The effect of host species on growth and variability of Echinorhynchus salmonis Müller, 1784 (Acanthocephala: Echinorhynchidae), with special reference to the status of the genus

Summary The sex and age (as measured by length) of Echinorhynchus salmonis Müller, 1784 and the host species of this acanthocephalan considerably affected worm body form and size, as well as size of proboscis, proboscis hooks, proboscis receptacle, lemnisci, testes and cement glands. Linear regression analysis indicated that curves describing the growth pattern of these characters by worm length were significantly different as a function of host species. The larger worms recovered from bloater (Coregonus hoyi: Salmonidae) almost invariably showed higher regression coefficient compared to those from smelt (Osmerus mordax: Osmeridae) in all characters. Taxonomic implications of these findings are discussed. Abnormalities in body wall, proboscis hook orientation, lemnisci and cement gland ducts as well as variations in proboscis hook and cement gland numbers are reported, some for the first time. Findings from studies of cement gland pattern invalidate Petrochenko's (1956) splitting up of the genus into three: Echinorhynchus, Metechinorhynchus and Pseudoechinorhynchus. It is proposed that the designation of the last two genera as junior synynoms of the first be accepted. ac]19791205     

Food and habitat partitioning between young-of-year alewives and rainbow smelt in southeastern Lake Ontario

Synopsis Thermal and depth distributions, diets and time of feeding of young-of-year (YOY) alewives and YOY rainbow smelt were compared for evidence of resource partitioning in southeastern Lake Ontario. YOY alewives were largely epilimnial during August and September, but moved toward the bottom during fall turnover. Alewives were most abundant in the warmest available water. YOY rainbow smelt were concentrated at depths between 10 and 30 m in August and September, but moved into deeper water at fall turnover. Depth distribution of YOY smelt was correlated with temperatures of 8–12°C Both species fed predominantly during day on zooplankton during August and September. Cyclopoid copepods were the most common prey, but bosminids, eubosminids, and occasionally calanoid copepods were frequently eaten. As YOY rainbow smelt grew (> 60 mm), they consumed more Mysis relicta and amphipods, which became the major prey of rainbow smelt by November. YOY alewives consumed mostly zooplankton in all months. Diet overlap of the two species was greatest in warm water (> 12.0° during October (94.3% similarity) and August (80.0% similarity) and lowest in November (16.9% similarity). Positive size-selection on zooplankton was found in all months for YOY rainbow smelt, but only in late September through November for YOY alewives. Thus, during thermal stratification, the species were spatially segregated by water temperature but had a high degree of overlap in time of feeding and types of prey eaten. In contrast, after fall turnover there was a greater separation in diet but a higher overlap in habitat use.     

Productive infection of Piscirickettsia salmonis in macrophages and monocyte‐like cells from rainbow trout,a possible survival strategy

Piscirickettsia salmonis is the etiologic agent of the salmonid rickettsial septicemia (SRS), an endemic disease which causes significant losses in salmon production. This intracellular bacterium is normally cultured in salmonid epithelial cell lines inducing characteristic cytopathic effects (CPEs). In this study we demonstrate that P. salmonis is able to infect, survive, replicate, and propagate in the macrophages/monocytes cell line RTS11 derived from rainbow trout spleen, without inducing the characteristic CPEs and the host cells showing the same expression levels as non‐infected control cell. On the other hand, bacteria were capable of expressing specific proteins within infected cells. Infected macrophages cease proliferation and a fraction of them detached from the plate, transform to non‐adhesive, monocyte‐like cells with proliferative activity. Productive infection of P. salmonis into salmonid macrophage/monocyte cells in culture provides an excellent model for the study of host–pathogen interactions, almost unknown in the case of P. salmonis. Our results suggest that the infection of cells from the salmonid innate immune system without inducing an important cell death response should lead to the persistence of the bacteria and consequently their dissemination to other tissues, favoring the evasion of the first line of defense against pathogens. J. Cell. Biochem. 108: 631–637, 2009. © 2009 Wiley‐Liss, Inc.     

Characterization of di‐ and tetranucleotide microsatellite markers in rainbow smelt (Osmerus mordax)

Twelve tetra‐ and di–tetra compound microsatellite loci are described for the rainbow smelt (Osmerus mordax). These loci were screened among a minimum of 79 individuals and the average number of alleles per locus was 16 with an average heterozygosity of 0.86. These microsatellites are being used to examine population structure and life‐history evolution among natural populations of rainbow smelt.     

<Emphasis Type="Italic">Echinorhynchus</Emphasis><Emphasis Type="Italic">hexagrammi</Emphasis> Baeva, 1965 (Acanthocephala: Echinorhynchidae) from marine fishes off Hokkaido,Japan, with morphological observations and new host records

Echinorhynchus hexagrammi Baeva, 1965 is redescribed on the basis of specimens collected from the saffron cod Eleginus gracilis (Tilesius) in Akkeshi Bay (western North Pacific) off Hokkaido, Japan. Eighteen museum specimens deposited as E. salmonis Müller, 1784 from Japanese coastal waters were also re-examined and re-identified as E. hexagrammi. Hexagrammos stelleri Tilesius, Hemitripterus villosus (Pallas), Podothecus sachi (Jordan & Snyder), Sebastes oblongus Günther and Verasper moseri Jordan & Gilbert are recognised as new hosts for E. hexagrammi. This acanthocephalan can be distinguished from three morphologically similar species, E. gadi Zoega in Müller, 1776, E. laurentianus Ronald, 1957 and E. salmonis, by the possession of the following characters: 12–16 (usually 14) rows of hook on the proboscis, a proboscis width of 170–240 μm in males and 195–270 μm in females, a hook root length of 35–45 μm in males and 40–50 μm in females, and linearly or almost linearly arranged cement glands in males.     

Ecological tracers reveal resource convergence among prey fish species in a large lake ecosystem

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Seasonal freeze resistance of rainbow smelt (Osmerus mordax) is generated by differential expression of glycerol-3-phosphate dehydrogenase, phosphoenolpyruvate carboxykinase, and antifreeze protein genes

In winter, rainbow smelt (Osmerus mordax) accumulate glycerol and produce an antifreeze protein (AFP), which both contribute to freeze resistance. The role of differential gene expression in the seasonal pattern of these adaptations was investigated. First, cDNAs encoding smelt and Atlantic salmon (Salmo salar) phosphoenolpyruvate carboxykinase (PEPCK) and smelt glyceraldehyde-3-phosphate dehydrogenase (GAPDH) were cloned so that all sequences required for expression analysis would be available. Using quantitative PCR, expression of beta actin in rainbow smelt liver was compared with that of GAPDH in order to determine its validity as a reference gene. Then, levels of glycerol-3-phosphate dehydrogenase (GPDH), PEPCK, and AFP relative to beta actin were measured in smelt liver over a fall-winter-spring interval. Levels of GPDH mRNA increased in the fall just before plasma glycerol accumulation, implying a driving role in glycerol synthesis. GPDH mRNA levels then declined during winter, well in advance of serum glycerol, suggesting the possibility of GPDH enzyme or glycerol conservation in smelt during the winter months. PEPCK mRNA levels rose in parallel with serum glycerol in the fall, consistent with an increasing requirement for amino acids as metabolic precursors, remained elevated for much of the winter, and then declined in advance of the decline in plasma glycerol. AFP mRNA was elevated at the onset of fall sampling in October and remained elevated until April, implying separate regulation from GPDH and PEPCK. Thus, winter freezing point depression in smelt appears to result from a seasonal cycle of GPDH gene expression, with an ensuing increase in the expression of PEPCK, and a similar but independent cycle of AFP gene expression.     

Predation by longfin smelt ( Spirinchus thaleichthys ) on the mysid Neomysis mercedis in Lake Washington

1. Longfin smelt and mysid samples collected in Lake Washington from 1988 to 1992 were examined, and historic data on smelt and mysid abundance were reviewed to assess the impacts of smelt predation on Neomysis mercedis. The feeding habits of smelt were also examined for differences between odd and even years, and seasonal and ontogenetic variations in smelt diet were described. 2. A reciprocal relationship in the abundance of 1+ smelt and Neomysis suggested that smelt regulate mysid abundance in the lake. 3. The proportion of smelt diet contributed by mysids was higher for less abundant year classes of smelt. 4. These results indicate that intraspecific competition for scarce mysids occurs in the even-year classes of smelt.     

Seasonal variation in the interactions between piscivorous fish,planktivorous fish and zooplankton in a shallow eutrophic lake

The interactions between the higher trophic levels in a shallow eutrophic lake were studied during the course of a year. Three fish species determined the main pathways of organic matter flow within the system: the predominantly planktivorous bream (Abramis brama), the obligate planktivorous smelt (Osmerus eperlanus), and the piscivorous pikeperch (Stizostedion lucioperca). Of the thirteen common zooplankton taxa Daphnia hyalina and cyclopoid copepods were utilized most by the planktivorous fish, while the large production of small cladocerans is almost left unutilized.The seasonal variations of production and consumption are large. This is mainly affected by seasonal variation of the water temperature. The production of O + smelt is efficiently utilized by the pikeperch. Being the most important zooplankton consumer, as well as the most important prey group, O + fish plays a key role in the Tjeukemeer food web.     

Feeding habits of sand smelt (Atherina boyeri, Risso 1810) in Trichonis Lake (Western Greece)

The feeding habits of sand smelt (Atherina boyeri, Risso, 1810) from Trichonis Lake (Western Greece) were investigated. Stomach contents were analyzed from 240 specimens with total lengths ranging from 35 to 112 mm. Samples were taken at monthly intervals (January–December 1997). Of the total number of stomachs examined, 53 were empty (22.1%). However, values varied greatly with season (maximum in January: 50%; minimum in August: 7.6%). Prey analyses of stomach contents identified 15 important items (%Rn > 0.05) belonging to six major groups: crustacean (copepods, cladocera), mollusca (bivalve: larvae), insects (larvae), cestode worms and finfish (fry and eggs). Dominant prey were larvae of the bivalve Dreissena polymorpha (%Rn = 33.8), the copepods Eudiadomus drieschi (%Rn = 26.4) and the cladocera Diaphanosoma brachyurum (%Rn = 24.2). The importance of cladocera and copepods decreased with increasing size of the sand smelt, while the importance of bivalve larvae, fish eggs and finfish fry increased with increasing sand smelt size. Seasonal changes in diet composition and prey abundance in sand smelt stomachs were recorded as coinciding with the seasonal composition and abundance of the zooplankton community in the surface layers of Trichonis Lake. Bivalve larvae were the dominant prey of A. boyeri during January to May, while copepods and cladocera dominated from June to December.     

The importance of escaped farmed rainbow trout (Oncorhynchus mykiss) as a vector for the salmon louse (Lepeophtheirus salmonis) depends on the hydrological conditions in the fjord

The objectives of the study were to see if escaped rainbow trout (Oncorhynchus mykiss) spread rapidly or not from fish farms, and to test whether the hydrological conditions in a fjord influence their vertical distribution and importance as vector for the salmon lice (Lepeophtheirus salmonis). Fifty farmed rainbow trout were tagged with acoustic transmitters including depth sensors and released from two of 11 fish farms in the fjord system. In addition, unintentionally escaped rainbow trout were recaptured for analysis of salmon lice and stomach content. Dispersal out of the fjord system was limited. Most fish stayed in the vicinity of and moved between the fish farms but fed primarily on a variety of indigestible items. They moved in the warm relatively fresh surface layer from late spring until early autumn where the risk of being infested with salmon lice was low. They swam gradually deeper and became much more infested with salmon lice as the surface layers cooled and salinity and temperature gradients became less distinct over the course of the winter. The observed post-escapement behavior may challenge the control of the spread of diseases and parasites between neighboring farms and to wild fish, but also increases opportunities for recapture.     

EVIDENCE FOR MULTIPLE ORIGINS AND SYMPATRIC DIVERGENCE OF TROPHIC ECOTYPES OF SMELT (OSMERUS) IN NORTHEASTERN NORTH AMERICA

The rainbow smelt, Osmerus mordax (Mitchill), is an osmerid fish that exhibits extensive life-history diversity throughout watersheds of northeastern North America. There are both ???sea-run (anadromous) and lake-resident (lacustrine) populations and the latter have diversified further into “dwarf-” and “normal-sized” life-history types. Anadromous and lacustrine smelt may inhabit the same watershed and there are several instances where dwarf and normal populations reside within the same lake. We assayed variation among smelt for morphological traits linked to feeding performance in fishes to see if trophic ecology might promote life-history diversity in Osmerus. We also examined mitochondrial DNA (mtDNA) restriction site variation among forms to assess their evolutionary interrelationships. Dwarf smelt had significantly more gill rakers, larger eyes, but shorter upper jaws than normal lake and anadromous smelt. The populations clustered into two trophic “morphotypes”; an anadromous/normal lake group of populations and a group consisting only of dwarf smelt. The mtDNAs of 444 smelt from 16 populations were digested with 12 restriction enzymes revealing 93 composite mtDNA genotypes that clustered (UPGMA) into two major phylogenetic groups differing by 0.78% in sequence. Both genetic groups were present in dwarf and normal smelt as well as in anadromous fish. Further, geographic proximity, rather than trophic morphotype, appeared to be the major determinant of genetic affinities among populations. In two lakes, however, dwarf and normal smelt populations had significantly different mtDNA genotype frequency distributions indicating that the forms are reproductively isolated within both lakes. A clustering analysis of population affinities suggested that the divergence of sympatric dwarf and normal populations had occurred independently in the two lakes. We concluded (1) that trophic ecology is an important factor promoting differentiation in smelt life histories; (2) that smelt ecotypes are polyphyletic and there have been multiple, independent divergences of Osmerus life-history types throughout northeastern North America; and (3) that the biological and mtDNA differences between coexisting dwarf and normal lake smelt argue strongly that their genetic isolation may have developed sympatrically.     

Seasonal and interannual effects of hypoxia on fish habitat quality in central Lake Erie

1. Hypoxia occurs seasonally in many stratified coastal marine and freshwater ecosystems when bottom dissolved oxygen (DO) concentrations are depleted below 2–3 mg O₂ L^?1. 2. We evaluated the effects of hypoxia on fish habitat quality in the central basin of Lake Erie from 1987 to 2005, using bioenergetic growth rate potential (GRP) as a proxy for habitat quality. We compared the effect of hypoxia on habitat quality of (i) rainbow smelt, Osmerus mordax mordax Mitchill (young‐of‐year, YOY, and adult), a cold‐water planktivore, (ii) emerald shiner, Notropis atherinoides Rafinesque (adult), a warm‐water planktivore, (iii) yellow perch, Perca flavescens Mitchill (YOY and adult), a cool‐water benthopelagic omnivore and (iv) round goby Neogobius melanostomus Pallas (adult) a eurythermal benthivore. Annual thermal and DO profiles were generated from 1D thermal and DO hydrodynamics models developed for Lake Erie’s central basin. 3. Hypoxia occurred annually, typically from mid‐July to mid‐October, which spatially and temporally overlaps with otherwise high benthic habitat quality. Hypoxia reduced the habitat quality across fish species and life stages, but the magnitude of the reduction varied both among and within species because of the differences in tolerance to low DO levels and warm‐water temperatures. 4. Across years, trends in habitat quality mirrored trends in phosphorus concentration and water column oxygen demand in central Lake Erie. The per cent reduction in habitat quality owing to hypoxia was greatest for adult rainbow smelt and round goby (mean: ?35%), followed by adult emerald shiner (mean: ?12%), YOY rainbow smelt (mean: ?10%) and YOY and adult yellow perch (mean: ?8.5%). 5. Our results highlight the importance of differential spatiotemporally interactive effects of DO and temperature on relative fish habitat quality and quantity. These effects have the potential to influence the performance of individual fish species as well as population dynamics, trophic interactions and fish community structure.     

Cladoceran Densities,Day-to-Day Variability in Food Selection by Smelt,and the Birth-Rate-Compensation Hypothesis

Late-evening gut inspection of a dominant planktivore (smelt) and evaluation of densities, fecundities, and body-size distributions in dominant zooplankton prey (cladocerans) were made in day-to-day sequences in June–July (24 days in 1999 and 24 days in 2000). This was conducted as a field test of the hypothesis that species-specific population densities in cladocerans result from size-selective predation by a dominant fish assumed to be a general predator, switching from one prey to another as relative abundance changes. Little of the expected coincidence has been revealed between population density declines and increased numbers of a given prey in smelt diet. However, the data were consistent with the notion that fish would switch from one prey to another depending on the prey relative abundance (the number of prey a fish would see in its reaction field volume). Each cladoceran population fluctuated around its species-specific density level, lower or higher, depending on individual susceptibility to smelt predation, from 0.2 ind. l^–1 in large-bodied Daphnia hyalina and Leptodora kindtii, to 30.0 ind. l^–1 in small-bodied Daphnia cucullata andBosmina thersites. In spite of high fish-to-fish and day-to-day variability in both smelt diet and smelt selectivity for different prey, all cladocerans (also copepods and midge larvae) were equally persistent in smelt diet, and smelt selectivity was similar for small- and large-bodied prey categories, but lower for elongated-(Daphnia, Diaphanosoma) than for compact-body (Bosmina, Chydorus) species, when integrated for the entire sampling time. Closer examination of D. cucullata and B. thersites revealed strong smelt selection for later instars and females with greater clutches, showing that size distribution in a cladoceran population might be structured by fish predation in a similar way to that a cladoceran community (species relative abundance) is structured in a lake habitat. The birth-rate-compensation hypothesis is offered to explain why the value of food selectivity index in a planktivorous fish would remain the same for alternate prey categories with similar life-history traits, unless they differ in susceptibility to predation before the time of first reproduction.     

Seasonal and regional occurrence of <Emphasis Type="Italic">Acanthocephalus</Emphasis> sp. (Acanthocephala: Echinorhynchidae) in fishes and isopods (<Emphasis Type="Italic">Asellus hilgendorfi</Emphasis>) in a lake system in northern Japan

 The echinorhynchid acanthocephalan Acanthocephalus sp. was collected and described from four species of fishes (rainbow trout Oncorhynchus mykiss, Sakhalin huchen Hucho perryi, Japanese pond smelt Hypomesus nipponensis, and threespine stickleback Gasterosteus aculeatus) from a lake system, the Tsugaru-Jūniko Lakes, in Aomori Prefecture, northern Japan. In rainbow trout, the prevalence and intensity of infection markedly differed between lakes, and the fish were most frequently and most heavily infected in the lakes with a dense population of the isopod intermediate host Asellus hilgendorfi. In isopods, the prevalence of acanthocephalan larvae increased in the late winter and reached its highest level in March or April. In rainbow trout, male worms were abundant from winter to spring, and female worms were immature during these seasons. Gravid females were abundant in summer and autumn. These findings indicate that Acanthocephalus sp. is an annual species and its recruitment from the intermediate host to the fish occurs mainly in winter and spring. Received: January 9, 2002 / Accepted: April 18, 2002 Acknowledgments We thank Professor Shōichi Saito, Faculty of Education, Hirosaki University, for his encouragement during this study. Thanks are also due to many students of the Nature Study Laboratory, Faculty of Education, Hirosaki University, for their assistance in the field. We are greatly indebted to the Iwasaki Village Office and Fukaura Forestry Office for giving us permission for the survey. Correspondence to:A. Ohtaka     

Occurrence of Proteocephalus tetrastomus (Rudolphi, 1810) (Cestoda: Proteocephalidea) in larval rainbow smelt (Osmerus mordax) in North America: identification of a potential pathogen confirmed

A morphological evaluation and genetic analysis (sequencing of ITS2 region of rDNA) of proteocephalidean cestodes from rainbow smelt (Osmerus mordax) in the Saint Lawrence Estuary, Canada, has shown their conspecificity with Proteocephalus tetrastomus, a specific parasite of smelt (Osmeridae), previously known only from northern Europe, Russia, and Japan. The parasite occurs only in larval, but not adult, smelt in the Saint Lawrence Estuary. Prevalence of larval smelt infection was 42% (n = 50), mean intensity 2.1 +/- 2.4 and mean abundance 1.1 +/- 1.0.     

Variable littoral‐pelagic coupling as a food‐web response to seasonal changes in pelagic primary production

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Genetic structure and relationships among anadromous and landlocked populations of rainbow smelt, Osmerus mordax, Mitchill, as revealed by mtDNA restriction analysis

Larvae of anadromous rainbow smelt originating in various spawning tributaries are retained in the St Lawrence estuary. We proposed that these smelt represent one population genetically differentiated from adjacent populations characterized by geographically distinct larval retention areas. We also analysed four landlocked populations to evaluate the phylogenetic basis of dwarf and normal-sized phenotypes and their relation to an adromous smelt. A phylogenetic distinction was revealed between anadromous and landlocked smelt, with only one of the two mtDNA phylogenetic groups of anadromous fish observed among landlocked smelt. Significant geographical heterogeneity in the distribution of mtDNA genotypes was observed among landlocked smelt, suggesting that dwarfism in smelt may be polyphyletic in origin. St Lawrence smelt were genetically identical but distinct from adjacent populations, supporting the proposition that population genetic structure reflects the number of larval retention zones rather than spawning sites.     

Proboscis profiler: a tool for detecting acanthocephalan morphotypes

Molecular studies conducted over the past 25 years have revealed previously unrecognised diversity in the phylum Acanthocephala. Several nominal species have been shown to represent complexes of morphologically cryptic biological species, a situation potentially confounding the analysis of ecological data. A software tool, ‘Proboscis profiler’, was developed to detect morphological heterogeneity in collections of superficially similar acanthocephalan worms based on the multivariate statistical analysis of proboscis hook dimensions. Proboscis profiler identifies objective, natural groups in a collection of acanthocephalans which may correspond to distinct biological species or populations. Initial trials demonstrate that Proboscis profiler can discriminate biological acanthocephalan species of the Echinorhynchus gadi Zoega in Müller, 1776 complex and differentiate between dorsal and ventral hook rows from the proboscis of E. salmonis Müller, 1784. Proboscis profiler is free software and can be downloaded from .