Genetic diversity and structure of two hybridizing anadromous fishes (Alosa pseudoharengus,Alosa aestivalis) across the northern portion of their ranges

Alewife (Alosa pseudoharengus) and blueback herring (Alosa aestivalis), also known collectively as either river herring or gaspereau, are anadromous clupeid fishes that display spatiotemporal overlap during riverine spawning migrations. Both species have experienced severe population declines within portions of their ranges. Evidence that they home to their natal rivers to spawn suggests the likelihood of ecologically significant population structure, yet this hypothesis has not been rigorously tested. We examined genetic diversity, differentiation and population structure in 34 alewife and four blueback herring populations spanning a 2,500 km portion of their northern range, using 14 microsatellite loci. Significant differentiation was detected among most rivers, and eight genetically defined alewife population clusters that largely corresponded to hydrographic regions were identified. Similar population structure was seen for blueback herring. Genetic isolation by distance was not significant among alewife populations in regions that have been historically influenced by dams, and/or stock transfers, but was highly significant in two regions that have not been subject to these influences. Genetic differentiation of alewife populations was strongest in the Bay of Fundy. Bottleneck tests revealed evidence of demographic bottlenecks in all of the alewife populations. Lastly, our results indicated that hybridization between alewife and blueback herring is common.     

Diet of alewives,Alosa pseudoharengus and blueback herring,A. aestivalis (Pisces: Clupeidae) in Minas Basin,Nova Scotia,a turbid,macrotidal estuary

Synopsis Stomach contents of anadromous alewives, Alosa pseudoharengus and blueback herring, A. aestivalis, obtained from brush weir and drift net collections in Minas Basin, N.S., were examined. Diets showed much overlap in terms of resource use, but the dietary importance of major prey categories differed substantially between species. Alewives favoured larger, more benthic prey (e.g. amphipods, mysids and crangonids), while blueback herring appeared to concentrate their feeding on microzooplankters (e.g. calanoid copepods, cypris larvae and molluscan veligers). Interspecific differences in diet composition are largely attributed to the planktivorous feeding habits of small (81–155 mm fork length) blueback herring. Differences in prey suggest that alewives utilize a particulate feeding strategy while blueback herring are predominantly filter-feeders. Although competition for food in the Basin seems unlikely since high secondary production yields a superabundance of prey, differences in feeding behaviour between younger, smaller individuals of both species could be a means of avoiding competitive interactions in an environment where there are space/access limitations imposed by the tidal cycle.     

Relationship between feeding of blueback herring and the zooplankton community of a Texas reservoir

Following the introduction of blueback herring (Alosa aestivalis), we examined herring food habits and the crustacean zooplankton community in Lake Theo, a 30-hectare reservoir located on a tributary of the Red River in north Texas. Prior to the introduction in spring, 1982, the reservoir contained an established fish community dominated by centrarchids. Blueback herring stomachs and zooplankton were sampled quarterly from summer, 1982 through winter, 1985. Cladocerans accounted for 89.4% of the zooplankton consumed by herring. Median lengths of cladocerans and copepods in the herring diet were significantly greater than those in the reservoir. No appreciable changes in lengths of cladocerans and copepods in the reservoir were observed after blueback herring introduction, but the zooplankton community shifted from cladoceran to copepod domination.     

Discovery and characterization of single nucleotide polymorphisms in two anadromous alosine fishes of conservation concern

Freshwater habitat alteration and marine fisheries can affect anadromous fish species, and populations fluctuating in size elicit conservation concern and coordinated management. We describe the development and characterization of two sets of 96 single nucleotide polymorphism (SNP) assays for two species of anadromous alosine fishes, alewife and blueback herring (collectively known as river herring), that are native to the Atlantic coast of North America. We used data from high‐throughput DNA sequencing to discover SNPs and then developed molecular genetic assays for genotyping sets of 96 individual loci in each species. The two sets of assays were validated with multiple populations that encompass both the geographic range and the known regional genetic stocks of both species. The SNP panels developed herein accurately resolved the genetic stock structure for alewife and blueback herring that was previously identified using microsatellites and assigned individuals to regional stock of origin with high accuracy. These genetic markers, which generate data that are easily shared and combined, will greatly facilitate ongoing conservation and management of river herring including genetic assignment of marine caught individuals to stock of origin.     

A thermal effluent as a sporadic cornucopia: effects on fish and zooplankton

Synopsis We report effects of a thermal effluent on fish and zooplankton in an impoundment in South Carolina. The effects are not directly due to heated effluent but are indirect effects on the trophic dynamics of the system and depend on the physical structure of the system, season, and geography. As effluent from a nuclear reactor cooling pond (Pond C) enters the larger Par Pond it sporadically carries with it zooplankton and/or dead fish. Pond C produces high zooplankton densities in two situations: (1) high and hot effluent flow during cold winter weather; and (2) low (cool) effluent flow during warmer seasons. High zooplankton densities attract Blueback herring, Alosa aestivalis, which attract largemouth bass, Micropterus salmoides. Dying and dead bluegills, Lepomis macrochirus from Pond C (heat killed) are eaten by Par Pond bass which swim into effluent temperatures as high as 46°C to take these easy prey. Blueback herring and largemouth bass distribution are affected by the presence of food rather than by temperature. Par Pond blueback herring appear to be available to bass mainly when herring are near the effluent from Pond C and when they spawn in spring in the littoral zone. Because Blueback herring in Par Pond live only one year and because their presence in bass habitat is seasonal there is a strong seasonal component to bass food abundance. A previously reported annual oscillation in bass condition (K), with a peak in winter, occurs throughout Par Pond but is extreme in the vicinity of the effluent. The strong seasonality of food abundance for bass at the effluent correlates the winter peak in bass condition.     

Quantifying alosine prey in the diets of marine piscivores in the Gulf of Maine

The objectives of this work were to quantify the spatial and temporal distribution of the occurrence of anadromous fishes (alewife Alosa pseudoharengus, blueback herring Alosa aestivalis and American shad Alosa sapidissima) in the stomachs of demersal fishes in coastal waters of the north‐west Atlantic Ocean. Results show that anadromous fishes were detectable and quantifiable in the diets of common marine piscivores for every season sampled. Even though anadromous fishes were not the most abundant prey, they accounted for c. 5–10% of the diet by mass for several marine piscivores. Statistical comparisons of these data with fish diet data from a broad‐scale survey of the north‐west Atlantic Ocean indicate that the frequency of this trophic interaction was significantly higher within spatially and temporally focused sampling areas of this study than in the broad‐scale survey. Odds ratios of anadromous predation were as much as 460 times higher in the targeted sampling as compared with the broad‐scale sampling. Analyses indicate that anadromous prey consumption was more concentrated in the near‐coastal waters compared with consumption of a similar, but more widely distributed species, the Atlantic herring Clupea harengus. In the context of ecosystem‐based fisheries management, the results suggest that even low‐frequency feeding events may be locally important, and should be incorporated into ecosystem models.     

Feeding chronology and habits of Alosa spp. (Clupeidae) juveniles from the lower Hudson River estuary, New York

Synopsis Feeding habits of juvenile alewife, A. pseudoharengus, blueback herring, A. aestivalis, and American shad, Alosa sapidissima, were examined over a 24 h period at an inshore location on the lower Hudson River. Feeding periodicity differed by species: alewives showed no diel differences, whereas shad fed least intensively near dawn and herring least intensively at night. Alewives fed primarily on chironomids and the amphipod Corophium lacustre, shad on Formicidae and larval chironomids, and herring on chironomids and copepods. Diel variation in prey selection was evident. Dietary overlap was generally greatest between alewife and herring and least between shad and herring.     

Ovarian dynamics and fecundity regulation in blueback herring,Alosa aestivalis,from the Connecticut River,US

We analyzed ovarian dynamics of anadromous blueback herring, Alosa aestivalis, in Connecticut River with the principal aim of exploring oocyte recruitment and how it shapes the fecundity pattern. We examined the oocyte release strategy and analyzed spawning cyclicity by linking oocyte growth to the degeneration of postovulatory follicles. Females were accordingly classified as pre-spawners, early and late active spawners, and oocyte recruitment intensity was compared among the different spawning phases. Oocyte recruitment occurred continuously and in parallel with spawning activity, a pattern which is diagnostic of indeterminate fecundity. However, both fecundity and oocyte recruitment intensity progressively decreased (tapered) throughout spawning, until the ovary was depleted of vitellogenic oocytes. There was no massive atresia of vitellogenic oocytes at the end of the spawning season, which is atypical of indeterminate spawners. We propose that tapering in oocyte recruitment and fecundity is an adaptation to the high energetic expenditure of the upstream spawning migration.     

Growth, ingestion rates and metabolic activity of walleye in lakes with and without lake herring

Growth efficiencies, ingestions rates and activity levels of walleye Sander vitreus were compared in lakes with and without lake herring Coregonus artedi . Yellow perch Perca flavescens were the main prey in lakes without lake herring. Walleye were sampled in September and October from 38 lakes in Ontario in 1998 and 1999, using multimesh monofilament gillnets. Ingestion rates were estimated from annual increments in somatic mercury and body mass, and the mercury content of yellow perch and lake herring. Walleye had higher growth efficiencies, and lower ingestion and activity rates in lakes with lake herring. Lake herring grow larger than yellow perch and therefore could provide more profitable prey for larger walleye. The results are consistent with optimal foraging theory that predicts that walleye feeding on optimal prey sizes should grow more efficiently, if the ratio of feeding benefit (energy) to cost (search and seizure) is a function of the ratio of predator and prey size.     

Factors affecting migration patterns of juvenile river herring in a coastal Massachusetts stream

Juvenile anadromous river herring (alewives, Alosa pseudoharengus, and blueback herring, Alosa aestivalis) spend 3–7 months in freshwater before migrating to the ocean, often exhibiting waves of early and late migrations. Migratory patterns and associated abiotic and biotic factors were examined for both species migrating in 2003 from Herring River in Bourne Massachusetts. Migrating herring were grouped into two temporal periods and separated by species. Relationships between abiotic (water temperature, lunar phase, and precipitation) and biotic (size, age, growth rate, and hatch date) factors and the timing of out-migration (transition to seawater) were examined. Blueback herring migration coincided with a sharp decrease in temperature consistent with the time of year. Peaks in alewive migration may reflect adaptive traits maximizing likelihood of survival through seasonal marginal stream habitats and also avoiding lethal water temperatures in winter. Migrating bluebacks exhibited a single migratory period (late September–October) which occurred between the early and late migratory pulses of the alewives. These patterns may also represent distinct migration strategies between early and late migrating alewives and between the two species of river herring. Early migration may be a mechanism by which factors such as competition and low food availability are mitigated. In contrast, environmental conditions during the post hatching time of late migrators may allow for a size maximizing strategy that promotes migration from the nursery area as late as possible.     

Detecting population structure in a high gene-flow species,Atlantic herring (Clupea harengus): direct,simultaneous evaluation of neutral vs putatively selected loci

In many marine fish species, genetic population structure is typically weak because populations are large, evolutionarily young and have a high potential for gene flow. We tested whether genetic markers influenced by natural selection are more efficient than the presumed neutral genetic markers to detect population structure in Atlantic herring (Clupea harengus), a migratory pelagic species with large effective population sizes. We compared the spatial and temporal patterns of divergence and statistical power of three traditional genetic marker types, microsatellites, allozymes and mitochondrial DNA, with one microsatellite locus, Cpa112, previously shown to be influenced by divergent selection associated with salinity, and one locus located in the major histocompatibility complex class IIA (MHC-IIA) gene, using the same individuals across analyses. Samples were collected in 2002 and 2003 at two locations in the North Sea, one location in the Skagerrak and one location in the low-saline Baltic Sea. Levels of divergence for putatively neutral markers were generally low, with the exception of single outlier locus/sample combinations; microsatellites were the most statistically powerful markers under neutral expectations. We found no evidence of selection acting on the MHC locus. Cpa112, however, was highly divergent in the Baltic samples. Simulations addressing the statistical power for detecting population divergence showed that when using Cpa112 alone, compared with using eight presumed neutral microsatellite loci, sample sizes could be reduced by up to a tenth while still retaining high statistical power. Our results show that the loci influenced by selection can serve as powerful markers for detecting population structure in high gene-flow marine fish species.     

Differential expression, induction, and stability of CYP1A4 and CYP1A5 mRNA in chicken and herring gull embryo hepatocytes

2,3,7,8-Tetrachlorodibenzo-p-dioxin (TCDD) induces cytochrome P4501A (CYP1A) catalyzed ethoxyresorufin-O-deethylase (EROD) activity in chickens and other avian species. To investigate mechanisms underlying the effectiveness of EROD activity as a biomarker for exposure to dioxin-like compounds in avian models, we characterized inter-species differences in isoform-specific CYP1A mRNA expression, induction, and stability in chickens (Gallus gallus domesticus) and herring gulls (Larus argentatus). Exposure to 100 nM TCDD significantly increased CYP1A4 and CYP1A5 mRNA expression in chicken and herring gull embryo hepatocyte cultures. Chicken CYP1A4 and CYP1A5 were induced 61-fold and 25-fold respectively. The herring gull isoforms were induced 2.2- and 4.3-fold respectively. In both species, the isoform that was preferentially induced exhibited lower constitutive expression. Half-lives of chicken CYP1A4, chicken CYP1A5, and herring gull CYP1A5 mRNA ranged from 5.0 to 7.0 h in cultured hepatocytes. The half-life of herring gull CYP1A4 mRNA was 2.5 h. Our findings indicate that expression, induction, and stability of CYP1A4 and CYP1A5 mRNA are differentially regulated in chickens and herring gulls. In particular, CYP1A4 is preferentially induced in chickens, while CYP1A5 is preferentially induced in herring gulls. We propose that CYP1A5 mRNA expression may be a sensitive biomarker of exposure to dioxin-like compounds in some avian species.     

Discrimination of Baltic herring populations with respect to Anisakis simplex larvae infection

Samples of herring Clupea harengus in the Baltic were collected during the spawning season in western, middle and eastern waters off the Polish coast. Two samples were taken per area, several morphometric characters were measured and some meristic characters (vertebrae and gill rakers) were counted. Infection with Anisakis simplex larvae, total length, age and maturity stage were recorded. Univariate and multivariate statistical tests were used to compare the samples, discriminant analysis was applied to separate herring populations and the fraction of herring populations in each area was estimated. The results obtained indicated that infected fish represented a mixed group of western and central Baltic herring. In the first samples collected from each area, the share of the western population decreased from 40% in the western to 18% in the middle and 13% in the eastern coasts. The share of western herring was higher in the second samples collected in the same area and decreased from 91% in the western coast to 14 and 32% in the middle and eastern ones, respectively.     

Distribution of viral haemorrhagic septicaemia virus in wild fish species of the North Sea, north east Atlantic Ocean and Irish Sea.

A surveillance programme was initiated on the occurrence and distribution of viral haemorrhagic septicaemia virus (VHSV) in wild marine fish. Six research cruises were undertaken in an 18 mo period during 1997 and 1998, covering the North Sea, the Atlantic waters off the north and west coasts of Scotland and the Irish Sea. A total of 19,293 fish were sampled from 23 different species including cod, haddock, Norway pout, herring and sprat. Individual fish lengths were recorded and the fish were checked for lesions, haemorrhaging and other signs of disease. Pools of organ samples were taken for virus assay. The majority of fish sampled did not display clinical signs indicative of viral haemorrhagic septicaemia. A small number of cod were found with skin lesions and haddock with skin haemorrhaging. Of the 2081 organ and skin sample pools collected, 21 tested positive for VHSV by tissue culture and enzyme-linked immunosorbent assay. Seventeen of the isolates originated from Norway pout Trisopterus esmarkii, one from cod Gadus morhua (skin lesion), one from herring Clupea harengus, one from whiting Merlangius merlangus, and one from a previously unreported host species, poor cod Trisopterus minutus.     

Parental effects on size variation in fish larvae

The growth and changes in the standard deviation about mean length were measured for sole, Solea solea L., and herring, Clupea harengus L., larvae from crosses of one or two males with one or two females. The sizes of sole larvae did not differ between crosses but the size variation increased faster with age when two females were used. Both maternal and paternal effects were detected for size at hatching in herring, but there was no parental influence on size variation. Size variation in herring larvae declined or was stable during the yolk sac stage and was not affected by the number of parents used for fertilizations.     

Feeding ecology of mackerel and dietary overlap with herring in Icelandic waters

The Northeast Atlantic mackerel (Scomber scombrus) has been extending its summer feeding distribution north and west, including around Iceland, since around 2006. The objective of this study is to quantify the weight gain and total food consumption of mackerel and to evaluate the food competition between mackerel and herring (Clupea harengus) feeding in the marine ecosystem around Iceland during the summers 2009–2011. Mackerel feeding in Icelandic waters gained around 43% on average in weight during these summers. Based on swept-area abundance estimates of mackerel from an international survey in 2011 and available estimates of food conversion efficiency in mackerel, the weight gain in Icelandic waters in 2011 corresponded to a total consumption of around 3.4 (2.4?4.5) million tonnes. Overall, 98% of 2314 mackerel, 91% of 398 Icelandic summer-spawning herring and 96% of 424 Norwegian spring-spawning herring stomachs, collected over the summers 2009?2011, contained food. The mean weight of the stomach content of mackerel was higher than for herring in all the years. While the stomach content weight of mackerel was generally highest in southeastern and southwestern areas, it was highest for herring in western, eastern and northern areas. Analysis of stomach contents showed that Copepoda were the most important food of mackerel in most areas, while Copepoda and Euphausiacea were the most important food items for herring. Fish prey contributed a higher proportion in stomachs of mackerel than herring, although relatively low for both species.     

Distribution patterns of Baltic herring larvae, Clupea harengus L., in the coastal waters off Helsinki, Finland

Changes in the spatial distribution of Baltic herring larvaethrough time were investigated in the waters off Helsinki, thenorthern Baltic Sea. The average larval densities were comparableto the relatively low levels (mean around 2 larvae m^–2)found in other studies in the Gulf of Finland. In shallow baysvirtually no herring larvae were found in early June. In lateJune and early July the densities of larvae longer than 10 mmincreased to 1–4 larvae m^–2 in shallow areas (depth<2 m), i.e. to levels higher than those recorded in deeperwaters. In combination with data on the size distributions ofthe larvae, these observations suggest that a significant proportionof the herring larvae are found in shallow water after the yolksac stage. We conclude that shallow coastal waters are potentiallyimportant nursery areas for herring larvae and that this featureof herring larval biology should be taken into account in planningsampling programs for larval herring.     

Vulnerability of late larval and early juvenile Atlantic herring, Clupea harengus, to predation by whiting,Merlangius merlangus

Laboratory experiments investigated changes in the vulnerability of herring to predation by whiting during growth from <34 to >80mm in length, Premetamorphic herring (<50mm) failed to react to c . 50% of attacks by the predator, but this was reduced to c . 20% in postmetamorphic fish. Premetamorphic herring failing to react had c. 30% probability of survival due to unforced errors by the predator (unsuccessful attacks that did not elicit a reaction by the prey), but these errors did not occur after herring metamorphosis. The loss of the advantage of lower conspicuousness following metamorphosis was counterbalanced by increased reactivity. Antipredator benefits should increase with schooling behaviour (first evident in experiments at c . 50 mm), so predation mortality should decrease in postmetamorphic herring in the sea. Metamorphosis itself would be associated with high predation risk since conspicuousness is enhanced but reactivity and schooling behaviour are still not fully developed.     

Estimating piscine prey size from partial remains: testing for shifts in foraging mode by juvenile bluefish

Knowledge of prey sizes consumed by a predator aids in the estimation of predation impact. Young-of-the-year bluefish, Pomatomus saltatrix, attack their prey tail-first and often bite their prey in half; this poses a unique problem in determining prey sizes from stomach content analysis. We developed a series of linear regressions to estimate original prey lengths from measurements of eye diameter and caudal peduncle depth for striped bass, Morone saxatilis, bay anchovy, Anchoa mitchilli, American shad, Alosa sapidissima, blueback herring, Alosa aestivalis, Atlantic silverside, Menidia menidia, and white perch, Morone americana. We then used these regressions to estimate original prey sizes from pieces of prey found in stomachs of bluefish collected in the Hudson River estuary from 1990–1993. Lengths of prey that were swallowed whole were compared to estimated lengths of prey that were consumed in pieces. Lengths of prey that were consumed in pieces were larger than prey that were consumed whole. We determined the prey length/predator length ratio at which bluefish began shifting from swallowing their prey whole to partial consumption. Shifting occurred at a ratio of approximately 0.35 irrespective of prey species, suggesting that prey length plays an important role in predator foraging decisions and may contribute to gape limitations. Shifts in foraging mode effectively reduce gape limitation and allow bluefish to consume larger prey sizes which may increase their effect on prey populations.     

Diet of Atlantic cod in the southern Gulf of St Lawrence as an index of ecosystem change, 1959–2000

Major temporal changes in the importance of euphausiids and Atlantic herring Clupea harengus in the diet of Atlantic cod Gadus morhua , 10–75 cm L _F, occurred in shallow waters (<100 m depths) of the southern Gulf of St Lawrence between 1959 and 2000. Euphausiids represented 6–70% of prey mass for Atlantic cod 31–60 cm L _F from 1959 to 1987 but only trace amounts were detected in stomachs collected from 1990 to 2000. Mysids and gammarid amphipods (for Atlantic cod ≤45 cm L _F), and Atlantic herring (for Atlantic cod >45 cm L _F) largely replaced euphausiids in the Atlantic cod diets from 1990 to 2000. This diet change suggested there has been a major perturbation of the food web of the southern Gulf of St Lawrence. The importance of fishes (mostly Atlantic herring) in the diet of Atlantic cod >45 cm L _F increased significantly between the periods 1959–1980 and 1987–2000. Atlantic herring comprised 0–4% (mean 1.3%) of prey mass of Atlantic cod 46–60 cm L _F from 1959 to 1980 and increased to 2–42% (mean 19.6%) of the diet from 1987 to 2000. Atlantic herring comprised 0–25% (mean 9.4%) of the prey mass of Atlantic cod 61–75 cm L _F from 1959 to 1980 and increased to 42–81% (mean 54.2%) of prey mass from 1987 to 2000. This increased consumption of Atlantic herring was consistent with observed changes in abundance of Atlantic herring in the ecosystem between the late 1970s and 2000. The large changes in consumption of euphausiids and Atlantic herring represent highly significant changes and would need to be included in the development and interpretation of ecosystem-based management models for this ecosystem.