Electrophoretic identity between allis shad, Alosa alosa (L.), and twaite shad, A.fallax (Lacépède)

Morphological characteristics as well as electrophoretic polymorphism have been analysed in eight samples of allis shad, Alosa alosa (L.) and twaite shad, Alosa fallax (Lacepede), collected in the Loire basin and the Gironde-Dordogne system. The morphological characteristics showed that allis and twaite shad were present in these samples. Moreover, specimens with intermediate characteristics were found in the Loire and assumed to be hybrids between the two forms. By contrast, the two species were monomorphic and electrophoretically indistingishable at the 22 loci analysed. Therefore, it cannot be excluded that these two forms correspond to a single species.     

Phylogenetic relationships among North American Alosa species (Clupeidae)

A phylogeny of the six North American species in the genus Alosa , with representatives of three Eurasian species, was generated using mtDNA sequences. This was accomplished by obtaining sequences for three North American species and additional geographical sampling of the other three species. The subgenus Alosa , including the formerly recognized subgenus Caspialosa , formed a strongly supported monophyletic group. Alosa alabamae was part of a polytomy with Alosa sapidissima , which was interpreted to support the recognition of A. alabamae as an incipient, yet distinct, species. The subgenus Pomolobus was not recovered as a monophyletic group. Alosa chrysochloris was basal to all other Alosa , although this position was only weakly supported. Previous work had indicated that Alosa pseudoharengus and Alosa aestivalis are not reciprocally monophyletic, but additional sampling in this study did not detect any further cases of shared haplotypes between the two species. The phylogeny supports previous hypotheses that the evolution of North American Alosa species in the Gulf of Mexico ( A. chrysochloris and A. alabamae ) was the result of two independent events. First, the ancestor of A. chrysochloris was isolated in the Gulf of Mexico, likely by the close of the Suwannee Straits, and this was followed later by dispersal of the ancestor of A. alabamae around the Florida peninsula into the Gulf of Mexico sometime during or after the Pleistocene.     

Development of hypoosmoregulatory ability in allis shad Alosa alosa

When different ages of juvenile allis shad Alosa alosa (18–74 days post-fertilization) were exposed for 48 h to elevated salinity (25 or 30), all ages could tolerate moderates salinity (25) whereas only the older stages could tolerate the higher salinity. In 88 days post-fertilization allis shad, acclimatation to full-strength sea water for 2 weeks did not lead to significant changes in total water content and gill Na⁺/K⁺-ATPase activity whereas total Na⁺ content increased within 5 days to values which remained stable thereafter. Together, these data suggest that juvenile allis shad develop a limited euryhalinity, which allows them to fully adapt to hyperosmotic environment immediately after transfer.     

Microsatellite DNA markers for American shad (Alosa sapidissima) and cross‐species amplification within the family Clupeidae

Sixteen microsatellite loci were identified and characterized for American shad (Alosa sapidissima). The number of alleles per locus observed ranged from eight to 32 and averaged 15.4 alleles. Average observed heterozygosity was 81.1%. The markers were screened using four other species from the family Clupeidae. Amplification success among Alosa species was 79.2% with 81.6% polymorphism among those markers that amplified successfully. Amplification success was poor in Dorosoma (31.3%). Due to allelic diversity and estimates of heterozygosity, these markers can be useful in A. sapidissima for population level analyses, parentage assignment and broodstock management.     

Metabolic rates in an anadromous clupeid, the American shad (Alosa sapidissima)

To assess the energetics of migration in an anadromous fish, adult American shad (Alosa sapidissima) were swum in a large respirometer at a range of speeds (1.0–2.3 body lengths (BL) s⁻¹, 13–24 °C). Metabolic rate (M_O2) was logarithmically related to swimming speed (Bl s⁻¹; r ² = 0.41, slope = 0.23 ± 0.037) and tailbeat frequency (beats × min⁻¹; r ² = 0.52, slope = 0.003 ± 0.0003). Temperature had a significant effect on metabolic rate (r ² = 0.41) with a Q₁₀ of 2.2. Standard metabolic rate (SMR), determined directly after immobilization with the neuroblocker gallamine triethiodide, ranged from 2.2–6.2 mmolO₂ kg⁻¹ h⁻¹ and scaled with mass (W) such that SMR = 4.0 (±0.03)W^{0.695(±0.15)}. Comparison of directly determined and extrapolated SMR suggests that swimming respirometry provides a good estimate of SMR in this species, given the differences in basal activity monitored by the two methods. Overall, American shad metabolic rates (M_O2 and SMR) were intermediate between salmonids and fast-swimming perciforms, including tunas, and may be a result of evolutionary adaptation to their active pelagic, schooling life history. This study demonstrates variability in metabolic strategy among anadromous fishes that may be important to understanding the relative success of different migratory species under varying environmental conditions. Accepted: 3 March 1999     

Mitochondrial genome of the American shad Alosa sapidissima

The complete mitochondrial genome of Alosa sapidissima has been determined. The total length of the mitogenome was 16,697 bp and had a gene content (13 protein-coding, 22 tRNAs and 2 rRNAs. Except for the seven tRNA and Nd6 genes, all other mitochondrial genes are encoded on the heavy strand. The overall base composition of the heavy strand is 28.3% A, 24.8% T, 28.9% C, 17.9% G, with an AT content of 53.1%. The DNA sequence of Alosa. sapidissima shared 97.1, 93.9, 88.8 and 82.3% sequence identity with that of Alosa alosa, Alosa pseudoharengus. Molecular data here presented provide a useful toll for evolutionary as well as population genetic studied.     

Genetic relationships among the shads (Alosa) revealed by mitochondrial DNA analysis

We surveyed restriction site differences in mitochondrial DNA. (mtDNA) among five species of shad ( Alosa ) from North America and Europe. Allis shad, Alosa alosa and twaite shad, Alosa fallax shared two divergent genotype groups, suggesting that the two forms are either a single species, or are distinct species that have hybridized. Phenetic and cladistic analyses of the relationships among the mitochondrial genotypes defined two groups of shad, corresponding to the subgenera, Alosa and Pomolobus . The mean estimated sequence divergence between the mtDNAs of these two groups of shad was 6.5%. Taken in conjunction with fossil data, this divergence estimate suggests that the rate of mtDNA divergence between the two subgenera has been almost 10-fold lower than the 'conventional' clock calibration for mtDNA.     

美国鲥鱼对我国淡水生态系统的潜在入侵风险

近年来美国鲥鱼已被引入到我国的上海和广东等地进行水产养殖。本实验报道了在纯淡水人工养殖条件下美国鲥鱼的繁殖和生长情况。人工养殖鲥鱼的生长速度接近美国北卡罗莱纳州沿岸野生鲥鱼的生长速度,而高于特拉华河野生鲥鱼的生长速度;人工养殖的2龄鲥鱼既可以在养殖池中自然产卵,也可以通过注射激素人工诱导产卵,其平均受精率、孵化率和仔鱼17天内的存活率分别约15%、75%和54%。这一结果表明,尽管鲥鱼是在海水中进行繁殖的洄游性鱼类,但在生长环境适宜的淡水条件下同样可以正常生长,并达到性成熟和繁育后代,有可能在适宜的淡水水体中形成自我维持的种群。结合本实验结果和美国鲥鱼的其他生物学特性,本文认为美国鲥鱼在我国天然水体生态系统中具有潜在的入侵性。     

Genetic monitoring of two decades of hybridization between allis shad (Alosa alosa) and twaite shad (Alosa fallax)

Habitat alteration has been implicated in driving hybridization between the sympatric migratory shads Alosa alosa and Alosa fallax. Morphological and molecular evidence is consistent with hybridization across the overlapping range of these species, but the temporal extent of hybrid occurrence and genetic consequences for populations have not been explored. Using eight nuclear microsatellite loci and samples collected between 1989 and 2008 in the Solway Firth (UK), we genetically identified hybrids, studied temporal changes in their frequency, and explored changes in allele frequencies of parental populations. These molecular data confirmed the hybrid status of individuals identified using morphology (number of rakers on the outer gill arch), and enabled separation of hybrids from purebred individuals. Mitochondrial cytochrome-b sequencing revealed the presence of two haplogroups, each predominantly occurring in one species. Heterospecific haplotypes were found in 22.3 and 12.8% of A. alosa and A. fallax individuals, respectively, consistent with backcrossing and suggesting that hybrids are fertile. On average, microsatellite-identified hybrids comprised 12.7% of all samples, but when individuals with cytonuclear discordance were also considered introgressed on average 25.4% of individuals were of hybrid ancestry. Overall, allelic richness remained largely unchanged within species, but there were declines in the inbreeding coefficient (F _IS) of both species and episodes of significant temporal allelic frequency change. Hybrids sampled between 2004 and 2008 showed no evidence of lower fecundity relative to purebred individuals. Together, results suggest that hybridization between shad species in northern Europe is prevalent, and has been ongoing over at least two decades. The challenge is now to understand the extent to which observed patterns are linked to immigration from other populations, and the mechanisms that have prevented species collapse despite apparent hybrid fertility and longstanding introgression of neutral markers.     

Isolation and characterization of eight dinucleotide microsatellite loci from two closely related clupeid species (Alosa alosa and A. fallax)

Eight dinucleotide microsatellite loci were developed through an enrichment protocol for allis shad (Alosa alosa) and twaite shad (A. fallax). Cross‐species amplification was successful for all loci isolated. The number of alleles per locus ranged from three to nine for A. alosa and from two to seven for A. fallax, while the observed heterozygosity ranged from 0.267 to 0.926 and from 0.240 to 0.727, respectively. These markers will constitute useful tools for studies of population structure and gene flow between two closely related hybridizing species.     

Observations on the ecology of a landlocked population of allis shad in Aguieira Reservoir, Portugal

A landlocked population of the anadromous clupeid Alosa alosa from Aguieira Reservoir in Portugal is described. The individuals are fast-growing, mature in their third year of life, live for 5+ years, and consequently spawn more than once.     

Circadian rhythms in juvenile american shad, Alosa sapidissima

Swimming activity (in cm s⁻¹) of a school (55 individuals) of young-of-the-year ( total length=110 mm) American shad, Alosa sapidissima , was determined under a variety of photoperiod conditions. These included a normal (ambient), a shifted, and constant-light day. Swimming activity was measured over 4-day periods. During normal days swimming speeds followed periods of about 24 h, with fast speeds (up to 45 cm s⁻¹) and schooling occurring during the photoperiod. Under dark conditions speeds were slower (8 cm s⁻¹) with most fish swimming as individuals. During a shifted day swimming speeds and schooling corresponded to the imposed day. Under constant light (equivalent to bright moonlight) no schooling was evident, and a constant, but slow, swimming speed was observed in each 24-h period. These shad demonstrated an exogenous rhythm with respect to the imposed day length. It is hypothesized that an endogenous circadian rhythm would only be of use to a fish required to hunt or chase its prey. Shad, being plankton feeders, do not chase prey and therefore can exhibit an exogenous circadian rhythm with no detrimental feeding results.     

Proximate composition,lipid utilization and validation of a non‐lethal method to determine lipid content in migrating American shad Alosa sapidissima

Lipid content forms the most important energy reserve in anadromous fish and can limit survival, migration and reproductive success. A fat meter was evaluated and compared with a traditional extractive method of measuring available lipid for migrating American shad Alosa sapidissima in the Connecticut River, U.S.A. The fat meter gives rapid (<10 s) and non‐lethal lipid measurements, whereas traditional methods require lethal sampling that is both time consuming and expensive. The fat‐meter readings had a strong relationship to traditional lipid extractions for 60 fish, 30 whole body (R² = 0·72) and 30 fillet only (R² = 0·81). Additional validation showed that fat‐meter readings captured the gradual decrease of lipid in individual fish over time, were not affected by removal of gonads or scales and were stable for fish exposed to water or air for 24 h after death. These experiments indicate that the fat meter can be used as a reliable tool for future A. sapidissima energetic studies, allowing for larger sample sizes and non‐lethal sampling.     

Homing behaviour facilitates subtle genetic differentiation among river populations of Alosa sapidissima: microsatellites and mtDNA

Significant but subtle differentiation was detected for both microsatellite DNA and mitochondrial DNA among four populations of American shad Alosa sapidissima . The data indicate that straying among rivers is sufficient to permit only marginal population differentiation in this species, but suggest that individual river populations should be managed as distinct stocks. Comparison of the Hudson and Columbia populations, the latter derived from the former over 100 years ago, revealed only a slight reduction in microsatellite DNA variation for the founded population but halving of mitochondrial DNA, consistent with the haploid maternal inheritance of the latter marker. The depleted and endangered James River (Virginia) population and two other Atlantic coast populations exhibited similar levels of microsatellite DNA variation, but mtDNA diversity in the James River was marginally lower than in other Atlantic populations, again consistent with the low effective population size of mtDNA.     

Retina-specific LDH isozyme in blueback herring, Alosa aestivalis (Mitchell), and alewife, Alosa pseudoharengus (Wilson).

The retina of 390 Alosa aestivalis and 410 Alosa pseudoharengus have been examined by means of starch-gel electrophoresis. The retina-specific E4 isozyme has been found to occur in all the fish examined. This study demonstrates for the first time that the E4 isozyme occurs in A. aestivalis. Because the E4 isozyme is not polymorphic and has an identical mobility in A. pseudoharengus and in A. aestivalis it is neither suitable for use as a species identification characteristic nor a population marker.     

Biology and traits of the formation of stock of big-eyed shad Alosa saposchnikowii

Special traits of distribution, reproduction, growth, feeding, and population abundance of the big-eyed shad Alosa saposchnikowii are discussed. In investigations of the formation of abundance, the focus is on the strategy of observations on the stock in the absence of fishery statistics data. The informational representativeness of two monitoring methods is noted: trawl survey of the annual abundance of underyrearlings and census of the relative abundance of spawners recalculated per 100 nets. The big-eyed shad was a secondary commercial species. At present it prospers and occupies a leading position among herrings. In the observed period of sea transgression, in the northern Caspian Sea where principal spawning grounds of herrings are situated, the freshened zone has considerably widened, with conditions favorable to the spawning of big-eyed shad. As a result, its abundance has increased. In spite of the increase in species abundance, its linear and weight growth also increased. The latter is attributed to the change in the diet of the big-eyed shad, namely, its transition to consumption of food consisting mainly of fish.     

Retrospective approach to investigating the early life history of a diadromous fish: allis shad Alosa alosa (L.) in the Gironde–Garonne–Dordogne watershed

In this study, the identification of major ontogenetic events and the duration of the fresh water and estuarine phases before the seaward exit of young allis shad Alosa alosa were investigated retrospectively, using the otoliths of spawners. Daily growth increment periodicity in the otoliths was validated with an accuracy of 4 days (9·4%) for fish >30 days. The specific limits of the Sr:Ca ratios in the otoliths, indicating fish movements between salinity 'compartments', were up to 1·1 ×10⁻³ for the freshwater phase, 1·1 to 1·9 × 10⁻³ for the estuarine phase and 1·9 × 10⁻³ for the marine phase. In early life history, a major event occured at 22 days, which was marked on the otoliths by a change in the direction of the otolith growth axis. This event was related to a change in larval swimming behaviour. Juvenile freshwater residence time was between 58 and 123 days, with a median duration of 88 days. Contrary to what has been estimated in previous studies, the present study showed that the estuarine phase lasted from 4 to 36 days, with a median value of 11 days. Seaward exit occurred in fish aged from 63 to 150 days, with a median age of 99 days. These results have important implications for the sustainable management of this species.     

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.     

Effects of prey density and short term food deprivation on the growth and survival of American shad larvae

Growth and survival of 16-and 18-day-old American larvae ( Alosa sapidissima ) increased with prey density (0, 500, and 1000 Artemia l⁻¹), though differences were not always significant. Significant differences were not observed in growth parameters and survival for larvae fed 500 v . 1000 Artemia 1⁻¹ in laboratory experiments. Food deprivation for as little as 2 days had significant effects on survival but growth effects were not detectable until 4 days of starvation. These findings have application to the Susquehanna River where c . 11 million 2.5-week-old shad larvae are released annually as part of a basin wide effort to restore the species. Larvae released at upriver sites where food may be limiting would drift for 2–3 days to reach reservoirs with higher prey densities. Based on these laboratory findings, survival of shad may be reduced significantly if larvae do not feed prior to reaching reservoir environments.     

Interspecific differentiation and intraspecific substructure in two closely related clupeids with extensive hybridization, Alosa alosa and Alosa fallax

In this study, the evolutionary relationships within and among populations of European shads, Alosa alosa and Alosa fallax , was investigated. Screening of allelic variation across eight allozyme loci and sequencing 448 bp of the mtDNA cytochrome b gene in 14 rivers throughout the range of the species supported that the two taxa were independent lineages (1·3% net nucleotide divergence) despite extensive hybridization. Genetic diversity and structure was considerably higher in A. fallax than A. alosa and the former species revealed evidence of distinct lineages in the Mediterranean and Atlantic basins. A Bayesian clustering approach combined with gill raker counts verified that individuals of the two species could be assigned to their parent group with relatively high confidence. Evaluation of hybridization in the Lima and Mondego Rivers in Portugal provided evidence that introgression is extensive but is not currently obscuring (through hybrid swarming) the diagnosability of the two species.