Genetic polymorphism of adenosine deaminase (ADA; E.C. 3.5.4.4.) in allis shad, Alosa alosa and twaite shad, Alosa fallax

A genetic polymorphism of adenosine deaminase (ADA), is described for the first time in European shad species, Alosa alosa and Alosa fallax . Significant heterogeneity in gene frequencies was found, both between species and within species.     

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.     

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.     

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.     

Photoresponse in allis shad larvae

Allis shad Alosa alosa photoresponse was investigated in 3 day-old to 1 month-old larvae. Regardless of age, larvae strongly preferred the lit compartment in aquaria experiments. Such response was consistent with other experimental studies which have suggested a pelagic behaviour of allis shad larvae from the time of hatch.     

Genetic polymorphism of a haemoglobin chain and adenosine deaminase in European shads: evidence for the existence of two distinct genetic entities with natural hybridization

The degree of genetic differentiation between European shads, Alosa alosa and A. fallax , was studied in populations from different Portuguese hydrographic basins. Using isoelectric focusing, polymorphic variation was detected in a haemoglobin (HB) chain and in adenosine deaminase (ADA). These polymorphisms are shared by the two species but show significant differences in their gene frequency distributions. Individuals with intermediate morphological characteristics were found to exhibit intermediate allele frequencies. While not completely excluding introgression, our results strongly support the existence of two distinct species that can hybridize.     

Observations sur l'activité de migration de la grande alose Alosa alosa L. en Loire (France)

We studied the migratory activity of shad (Alosa alosa Linné) in the middle part of the Loire river, using stop net catches, in 1984.The migration started in the beginning of April, instead of February, as more usual. The end of migration is not precisely known, because of adverse hydrological conditions.During migration, the shad principally used the channel with the highest current velocity. The migration was disturbed by the presence of obstacles (weirs, dams of nuclear power stations).The daily and hourly activity of migration was strongly correlated with variations in water temperature. Shads did not run up at temperatures below 11°C. The upstream limit of ditribution was situated at more than 500 km from the estuary.

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Observations sur l'activité de migration de la grande alose Alosa alosa L. en Loire (France)

     

The occurrence of Eubothrium fragile (Cestoda: Pseudophyllidae) in twaite shad, Alosa fallax (Lacépède) in the River Severn

The occurrence, size and maturity changes of Eubothrium fragile have been studied in postlarvae, juveniles and adult twaìte shad, Alosa fallax , from several locations in the River Severn. Parasites were only found in adult shad and not in post-larvae or juveniles. No juvenile or recently acquired cestodes were identified as such, but adults were present in shad throughout the whole period of their spawning migration. A large proportion of the parasites were gravid upon arrival in the river and, although eggs were subsequently released into fresh water, there was no loss of cestodes from the fish. It was concluded that E. fragile is a marine species, that the parasites found in adult shad in fresh water were the residue of a marine life cycle and that the eggs released in fresh water were part of the parasite's natural reproductive wastage. The distribution and biology of E. fragile were discussed and it is considered that it is typical of the marine species of the genus.     

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.     

Weak population differentiation in northern European populations of the endangered anadromous clupeid Alosa fallax

Examination of genetic structure among different spawning populations of the endangered twaite shad Alosa fallax showed statistically significant but weak differentiation estimated using nine microsatellite loci. Results point to substantial gene flow among the majority of individual estuary/river spawning locations, and no evidence that habitat disruption has led to increased local genetic drift.     

Morphometric, meristic character and electrophoretic analyses of two Irish populations of twaite shad, Alosa fallax (Lacépède)

Since 1911 a rare lacustrine form of twaite shad, Alosa fallax killarnensis Regan (1916), has been known to occur in the Killarney Lakes. During the summers of 1985 and 1986 a gill netting programme in Lough Leane for brown trout, Salmo trutta L., yielded 32 shad (1985) and 64 shad (1986) as a by-catch. These fish were deep frozen and compared with a sample of 59 marine twaite shad, taken on rod and line from St Mullin's, Co. Carlow, which had entered the River Barrow to spawn.
Morphometric, meristic and electrophoretic character analyses were carried out on the two populations. The three analyses, particularly the isoelectric focusing, confirmed a high degree of genetic similarity. The two major differences found were the dwarfed size of the Killarney shad and the increased numbers of gill rakers carried on the first branchial arch. The merits and demerits of assigning the term subspecies to the Killarney shad are discussed in the light of the available evidence produced.     

Growth and movements of the twaite shad, Alosa fallax (Lacépède), in the Severn Estuary

Between July 1974 and April 1977, appreciable numbers of 0 group twaite shad, Alosa fallax , have been collected from the cooling water intake screens of the Nuclear Power Stations at Oldbury-upon-Severn and Berkeley on the Severn Estuary and at Hinkley Point on the Bristol Channel, England. Young of the year first appeared at Oldbury in July at a size of approximately 32 mm standard length, and grew to approximately 61 mm by October. The downstream migration in the late summer and autumn appeared to be closely correlated with water temperature since movement reached a peak soon after the temperature in the Estuary had declined below 19° C, and ceased altogether below 9° C. Thereafter, there was virtually no movement of shad until temperatures rose sharply above 7° C in the spring. Maturing adult twaite shad were captured in late April and May as they moved into freshwater to spawn, probably in mid-June. Males were recorded slightly earlier during the migration and were generally younger than females. Furthermore, mean lengths and weights for individuals belonging to the different age classes indicated that males were also smaller. Logarithmic relationships between length and weight of young and adult A. fallax and for the variation with size in the number of rakers on the first gill arch are presented.     

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.     

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.     

Genetic structure of the spawning population of Danube shad, Alosa pontica, Eichwaldt 1838 (Clupeiformes, Alosiinae)

The genetic structure of the population of Danube shad, Alosa pontica, has been studied by means of analysis of 3 polymorphic biochemical gene loci. The results of the study provide evidence of its genetic heterogeneity which is expressed by: firstly, an unbalanced ratio of genotype loci manifested by an excess of rare homozygotes and deficite of the correspoding heterozygotes, and secondly, differences in the allele frequencies between shad populations arriving for spawning in March-April and May-June. It is suggested that there could be two causes of heterogeneity: one due to the introgressive hybridization between various forms of A. pontica and another to the differences between early and late spring races of shad which to a certain extent are reproductively autonomous.     

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.     

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.     

Macrotidal estuaries: a region of collision between migratory marine animals and tidal power development

Macrotidal estuaries of the inner Bay of Fundy are utilized by large numbers of migratory fishes, particularly dogfish, sturgeon, herring, shad, Atlantic salmon and striped bass as well as by other migratory marine animals, many of which have large body sizes (squid, Lamnid sharks, seals and whales). Tagging experiments indicate the fishes originate from stocks derived over the entire North American Atlantic coast from Florida to Labrador. Population estimates suggest up to 2.0 times 10⁶adult American shad (Alosa sapidissima) migrate through an individual embayment each year. These migrations are an integral part of the life history of the respective species and appear to be controlled in part by the near shore movements of ocean currents. In other regions of the world similar macrotidal estuaries exist (Cook Inlet, Alaska; Severn Estuary, U.K.) and they, like the Bay of Fundy, are linked in continuum to the local ocean currents. We propose that marine animals utilize all these regions in a manner similar to the Bay of Fundy estuaries and properly designed surveys will reveal their presence. Fish passage studies utilizing the Annapolis estuary low-head, tidal turbine on the Bay of Fundy have shown that turbine related mortality of 20–80% per passage occurs depending on fish species, fish size and the efficiency of turbine operation. We suggest that introduction of tidal turbines into open ocean current systems will cause widespread impact on marine populations resulting in significant declines in abundance.     

Length–weight and length–length relationships for four shad species along the western Black Sea coast of Turkey

This study investigates length–weight and length–length relationships of four shad species (Alosa caspia caspia, Alosa immaculata, Alosa maeotica, Alosa tanaica) captured in the western Black Sea coast of Turkey. A total of 686 specimens, 355 females and 331 males, were caught by gill net, trammel net and longline from 2006 through 2007 along the western Black Sea coast of Turkey. The relationships of total length (TL), fork length (FL) and standard length (SL) are presented for inhabiting in the Black Sea four shad species and the relationships between total length (TL) and body weight (BW) from the two localities, Sile and Karasu in the western Black Sea coast of Turkey. The Length–weight relationships generally indicated positive allometric growth in Alosa species for both males and females and an isometric growth for A. immaculata females. The coefficient b ranged from a minimum 2.97 for females of A. immaculata to a maximum 3.75 for both sexes A. c. caspia. Results indicated that the length–length relationship between the three length measurements were highly correlated (r² > 0.99, P < 0.001).     

A review of the past and present status of anadromous fish species in the Netherlands: is restocking the Rhine feasible?

The paper reviews the past, present and future of eight anadromous fish species inhabiting the Lower Rhine (The Netherlands), viz. -sturgeon (Acipenser sturio), whitefish and houting (Coregonus lavaretus, C. oxyrinchus), smelt (Osmerus eperlanus), allis and twaite shad (Alosa alosa, A. fallax), sea trout (Salmo trutta) and salmon (Salmo salar). All species are under threat or became extinct (e.g. sturgeon, allis shad). It is not possible to single out a specific factor for the decline or disappearance. A combination of factors is responsible, as the degradation of the spawning and nursery areas, river correction for shipping, building of sluices and hydropower dams, extraction of sand and gravel and river pollution. The likelihood that a species will return via natural recovery, or restocking is assessed. The return of the sturgeon is unlikely. Present observations of sturgeon can be attributed to releases of unwanted sturgeon hybrids. A natural stock of coregonids in Dutch waters seems not feasible any more due to irreversible habitat degradation. Present day catches originate from German releases. The anadromous smelt, heavily reduced in numbers, still inhabits some of our waters, the non-migratory smelt is still very common. The allis shad is extinct and unlikely to recover. The species never spawned in the Dutch part of the Rhine. Twaite shad, declining in numbers, are still observed in the lower reaches of Rhine and Meuse. Sea trout is presumably still present in the same varying numbers as before. Spawning in our waters has not been documented. The salmon, once fished in large numbers, is now the subject of restocking programmes in Germany. Observations of individuals can partly be attributed to these programmes but also to straying salmon. Restocking programmes should be considerably improved before noticeable success is to be met.