in importance in lower salinity waters. Spot (Leiostomus xanthurus) and other sciaenid fishes and anadromous herrings (Alosa spp.) also contibuted large percentages of striped bass diet. Although pelagic schooling fishes formed the majority of the diet, benthic fishes contributed a higher percentage to the diet than in previous studies of striped bass diet composition.

Diving behavior of immature Steller sea lions (Eumetopias jubatus)

Understanding the ontogenetic relationship between juvenile Steller sea lions (Eumetopias jubatus) and their foraging habitat is key to understanding their relationship to available prey and ultimately their survival. We summarize dive and movement data from 13 young-of-the-year (YOY) and 12 yearling Steller sea lions equipped with satellite dive recorders in the Gulf of Alaska and Aleutian Islands (n=18), and Washington (n=7) from 1994 to 2000. A total of 1413 d of transmission (x =56.5 d, range: 14.5–104.1 d) were received. We recorded 222,073 dives, which had a mean depth of 18.4 m (range of means: 5.8−67.9 m; SD=16.4). Alaska YOY dived for shorter periods and at shallower depths (mean depth=7.7 m, mean duration=0.8 min, mean maximum depth=25.7 m, and maximum depth=252 m) than Alaska yearlings (x =16.6 m, 0=1.1 min, x = 63.4 m, 288 m), whereas Washington yearlings dived the longest and deepest (mean depth=39.4 m, mean duration=1.8 min, mean maximum depth=144.5 m, and maximum depth=328 m). Mean distance for 564 measured trips was 16.6 km; for sea lions ≤10 months of age, trip distance (7.0 km) was significantly less than for those >10 months of age (24.6 km). Mean trip duration for 10 of the 25 sea lions was 12.1 h; for sea lions ≤10 months of age, trip duration was 7.5 h and 18.1 h for those >10 months of age. We identified three movements types: long-range trips (>15 km and >20 h), short-range trips (<15 km and <20 h) during which the animals left and returned to the same site, and transits to other haul-out sites. Long-range trips started around 9 months of age and occurred most frequently around the assumed time of weaning, whereas short-range trips happened almost daily (0.9 trips/day, n=426 trips). Transits began as early as 7 months of age, occurred more often after 9 months of age, and ranged between 6.5 and 454 km. The change in dive characteristics coincided with the assumed onset of weaning. These yearling sea lion movement patterns and dive characteristics suggest that immature Steller sea lions are as capable of making the same types of movements as adults.

Diets of thornback ray (Raja clavata) and tope shark (Galeorhinus galeus) in the bottom longline fishery of the Azores, northeastern Atlantic

Tope shark (Galeorhinus galeus) and thornback ray (Raja clavata) are the two most captured elasmobranch species by the Azorean bottom longline fishery. In order to better understand the trophic dynamics of these species in the Azores, the diets of thornback ray and tope shark caught in this area during 1996 and 1997 were analyzed to describe feeding patterns and to investigate the effect of sex, size, and depth and area of capture on diet. Thornback rays fed mainly upon fishes and reptants, but also upon polychaetes, mysids, natant crustaceans, isopods, and cephalopods. In the Azores, this species preyed more heavily upon fish compared with the predation patterns described in other areas. Differences in the diet may be due to differences in the environments (e.g. in the Azores, seamounts and oceanic islands are the major topographic features, whereas in all other studies, continental shelves have been the major topographic feature). No differences were observed in the major prey consumed between the sexes or between size classes (49−60, 61−70, 71−80, and 81−93 cm TL). Our study indicates that rays inhabiting different depths and areas (coastal or offshore banks) prey upon different resources. This appears to be related to the relative abundance of prey with habitat. Tope sharks were found to prey almost exclusively upon teleost fish: small shoaling fish, mainly boarfish (Capros aper) and snipefish (Macroramphosus scolopax), were the most frequent prey. This study illustrates that thornback rays and tope sharks are top predators in waters off the Azores.

Physiological ecology of juvenile chinook salmon (Oncorhynchus tshawytscha) at the southern end of their distribution, the San Francisco Estuary and Gulf of the Farallones, California

Juvenile chinook salmon, Oncorhynchus tshawytscha, from natal streams in California’s Central Valley demonstrated little estuarine dependency but grew rapidly once in coastal waters. We collected juvenile chinook salmon at locations spanning the San Francisco Estuary from the western side of the freshwater delta—at the confluence of the Sacramento and San Joaquin Rivers—to the estuary exit at the Golden Gate and in the coastal waters of the Gulf of the Farallones. Juveniles spent about 40 d migrating through the estuary at an estimated rate of 1.6 km/d or faster during their migration season (May and June 1997) toward the ocean. Mean growth in length (0.18 mm/d) and weight (0.02 g/d) was insignificant in young chinook salmon while in the estuary, but estimated daily growth of 0.6 mm/d and 0.5 g/d in the ocean was rapid (P≤0.001). Condition (K factor) declined in the estuary, but improved markedly in ocean fish. Total body protein, total lipid, triacylglycerols (TAG), polar lipids, cholesterol, and nonesterified fatty acids concentrations did not change in juveniles in the estuary, but total lipid and TAG were depleted in ocean juveniles. As young chinook migrated from freshwater to the ocean, their prey changed progressively in importance from invertebrates to fish larvae. Once in coastal waters, juvenile salmon appear to employ a strategy of rapid growth at the expense of energy reserves to increase survival potential. In 1997, environmental conditions did not impede development: freshwater discharge was above average and water temperatures were only slightly elevated, within the species’ tolerance. Data suggest that chinook salmon from California’s Central Valley have evolved a strong ecological propensity for a ocean-type life history. But unlike populations in the Pacific Northwest, they show little estuarine dependency and proceed to the ocean to benefit from the upwelling-driven, biologically productive coastal waters.

Improving pinniped diet analyses through identification of multiple skeletal structures in fecal samples

Marine mammal diet is typically characterized by identifying fish otoliths and cephalopod beaks retrieved from stomachs and fecal material (scats). The use and applicability of these techniques has been the matter of some debate given inherent biases associated with the method. Recent attempts to identify prey using skeletal remains in addition to beaks and otoliths are an improvement; however, difficulties incorporating these data into quantitative analyses have limited results for descriptive analyses such as frequency of occurrence. We attempted to characterize harbor seal (Phoca vitulina) diet in an area where seals co-occur with several salmon species, some endangered and all managed by state or federal agencies, or both. Although diet was extremely variable within sampling date, season, year, and between years, the frequency and number of individual prey were at least two times greater for most taxa when prey structures in addition to otoliths were identified. Estimating prey mass in addition to frequency and number resulted in an extremely different relative importance of prey in harbor seal diet. These data analyses are a necessary step in generating estimates of the size, total number, and annual biomass of a prey species eaten by pinnipeds for inclusion in fisheries management plans.

Pinniped diet composition: a comparison of estimation models

Along the west coast of the United States, the potential impact of increasing pinniped populations on declining salmonid (Oncorhynchus spp.) stocks has become an issue of concern. Fisheries managers need species-specific estimates of consumption by pinnipeds to evaluate their impact on salmonid stocks. To estimate consumption, we developed a model that estimates diet composition by reconstructing prey biomass from fecal samples. We applied the model to data collected from harbor seals (Phoca vitulina) that are present year-round in the lower Columbia River where endangered stocks of salmonids pass as returning adults and as seaward-migrating smolts. Using the same data, we applied the split-sample frequency of occurrence model, which avoids reconstructing biomass by assuming that each fecal sample represents an equal volume of consumption and that within each sample each prey item represents an equal proportion of the volume. The two models for estimating diet composition yielded size-specific differences in consumption estimates that were as large as tenfold for the smallest and largest prey. Conclusions about the impact of harbor seal predation on adult salmonids, some of their largest prey species, remain uncertain without some appropriate rationale or further information (e.g. empirical captive studies) to discriminate between these models.