Application of Acoustic Telemetry to Assess Residency and Movements of Rockfish and Lingcod at Created and Natural Habitats in Prince William Sound

Loss and/or degradation of nearshore habitats have led to increased efforts to restore or enhance many of these habitats, particularly those that are deemed essential for marine fishes. Copper rockfish (Sebastes caurinus) and lingcod (Ophiodon enlongatus) are dominant members of the typical reef fish community that inhabit rocky and high-relief substrates along the Pacific Northwest. We used acoustic telemetry to document their residency and movements in the nearshore waters of Prince William Sound, Alaska in order to assess use of created reef habitat in an individual-based manner. A total of 57 fish were surgically implanted with acoustic transmitters. Forty-five fish were captured and monitored in three habitats: artificial reef, low-relief natural reef, and patchy high-relief natural reef. Within each habitat, both rockfish and lingcod exhibited long periods of residency with limited movements. Twelve rockfish were captured at the natural reefs and displaced a distance of 4.0 km to the artificial reef. Five of the 12 rockfish returned within 10 d of their release to their initial capture site. Another five of the 12 displaced fish established residency at the artificial reef through the duration of our study. Our results suggest the potential for artificial reefs to provide rockfish habitat in the event of disturbances to natural habitat.     

Diel differences in fish assemblages in nearshore eelgrass and kelp habitats in Prince William Sound,Alaska

The importance of a particular habitat to nearshore fishes can be best assessed by both diurnal and nocturnal sampling. To determine diel differences in fish assemblages in nearshore eelgrass and understory kelp habitats, fishes were sampled diurnally and nocturnally at six locations in western Prince William Sound, Alaska, in summer 2007. Abundance of fish between day and night were similar, but species composition and mean size of some fish changed. Species richness and species diversity were similar in eelgrass during the day and night, whereas in kelp, species richness and species diversity were greater at night than during the day. In eelgrass, saffron cod (Eleginus gracilis) was the most abundant species during the day and night. In kelp, the most abundant species were Pacific herring (Clupea pallasii) during the day and saffron cod at night. Diel differences in fish size varied by species and habitat. Mean length of saffron cod was similar between day and night in eelgrass but was greatest during the day in kelp. Pacific herring were larger at night than during the day in kelp. Diel sampling is important to identity nearshore habitats essential to fish and help manage fish stocks at risk.     

Seasonal foraging movements and migratory patterns of female Lamna ditropis tagged in Prince William Sound, Alaska

Conventional and electronic tags were used to investigate social segregation, distribution, movements and migrations of salmon sharks Lamna ditropis in Prince William Sound, Alaska. Sixteen salmon sharks were tagged with satellite transmitters and 246 with conventional tags following capture, and were then released in Prince William Sound during summer 1999 to 2001. Most salmon sharks sexed during the study were female (95%), suggesting a high degree of sexual segregation in the region. Salmon sharks congregated at adult Pacific salmon Oncorhynchus spp. migration routes and in bays near Pacific salmon spawning grounds in Prince William Sound during July and August. Adult Pacific salmon were the principal prey in 51 salmon shark stomachs collected during summer months in Prince William Sound, but the fish appeared to be opportunistic predators and consumed sablefish Anoplopoma fimbria, gadids, Pacific herring Clupea pallasi, rockfish Sebastes spp. and squid (Teuthoidea) even when adult Pacific salmon were locally abundant. As Pacific salmon migrations declined in late summer, the salmon sharks dispersed; some continued to forage in Prince William Sound and the Gulf of Alaska into autumn and winter months, while others rapidly moved south‐east thousands of kilometres toward the west coasts of Canada and the U.S. Three movement modes are proposed to explain the movement patterns observed in the Gulf of Alaska and eastern North Pacific Ocean: ‘focal foraging’ movements, ‘foraging dispersals’ and ‘direct migrations’. Patterns of salmon shark movement are possibly explained by spatio‐temporal changes in prey quality and density, an energetic trade‐off between prey availability and water temperature, intra‐specific competition for food and reproductive success. Transmissions from the electronic tags also provided data on depth and water temperatures experienced by the salmon sharks. The fish ranged from the surface to a depth of 668 m, encountered water temperatures from 4·0 to 16·8° C and generally spent the most time above 40 m depth and between 6 and 14° C (60 and 73%, respectively).     

Production of the 57 kDa major surface antigen by a non-agglutinating strain of the fish pathogen Renibacterium salmoninarum

Both the prevalence and tissue titer of viral hemorrhagic septicemia virus (VHSV) increased in Pacific herring Clupea pallasi following their introduction into net pens (pounds) used in the closed pound spawn-on-kelp (SOK) fishery in Prince William Sound, Alaska. VHSV was also found in water samples from inside and outside the SOK pounds after herring had been confined for several days; however, water samples taken near wild free-ranging, spawning herring either failed to test positive or tested weakly positive for virus. Little or no virus was found in tissue samples from free-ranging, spawning herring captured from the vicinity of the pounds, nor did the prevalence of VHSV increase following spawning as it did in impounded herring. The data indicated that increased prevalences of VHSV were correlated with confinement of herring for the closed pound SOK fishery and that infection was spread within the pounds through waterborne exposure to virus particles originating from impounded fish. In addition, pounds containing predominantly young fish had higher prevalences of VHSV, suggesting that older fish may be partially immune, perhaps as a result of previous infection with the virus. Operation of SOK pounds during spawning seasons in which young herring predominate may amplify the disease and possibly exacerbate the population fluctuations observed in wild herring stocks.     

Limited realized dispersal and introgressive hybridization influence genetic structure and conservation strategies for brown rockfish, <Emphasis Type="Italic">Sebastes auriculatus</Emphasis>

Understanding patterns of connectivity among marine fish populations with demersal adults and pelagic larvae is critical for effective conservation of west coast rockfishes. The brown rockfish (Sebastes auriculatus) occurs in nearshore habitat and is common from northern Baja California, Mexico to northern California, rare off the outer coast of Oregon and Washington and again common in the inland waters of Puget Sound, Washington. Here we examine patterns of microsatellite DNA diversity from throughout the species’ range as an indirect measure of long-term trends in larval dispersal. Genetic divergence was large and highly significant over all populations (F _ST=0.056, P<0.0001), and was significantly correlated with geographic distance when considering coastal populations. The best estimates of mean coastal dispersal distance were on the order of 10 km or less per generation. Diversity was relatively low in the Puget Sound, suggesting that Puget Sound rockfish populations experienced a post-glacial founder effect followed by genetic isolation and low effective population size. Puget Sound individuals appeared to have recent mixed ancestry as a result of introgression with S. maliger and S. caurinus. Genetic isolation of Puget Sound fish provides a basis for consideration as a Distinct Population Segment (DPS) under the provisions of the Endangered Species Act. We recommend that coastal brown rockfish fisheries be managed at regional rather than coast-wide scales, and that design of marine reserve networks considers the surprisingly low realized dispersal distance of some species with high dispersal potential.     

Genetic structuring among Alaskan Pacific herring populations identified using microsatellite variation

Five highly variable microsatellite loci were used to investigate population structuring in Pacific herring Clupea pallasi collected from Kodiak Island, two sites in the Bering Sea and four sites within Prince William Sound, Alaska. All loci revealed high levels of variability with heterozygosity estimates ranging from 86 to 97% (mean heterozygosity: 89%). The variation was structured significantly among sites suggesting that the samples investigated were genetically distinct from each other. Genetic divergence was greatest between populations from the Bering Sea and those from Prince William Sound. The Kodiak Island and Point Chalmers samples appeared to be distinct from the Prince William Sound and Bering Sea populations. The observed genetic distance relationships among samples could be explained largely in terms of geographical separation.     

Northern pike (Esox lucius L.) and aquatic vegetation,tools in the management of fisheries and water quality in shallow waters

Species and size composition of fish communities in shallow stagnant waters appear to be associated with the type, abundance and pattern of the vegetation. Man-induced impacts as eutrophication, and suppression of vegetation for reasons of water quantity management or angling pleasure may induce irreversible changes in the aquatic ecosystem. Water quality management should aim at restoring former pike habitat. Submerged weeds are important to that effect. These habitats are characterized by relatively low densities of fish stocks. Stocking of bream and carp interferes strongly with these objectives. With respect to fish stocks, interests of water quality and fishery management may be opposed.     

Spatial and Temporal Variability in Juvenile Pacific Herring, Clupea pallasi, Growth in Prince William Sound, Alaska

We examined the spatial and temporal variability of juvenile Pacific herring, Clupea pallasi, growth within Prince William Sound, Alaska. Pacific herring, ranging from post-larval to mature fish, were collected from four spatially segregated bays between October 1995 and March 1998. Linear growth equations from each bay were similar. However, growth rates and wet weight-at-length, reflecting condition, of juvenile Pacific herring cohorts varied seasonally and annually. The short term spatial variability in juvenile Pacific herring growth suggested that each bay was a unique nursery area. The physical and biological conditions within each bay appeared to dictate Pacific herring growth rate.     

Cultivation of Macrocystis integrifolia (Laminariales,Phaeophyta) in southeastern Alaskan waters

The artificial culture of Macrocystis integrifolia in southeastern Alaskan waters has been initiated as a first step in a project to augment the existing herring roe-on-kelp fishery in the state. The growing of Macrocystis in Prince William Sound has the potential of considerably enhancing this fishery by eliminating the costly importation of this kelp from southeast Alaska. In an ongoing feasibility study, Macrocystis has been cultured under laboratory conditions through the embryonic sporophyte stage and outplanted in waters near Sitka, Alaska (57° N). Growth of the outplanted kelps has been monitored as a function of the time and depth of the outplanting. Preliminary results suggest that light is limiting for growth in winter and that the optimal depth for outplanting will vary with the season.     

Integrating fish and parasite data as a holistic solution for identifying the elusive stock structure of Pacific sardines (Sardinops sagax)

There is an urgent need to clarify how different stocks, or subpopulations of fish species, are vulnerable to fishing pressure and unfavorable ocean conditions because of the increasing demand on fisheries for human consumption. For marine fishes, the potential for high gene flow increases the difficulty in determining the number of subpopulations managed in a specific fishery. Although the use of molecular data has become a common method in the past 15 years to identify fish subpopulations, no single technique or suite of techniques has been established for fish stock structure studies. We review the use of fish morphometrics, artificial tags, fish genetics, parasite genetics, and parasites as biological tags to identify subpopulations of marine fishes with a focus on the Pacific sardine (Sardinops sagax) fishery off the west coast of North America. We suggest an integration of fish- and parasite-based techniques for future stock structure studies, particularly for pelagic fish species whose stock structure can be elusive. An integration of techniques may also resolve fish stock structure over small geographic areas by increasing the number of spatial and temporal scales studied simultaneously leading to methods for successful management of marine fish species.     

Hypotheses concerning the decline and poor recovery of Pacific herring in Prince William Sound, Alaska

This paper updates previous reviews of the 1993 stock decline of Pacific herring (Clupea pallasi) in Prince William Sound, Alaska, and focuses on hypotheses about subsequent poor recovery. Recent age structured assessment modeling with covariate analysis indicates that the population dynamics of the sound’s herring are influenced by oceanic factors, nutrition, and, most substantially, hatchery releases of juvenile pink salmon. For the 1993 decline, poor nutrition remains the most probable cause with disease a secondary response. Concerning poor recovery, we examined 16 potential factors and found three to be causal: oceanic factors, poor nutrition, and hatchery releases of juvenile pink salmon. Absences of strong year classes at both Sitka and Prince William Sound after 1993 indicate the action of large-scale ocean processes. Beyond regional-scale environmental factors, two factors specific to the sound influence the population dynamics of herring and are likely impeding recovery. First, pink salmon fry releases have increased to about 600 million annually and may disrupt feeding in young herring, which require adequate nutrition for growth and overwintering survival. Juvenile pink salmon and age-1 herring co-occur in nearshore areas of bays in late spring and summer, and available data on dietary overlap indicates potential competition between the age-1 juvenile herring and juvenile pink salmon. Field studies demonstrate that juvenile herring reduce food intake substantially in the presence of juvenile pink salmon. Second, overwintering humpback whales may consume potentially large amounts of adult herring, but further studies must confirm to what extent whale predation reduces herring biomass.     

Association of plasma IgM with body size, histopathologic changes, and plasma chemistries in adult Pacific herring Clupea pallasi

Pacific herring Clupea pallasi immunoglobulin is an IgM-like molecule comprised of heavy and light chains with molecular weights of 79 and 25 to 27 kD, respectively. Purified immunoglobulin was used to generate highly specific polyclonal antibodies for development of a sandwich ELISA. The ELISA was used to quantify total plasma IgM in 602 Pacific herring captured in Prince William Sound and Sitka Sound, Alaska, USA. Plasma IgM concentrations ranged from 0.13 to 5.32 mg ml-1. Using multiple stepwise regression analysis, plasma IgM was highly correlated (p < or = 0.01) with body length, Ichthyophonus hoferi infection, plasma albumin, plasma cholesterol, liver macrophage aggregates, and focal skin reddening. I. hoferi was the only organism significantly associated with plasma IgM. Gender, site, and season (spring vs fall) did not contribute to significant differences in plasma IgM. This study contributes to the understanding of the interaction of body size, plasma chemistries, and pathological changes upon circulating immunoglobulins in fish.     

Genetic population structure of the endemic fourline wrasse (Larabicus quadrilineatus) suggests limited larval dispersal distances in the Red Sea

The connectivity among marine populations is determined by the dispersal capabilities of adults as well as their eggs and larvae. Dispersal distances and directions have a profound effect on gene flow and genetic differentiation within species. Genetic homogeneity over large areas is a common feature of coral reef fishes and can reflect high dispersal capability resulting in high levels of gene flow. If fish larvae return to their parental reef, gene flow would be restricted and genetic differentiation could occur. Larabicus quadrilineatus (Labridae) is considered as an endemic fish species of the Red Sea and Gulf of Aden. The juveniles of this species are cleaner fish that feed on ectoparasites of other fishes. Here, we investigated the genetic population structure and gene flow in L. quadrilineatus among five locations in the Red Sea to infer connectivity among them. To estimate genetic diversity, we analysed 369 bp of 237 mitochondrial DNA control region sequences. Haplotype and nucleotide diversities were higher in the southern than in the northern Red Sea. Analysis of molecular variance (amova) detected the highest significant genetic variation between northern and central/southern populations (Phi(CT) = 0.01; P < 0.001). Migration analysis revealed a several fold higher northward than southward migration, which could be explained by oceanographic conditions and spawning season. Even though the Phi(ST) value of 0.01 is rather low and implies a long larval dispersal distance, estimates based on the isolation-by-distance model show a very low mean larval dispersal distance (0.44-5.1 km) compared to other studies. In order to enable a sustainable ornamental fishery on the fourline wrasse, the results of this study suggest that populations in the northern and southern Red Sea should be managed separately as two different stocks. The rather low larval dispersal distance of about 5 km needs to be considered in the design of marine protected areas to enable connectivity and self-seeding.     

FORAGING STRATEGIES OF SYMPATRIC KILLER WHALE (ORCINUS ORCA) POPULATIONS IN PRINCE WILLIAM SOUND, ALASKA

Killer whales ( Orcinus orca ) feed on a wide variety of fish, cephalopods, and marine mammals throughout their cosmopolitan range; however, the dietary breadth that characterizes the species is not reflected in all populations. Here, we present the findings of a 14-yr study of the diet and feeding habits of killer whales in Prince William Sound, Alaska. Two non-associating forms of killer whale, termed resident and transient (Bigg et al. 1987), were identified. All prey seen taken by transients were marine mammals, including harbor seals ( Phoca vitulina ), Dall's porpoises ( Phocoenoides dalli ), Steller sea lions ( Eumetopias jubatus ), and harbor porpoises ( Phocoena phocoena ). Resident killer whales appeared to prey principally on salmon ( Oncorhynchus spp.), preferring coho salmon ( O. kisutch ) over other, more abundant salmon species. Pacific herring ( Clupea pallasi ) and Pacific halibut ( Hippocampus stenolepis ) were also taken. Resident killer whales frequently were seen to interact in non-predatory ways with Steller sea lions and Dall's porpoises, while transients were not. Differences in the social organization and behavior of the resident and transient killer whales in Prince William Sound are discussed in the light of the dietary differences documented here.     

Gene flow and functional connectivity in the natterjack toad

Functional connectivity is a key factor for the persistence of many specialist species in fragmented landscapes. However, connectivity estimates have rarely been validated by the observation of dispersal movements. In this study, we estimated functional connectivity of a real landscape by modelling dispersal for the endangered natterjack toad (Bufo calamita) using cost distance. Cost distance allows the evaluation of 'effective distances', which are distances corrected for the costs involved in moving between habitat patches in spatially explicit landscapes. We parameterized cost-distance models using the results of our previous experimental investigation of natterjack's movement behaviour. These model predictions (connectivity estimates from the GIS study) were then confronted to genetic-based dispersal rates between natterjack populations in the same landscape using Mantel tests. Dispersal rates between the populations were inferred from variation at six microsatellite loci. Based on these results, we conclude that matrix structure has a strong effect on dispersal rates. Moreover, we found that cost distances generated by habitat preferences explained dispersal rates better than did the Euclidian distances, or the connectivity estimate based on patch-specific resistances (patch viscosity). This study is a clear example of how landscape genetics can validate operational functional connectivity estimates.     

Streams over mountains: influence of riparian connectivity on gene flow in the Pacific jumping mouse (Zapus trinotatus)

In species affiliated with heterogeneous habitat, we expect gene flow to be restricted due to constraints placed on individual movement by habitat boundaries. This is likely to impact both individual dispersal and connectivity between populations. In this study, a GIS-based landscape genetics approach was used, in combination with fine-scale spatial autocorrelation analysis and the estimation of recent intersubpopulation migration rates, to infer patterns of dispersal and migration in the riparian-affiliated Pacific jumping mouse (Zapus trinotatus). A total of 228 individuals were sampled from nine subpopulations across a system of three rivers and genotyped at eight microsatellite loci. Significant spatial autocorrelation among individuals revealed a pattern of fine-scale spatial genetic structure indicative of limited dispersal. Geographical distances between pairwise subpopulations were defined following four criteria: (i) Euclidean distance, and three landscape-specific distances, (ii) river distance (distance travelled along the river only), (iii) overland distance (similar to Euclidean, but includes elevation), and (iv) habitat-path distance (a least-cost path distance that models movement along habitat pathways). Pairwise Mantel tests were used to test for a correlation between genetic distance and each of the geographical distances. Significant correlations were found between genetic distance and both the overland and habitat-path distances; however, the correlation with habitat-path distance was stronger. Lastly, estimates of recent migration rates revealed that migration occurs not only within drainages but also across large topographic barriers. These results suggest that patterns of dispersal and migration in Pacific jumping mice are largely determined by habitat connectivity.     

Habitat-related dispersal in the rock-dwelling land snail Chondrina clienta

Habitat type may influence dispersal in a variety of animal species We examined dispersal in the rock-dwelling land snail Chondrina clienta m a limestone pavement, on vertical rock walls, a pile of stones and a stone wall on the Baltic island of Oland, Sweden Dispersal was estimated by recording movements of marked C clienta in natural populations over 3 yr Dispersal differed significantly between habitat types The largest distances dispersed were recorded in the limestone pavement (264 cm yr¹, median distance) and on vertical rock walls (96 cm yr¹), simple habitats for travelling snails Dispersal was less in the stone pile (68 cm yr¹) and on the stone wall (88 cm yr¹) that are more complex habitats with multiple layers of pieces of stones Distances dispersed also varied among vertical rock walls, indicating that other factors such as exposure of the rock surface and size of the habitat may be important in determining snail dispersal The results of two experiments indicated that grassland vegetation may inhibit dispersal in C clienta , and that isolated stones covered with lichens might serve as stepping stones for dispersing snails in otherwise unsuitable grassland Snail size (age) influenced distances moved, but might only be important in determining daily movements, not dispersal over longer periods Dispersal in C clienta is habitat-specific and cannot be characterized by a single parameter     

Current and historical patterns of drainage connectivity in eastern Australia inferred from population genetic structuring in a widespread freshwater fish Pseudomugil signifer (Pseudomugilidae)

Dispersal can play an important role in the genetic structuring of natural populations. In this regard, freshwater fishes often exhibit extensive population genetic subdivision and are ideal subjects for investigating current and historical patterns of connection and dissociation between drainages. We set out to generate a comprehensive molecular phylogeny for a widespread freshwater fish from eastern Australia, the Pacific blue-eye Pseudomugil signifer. Although movement via flood events may be important in the southern end of the species' range, genetic structuring revealed the importance of historical drainage connections and dissociations in mediating or disrupting dispersal. A dominant feature of our phylogeny is a split between northern and southern populations, which appears to be congruent with a biogeographical barrier recently implicated as important for the connectivity of freshwater organisms in eastern Australia. The extent of the split also has taxonomic implications consistent with suggestions that the Pacific blue-eye may represent more than a single species.     

Constraints on recovery: using molecular methods to study connectivity of aquatic biota in rivers and streams

1. The ‘Field of Dreams Hypothesis’ states ‘if we build it, they will come’, referring to the assumption that if habitats are restored, species will recolonise them. However, the ability of a species to recolonise a restored site will depend not only on the appropriate habitat being present, but also on the ability to get there. This is likely to depend on both the species’ dispersal behaviour and the position of a site in the landscape. 2. Animals with good potential for dispersal are more likely to be able to disperse to newly restored sites. Similarly, sites in lowland streams with limited altitudinal differences between sites may be easier to reach than upstream sites. This is because upstream sites are connected to one another via lowland streams that have different characteristics and therefore may be difficult for animals to traverse. 3. In this paper, genetic data from a range of freshwater species that have been analysed in my laboratory are used to assess the importance of life cycle and position in the landscape (i.e. upland versus lowland streams) on connectivity patterns (and thus recolonisation potential) among populations. 4. In general, contemporary dispersal across catchment boundaries is negligible, except for aquatic insects with an adult flight stage. Dispersal among streams within catchments appears to be more limited than was predicted from knowledge on life histories, except for fish in lowland rivers and streams. 5. As predicted, dispersal of fish, crustaceans and molluscs among streams within catchments is significantly greater in lowland rivers than in upland streams. 6. Overall, these analyses suggest that, with the exception of most insects, and fishes in lowland rivers, natural recolonisation of restored sites is only likely from sites within the same stream. If a species has disappeared from the whole stream, then restoration of habitat alone may not be sufficient for its re‐establishment.     

Between‐patch natal dispersal declines with increasing natal patch size and distance to other patches in the endangered Southern Dunlin Calidris alpina schinzii

Natal dispersal has profound consequences for populations through the movement of individuals and genes. Habitat fragmentation reduces structural connectivity by decreasing patch size and increasing isolation, but understanding of how this impacts dispersal and the functional connectivity of landscapes is limited because many studies are constrained by the size of the study areas or sample sizes to accurately capture natal dispersal. We quantified natal dispersal probability and natal dispersal distances in a small migratory shorebird, the Southern Dunlin Calidris alpina schinzii, with data from two extensively monitored endangered metapopulations breeding in Sweden and Finland. In both metapopulations philopatry was strong, with individuals returning to or close to their natal patches more often than expected by chance, consistent with the patchy distribution of their breeding habitat. Dispersal probabilities were lower and dispersal distances were shorter in Sweden. These results provide a plausible explanation for the observed inbreeding and population decline of the Swedish population. The differences between Sweden and Finland were explained by patch‐specific differences. Between‐patch dispersal decreased with increasing natal patch size and distance to other patches. Our results suggest that reduced connectivity reduces movements of the philopatric Dunlin, making it vulnerable to the effects of inbreeding. Increasing connectivity between patches should thus be one of the main goals when planning future management. This may be facilitated by creating a network of suitably sized patches (20–100 ha), no more than 20 km apart from each other, from existing active patches that may work as stepping stones for movement, and by increasing nest success and pre‐fledging survival in small patches.