Prediction and confirmation of seasonal migration of Pacific sardine (Sardinops sagax) in the California Current Ecosystem

During the last century, the population of Pacific sardine (Sardinops sagax) in the California Current Ecosystem has exhibited large fluctuations in abundance and migration behavior. From approximately 1900 to 1940, the abundance of sardine reached 3.6 million metric tons and the “northern stock” migrated from offshore of California in the spring to the coastal areas near Oregon, Washington, and Vancouver Island in the summer. In the 1940s, the sardine stock collapsed and the few remaining sardine schools concentrated in the coastal region off southern California, year-round, for the next 50 years. The stock gradually recovered in the late 1980s and resumed its seasonal migration between regions off southern California and Canada. Recently, a model was developed which predicts the potential habitat for the northern stock of Pacific sardine and its seasonal dynamics. The habitat predictions were successfully validated using data from sardine surveys using the daily egg production method; scientific trawl surveys off the Columbia River mouth; and commercial sardine landings off Oregon, Washington, and Vancouver Island. Here, the predictions of the potential habitat and seasonal migration of the northern stock of sardine are validated using data from “acoustic–trawl” surveys of the entire west coast of the United States during the spring and summer of 2008. The estimates of sardine biomass and lengths from the two surveys are not significantly different between spring and summer, indicating that they are representative of the entire stock. The results also confirm that the model of potential sardine habitat can be used to optimally apply survey effort and thus minimize random and systematic sampling error in the biomass estimates. Furthermore, the acoustic–trawl survey data are useful to estimate concurrently the distributions and abundances of other pelagic fishes.

Distributions and abundances of Pacific sardine (Sardinops sagax) and other pelagic fishes in the California Current Ecosystem during spring 2006, 2008, and 2010, estimated from acoustic–trawl surveys

The abundances and distributions of coastal pelagic fish species in the California Current Ecosystem from San Diego to southern Vancouver Island, were estimated from combined acoustic and trawl surveys conducted in the spring of 2006, 2008, and 2010. Pacific sardine (Sardinops sagax), jack mackerel (Trachurus symmetricus), and Pacific mackerel (Scomber japonicus) were the dominant coastal pelagic fish species, in that order. Northern anchovy (Engraulis mordax) and Pacific herring (Clupea pallasii) were sampled only sporadically and therefore estimates for these species were unreliable. The estimates of sardine biomass compared well with those of the annual assessments and confirmed a declining trajectory of the “northern stock” since 2006. During the sampling period, the biomass of jack mackerel was stable or increasing, and that of Pacific mackerel was low and variable. The uncertainties in these estimates are mostly the result of spatial patchiness which increased from sardine to mackerels to anchovy and herring. Future surveys of coastal pelagic fish species in the California Current Ecosystem should benefit from adaptive sampling based on modeled habitat; increased echosounder and trawl sampling, particularly for the most patchy and nearshore species; and directed-trawl sampling for improved species identification and estimations of their acoustic target stren

Food habits of Steller sea lions (Eumetopias jubatus) off Oregon and northern California, 1986–2007

We described the diet of the eastern stock of Steller sea lions (Eumetopias jubatus) from 1416 scat samples collected from five sites in Oregon and northern California from 1986 through 2007. A total of 47 prey types from 30 families were identified. The most common prey was Pacific hake (Merluccius productus), followed by salmonids (Oncorhynchus spp.), skates (Rajidae), Pacific lamprey (Lampetra tridentata), herrings (Clupeidae), rockfish (Sebastes spp.), and northern anchovy (Engraulis mordax). Steller sea lion diet composition varied seasonally, annually, and spatially. Hake and salmonids were the most commonly identified prey in scats collected during the summer (breeding season), whereas hake and skate were most common in the nonbreeding season. Continued research on Steller sea lion diet and foraging behavior in the southern extent of their range is necessary to address issues such as climate change, interaction with competing California sea lions, and predation impacts on valuable or sensitive fish stocks.

Regional and seasonal patterns of epipelagic fish assemblages from the central California Current

The coastal Pacific Ocean off northern and central California encompasses the strongest seasonal upwelling zone in the California Current ecosystem. Headlands and bays here generate complex circulation features and confer unusual oceanographic complexity. We sampled the coastal epipelagic fish community of this region with a surface trawl in the summer and fall of 2000–05 to assess patterns of spatial and temporal community structure. Fifty-three species of fish were captured in 218 hauls at 34 fixed stations, with clupeiform species dominating. To examine spatial patterns, samples were grouped by location relative to a prominent headland at Point Reyes and the resulting two regions, north coast and Gulf of the Farallones, were plotted by using nonmetric multidimensional scaling. Seasonal and interannual patterns were also examined, and representative species were identified for each distinct community. Seven oceanographic variables measured concurrently with trawling were plotted by principal components analysis and tested for correlation with biotic patterns. We found significant differences in community structure by region, year, and season, but no interaction among main effects. Significant differences in oceanographic conditions mirrored the biotic patterns, and a match between biotic and hydrographic structure was detected. Dissimilarity between assemblages was mostly the result of differences in abundance and frequency of occurrence of about twelve common species. Community patterns were best described by a subset of hydrographic variables, including water depth, distance from shore, and any one of several correlated variables associated with upwelling intensity. Rather than discrete communities with clear borders and distinct member species, we found gradients in community structure and identified stations with similar fish communities by region and by proximity to features such as the San Francisco Bay.

Influence of ocean conditions on the timing of early life history events for blue rockfish (Sebastes mystinus) off California

Settled juvenile blue rockfish (Sebastes mystinus) were collected from two kelp beds approximately 335 km apart off Mendocino in northern California and Monterey in central California. A total of 112 rockfish were collected from both sites over 5 years (1993, 1994, 2001, 2002, and 2003). Total age, settlement date, age at settlement, and birth date were determined from otolith microstructure. Fish off Mendocino settled mostly in June and fish off Monterey settled mostly in May (average difference in settlement=23 days). Although the difference in the timing of settlement followed this same pattern for both areas over the five years, settlement occurred later in 2002 and 2003 than in the prior years of sampling. The difference in the timing of settlement was due primarily to differences in birth dates for the two areas. The time of settlement was positively related to upwelling and negatively related to sea level anomaly for most of the months before settlement. Knowledge of the timing of settlement has implications for design and placement of marine protected areas because protection of nursery grounds is frequently a major objective of these protected areas. The timing of settlement is also an important consideration in the planning of surveys of early recruits because mistimed surveys (caused by latitudinal differences in the timing of settlement) could produce biased estimates.

Larval development of the spotfin tonguefish (Symphurus oligomerus Mahadeva and Munroe, 1990) (Pleuronectiformes: Cynoglossidae) from the Gulf of California, Mexico

Larval and early juvenile stages of Symphurus oligomerus are described from 24 specimens from the Gulf of California. Meristic features were 48 – 49 total vertebrae, 87–94 dorsal-fin rays, 73–77 anal-fin rays, 12 caudal-fin rays, and five hypural bones. Seven larvae and one juvenile were cleared and stained to obtain the pterygiophore formula (1-3-2-2-2) that confirmed the identification of S. oligomerus. The pigment pattern from preflexion to juvenile stage consists of three bands on the dorsal margin and two bands on the ventral margin formed by star-shaped melanophores on the left side of the body. The intestine in preflexion to postflexion larvae forms an abdominal projection that ends in a short conical appendix. The intestine is supported by three cartilaginous struts; larvae with these physical attributes are called exterilium larvae. Preflexion larvae have two elongated dorsal-fin rays, and in flexion to postflexion larvae the second to the fourth dorsalfin rays are elongate. We found an apparent connection between the size at metamorphosis of the species of Symphurus and the depth distribution range of adults such that the fish species that metamorphose at a larger size have a deeper distribution as adults and exterilium larvae seem to correspond to species that have deeper distributions.

Assessment of habitat quality for juvenile California halibut (Paralichthys californicus) in a seasonally arid estuary

We evaluated habitat quality for juvenile California halibut (Paralichthys californicus) in a Pacific Coast estuary lacking in strong salinity gradients by examining density, recent otolith growth rates, and gut fullness levels of wild-caught and caged juveniles for one year. Juveniles <200 mm standard length were caught consistently in the inner, central, and outer sections of the estuary. The density of juveniles was two times higher in the inner estuary during most of the year, consistent with active habitat selection by part of the population. A generalized linear model indicated temperature, sampling time, and the interaction between salinity and temperature were significantly related to density. However, the model explained only 21% of the variance. Gut fullness levels of wild-caught juveniles were highest during the summer, but recent otolith growth rates were not related to temperature. The proportion of individuals feeding successfully indicated that seasonal differences in food availability are more important than spatial variation in prey abundance in driving feeding success. Feeding success of caged fishes was limited, precluding the use of growth rates as indicators of local habitat quality. However, marginal increment widths were reliable indicators of somatic growth at low growth rates over two-week periods. The relatively high growth rates and abundance of small wild-caught juveniles found throughout the estuary indicates that the entire estuary system has the potential for serving as nursery habitat.

All their eggs in one basket: a rocky reef nursery for the longnose skate (Raja rhina Jordan & Gilbert, 1880) in the southern California Bight

Skates (family Rajidae) are oviparous and lay tough, thick-walled eggs. At least some skate species lay their eggs in spatially restricted nursery grounds where embryos develop and hatch (Hitz, 1964; Hoff, 2007). After hatching, neonates may quickly leave the nursery grounds (Hoff, 2007). Egg densities in these small areas may be quite high. As an example, in the eastern Bering Sea, a site <2 km2 harbored eggs of Alaska skate (Bathyraja parmifera) exceeding 500,000/km2. All skate nursery grounds have been identified over soft sea floors (Lucifora and García, 2004; Hoff, 2007).

Abundance and population density of cetaceans in the California Current ecosystem

The abundance and population density of cetaceans along the U.S. west coast were estimated from ship surveys conducted in the summer and fall of 1991, 1993, 1996, 2001, and 2005 by using multiple-covariate, line-transect analyses. Overall, approximately 556,000 cetaceans of 21 species were estimated to be in the 1,141,800-km2 study area. Delphinoids (Delphinidae and Phocoenidae), the most abundant group, numbered ~540,000 individuals. Abundance in other taxonomic groups included ~5800 baleen whales (Mysticeti), ~7000 beaked whales (Ziphiidae), and ~3200 sperm whales (Physeteridae). This study provides the longest time series of abundance estimates that includes all the cetacean species in any marine ecosystem. These estimates will be used to interpret the impacts of human-caused mortality (such as that documented in fishery bycatch and that caused by ship strikes and other means) and to evaluate the ecological role of cetaceans in the California Current ecosystem.

Diets of and trophic relationships among dominant marine nekton within the northern California Current ecosystem

In this study we analyzed the diets of 26 nekton species collected from two years (2000 and 2002) off Oregon and northern California to describe dominant nekton trophic groups of the northern California Current (NCC) pelagic ecosystem. We also examined interannual variation in the diets of three nekton species. Cluster analysis of predator diets resulted in nekton trophic groups based on the consumption of copepods, euphausiids, brachyuran larvae, larval juvenile fishes, and adult nekton. However, many fish within trophic groups consumed prey from multiple trophic levels—euphausiids being the most widely consumed. Comparison of diets between years showed that most variation occurred with changes in the contribution of euphausiids and brachyuran larvae to nekton diets. The importance of euphausiids and other crustacean prey to nekton indicates that omnivory is an important characteristic of the NCC food web; however it may change during periods of lower or higher upwelling and ecosystem production.

Abundance, distribution, and habitat of leatherback turtles (Dermochelys coriacea) off California, 1990−2003

Leatherback turtles (Dermochelys coriacea) are regularly seen off the U.S. West Coast, where they forage on jellyfish (Scyphomedusae) during summer and fall. Aerial line-transect surveys were conducted in neritic waters (<92 m depth) off central and northern California during 1990−2003, providing the first foraging population estimates for Pacific leatherback turtles. Males and females of about 1.1 to 2.1 m length were observed. Estimated abundance was linked to the Northern Oscillation Index and ranged from 12 (coefficient of variation [CV] =0.75) in 1995 to 379 (CV= 0.23) in 1990, averaging 178 (CV= 0.15). Greatest densities were found off central California, where oceanographic retention areas or upwelling shadows created favorable habitat for leatherback turtle prey. Results from independent telemetry studies have linked leatherback turtles off the U.S. West Coast to one of the two largest remaining Pacific breeding populations, at Jamursba Medi, Indonesia. Nearshore waters off California thus represent an important foraging region for the critically endangered Pacific leatherback turtle.

Multiscale habitat associations of deepwater demersal fishes off central California

Fish-habitat associations were examined at three spatial scales in Monterey Bay, California, to determine how benthic habitats and landscape configuration have structured deepwater demersal fish assemblages. Fish counts and habitat variables were quantified by using observer and video data collected from a submersible. Fish responded to benthic habitats at scales ranging from cm’s to km’s. At broad-scales (km’s), habitat strata classified from acoustic maps were a strong predictor of fish assemblage composition. At intermediate-scales (m’s−100 m’s), fish species were associated with specific substratum patch types. At fine-scales (<1 m), microhabitat associations revealed differing degrees of microhabitat specificity, and for some species revealed niche separation within patches. The use of habitat characteristics in ecosystembased management, particularly as a surrogate for species distributions, will depend on resolving fish-habitat associations and habitat complexity over multiple scales.

Relationship between abundance of juvenile rockfishes (Sebastes spp.) and environmental variables documented off northern California and potential mechanisms for the covariation

We estimated annual abundance of juvenile blue (Sebastes mystinus), yellowtail (S. f lavidus), and black (S. melanops) rockfish off northern California over 21 years and evaluated the relationship of abundance to oceanographic variables (sea level anomaly, nearshore temperature, and offshore Ekman transport). Although mean annual abundance was highly variable (0.01−181 fish/minute), trends were similar for the three species. Sea level anomaly and nearshore temperature had the strongest relationship with interannual variation in rockfish abundance, and offshore Ekman transport did not correlate with abundance. Oceanographic events occurring in February and March (i.e., during the larval stage) had the strongest relationship with juvenile abundance, which indicates that year-class strength is determined during the larval stage. Also of note, the annual abundance of juvenile yellowtail rockfish was positively correlated with year-class strength of adult yellowtail rockfish; this finding would indicate the importance of studying juvenile abundance surveys for management purposes.

The relationships between fish assemblages and the amount of bottom horizontal beam exposed at California oil platforms: fish habitat preferences at man-made platforms and (by inference) at natural reefs

Between 1995 and 2002, we surveyed fish assemblages at seven oil platforms off southern and central California using the manned research submersible Delta. At each platform, there is a large horizontal beam situated at or near the sea floor. In some instances, shells and sediment have buried this beam and in other instances it is partially or completely exposed. We found that fish species responded in various ways to the amount of exposure of the beam. A few species, such as blackeye goby (Rhinogobiops nicholsii), greenstriped rockfish (Sebastes elongatus), and pink seaperch (Zalembius rosaceus) tended to avoid the beam. However, many species that typically associate with natural rocky outcrops, such as bocaccio (S. paucispinis), cowcod (S. levis), copper (S. caurinus), greenblotched (S. rosenblatti), pinkrose (S. simulator) and vermilion (S. miniatus) rockfishes, were found most often where the beam was exposed. In particular, a group of species (e.g., bocaccio, cowcod, blue (Sebastes mystinus), and vermilion rockfishes) called here the “sheltering habitat” guild, lived primarily where the beam was exposed and formed a crevice. This work demonstrates that the presence of sheltering sites is important in determining the species composition of man-made reefs and, likely, natural reefs. This research also indicates that adding structures that form sheltering sites in and around decommissioned platforms will likely lead to higher densities of many species typical of hard and complex structure.

Comparing the identification of southern California juvenile rockfishes (genus Sebastes spp.) by restriction site analysis of the mitochondrial ND3/ND4 region and by morphological characteristics

Rockfish (Sebastes spp.) juveniles are often difficult to identify by using morphological characters. This study independently applies morphological characters and a key based on mitochondrial restriction site variation to identify juvenile rockf ishes collected in southern California during juvenile rockfish surveys. Twenty-four specimens of Sebastes were examined genetically without knowledge of the morphological assignment. Seventeen fish were identified genetically as S. semicinctus, S. goodei, S. auriculatus, S. jordani, S. levis, S. rastrelliger, and S. saxicola. Identities for the remaining fish were narrowed to two or three species: 1) three fish were either S. carnatus or S. chrysomelas; 2) one fish was either S. chlorosticus, S. eos, or S. rosenblatti; and 3) three fish could have been either S. hopkinsi or S. ovalis, the latter for which we now have distinguishing mitochondrial markers. The genetic and morphological assignments concurred except for the identity of one fish that could only be narrowed down to S. hopkinsi or S. semicinctus by using morphological characters. Genetics excluded more species from multispecies groupings than did the morphological approach, especially species within the subgenus Sebastomus. Species in the genetically unresolvable groups may be similar because of recent divergence or because of interspecies introgression.