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.     

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.     

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.     

Diet of the twaite shad Alosa fallax (Lacépède) (Clupeidae) in the River Tagus Estuary, Portugal

The diet of Alosafallax was studied by means of the frequency of occurrence of prey items. In the Tagus Estuary, shads feed preferentially on fish, with mysids, shrimp, isopods and insects occurring as secondary prey.     

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.     

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.     

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.     

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.     

Ecology of Allis Shad Alosa alosa and Twaite Shad Alosa fallax in the Solway Firth, Scotland

The basis of this study were 132 adult Allis Shad and 150 Twaite Shad collected as bycatches from salmon stake nets in Scotland on the north side of the Solway Firth. Most (60%) of the Allis Shad were immature fish 2–3 years of age (mean length 305 mm). Mature males were younger (3–5 years) and smaller (mean length 421 mm) than females (4–6 years and 481 mm mean length). The largest Allis Shad was a female of 515 mm and 2183 g. In contrast, almost all the Twaite Shad were mature, the males younger (3–5 years) and smaller (mean length 341 mm) than the females (4–6 years and 364 mm mean length). The largest Twaite Shad was a female of 400 mm and 1213 g. The food of Allis Shad consisted mainly of small zooplankton with some fish and larger Crustacea. Fine vegetable fragments were common in the stomachs and attributed to filter feeding. The food of Twaite Shad was mainly small fish with some Crustacea. Mature Allis Shad of both sexes with large gonads (maximum female GSI: 20.63) were found throughout the spring and summer but no definite evidence of local spawning was obtained. Mature Twaite Shad of both sexes with large gonads (maximum female GSI: 23.32) were found until early July, thereafter most of the fish were spent. It appears that Twaite Shad spawn locally in June. Some hybrids between the two species were found.     

The biology of the twaite shad, Alosa fallax fallax (Lacépède), in the Severn Estuary

The estuarine biology of the twaite shad was studied in the Severn Estuary. Adults enter the estuary at the start of the freshwater phase of their spawning migration between April and June. Peak immigration generally occurs in May and is associated with temperatures in the range 10.6–12.3°C. The mean (± s.d. ) instantaneous mortality rate for the mature population was 0.53±0.18. The effect of additional mortality on the spawning population was modelled assuming constant recruitment and no density-dependent effects.
Juvenile twaite shad are present in the estuary from July until they emigrate seaward during the autumn. A portion of these fish re-enter the estuary the following April–May and remain until late summer/early autumn before once more migrating seaward.
The 0 + age group feed mainly on harpacticoid and calanoid copepods and mysids, the relative preponderance of these in the diet being apparently related to tidal conditions. The possible implications of the proposed tidal power barrage in the Severn Estuary on the twaite shad population are discussed in relation to movement, diet and additional mortality of the mature population.     

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.     

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.     

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.     

Juvenile downstream migration patterns of an anadromous fish,allis shad (Alosa alosa), before and after the population collapse in the Gironde system,France

Diadromous fish have exhibited a dramatic decline since the end of the 20th century. The allis shad (Alosa alosa) population in the Gironde-Garonne-Dordogne (GGD) system, once considered as a reference in Europe, remains low despite a fishing ban in 2008. One hypothesis to explain this decline is that the downstream migration and growth dynamics of young stages have changed due to environmental modifications in the rivers and estuary. We retrospectively analysed juvenile growth and migration patterns using otoliths from adults caught in the GGD system 30 years apart during their spawning migration, in 1987 and 2016. We coupled otolith daily growth increments and laser ablation inductively-coupled plasma mass spectrometry measurements of Sr:Ca, Ba:Ca, and Mn:Ca ratios along the longest growth axis from hatching to an age of 100 days (i.e., during the juvenile stage). A back-calculation allowed us to estimate the size of juveniles at the entrance into the brackish estuary. Based on the geochemistry data, we distinguished four different zones that juveniles encountered during their downstream migration: freshwater, fluvial estuary, brackish estuary, and lower estuary. We identified three migration patterns during the first 100 days of their life: (a) Individuals that reached the lower estuary zone, (b) individuals that reached the brackish estuary zone, and (c) individuals that reached the fluvial estuary zone. On average, juveniles from the 1987 subsample stayed slightly longer in freshwater than juveniles from the 2016 subsample. In addition, juveniles from the 2016 subsample entered the brackish estuary at a smaller size. This result suggests that juveniles from the 2016 subsample might have encountered more difficult conditions during their downstream migration, which we attribute to a longer exposure to the turbid maximum zone. This assumption is supported by the microchemical analyses of the otoliths, which suggests based on wider Mn:Ca peaks that juveniles in 2010s experienced a longer period of physiological stress during their downstream migration than juveniles in 1980s. Finally, juveniles from the 2016 subsample took longer than 100 days to exit the lower estuary than we would have expected from previous studies. Adding a new marker (i.e., Ba:Ca) helped us refine the interpretation of the downstream migration for each individual.     

Extensive polymorphism at adenosine deaminase in the marine fish Sciaenops ocellatus (L.)

Summary Eleven different allelic variants at the adenosine deaminase (ADA) locus have been detected using vertical starch-gel electrophoresis among 474 individuals of the marine fish Sciaenops ocellatus (L.). Thirty-five of the 66 possible genotypes were observed, and the heterozygosity level at ADA was estimated to be 70·3%. The extensive polymorphism at ADA may prove useful in terms of providing genetic markers for stocking programs using hatchery-raised fish.     

The presence of anadromous twaite shad (Alosa fallax) in the Ebro River (western Mediterranean,Spain): an indicator of the population's recovery?

Although the Ebro River is one of the largest Mediterranean rivers, the presence of anadromous fish species was near to complete extirpation since the second half of the 20th century. However, many mature twaite shads (Alosa fallax) were captured at the lower reaches of this river (Xerta's weir site, 63 river km) in spring 2005, suggesting a natural recovery of this population. In this study, the first biological data on twaite shad from the Ebro River are presented, and their significance for the species conservation and recovery are discussed. The historical reasons for the decline of this species are reviewed, and data supports the proposal for urgent construction of a proper fish passage for anadromous species at the Xerta's weir. This action would allow fish access to a further 60 km of river upstream of the weir, representing a 200% increase in the amount of freshwater habitat and potential spawning areas. Improved access together with restrictive fishery regulations will assist with renewed interest in the fishery and conservation of the species.     

Two new common alleles of erythrocyte adenosine deaminase (ADA) in pigs

New adenosine deaminase variants ADA C and ADA D were found by means of agarose gel electrophoresis in pig erythrocytes. Family data supported the hypothesis that these are controlled by codominant alleles ADAC and ADAD. The ADAC allele was present in Large White (q = 0.076), Landrace (q = 0.037) and their crosses with other breeds. The ADAD allele was present in Duroc (q = 0.067) and its crosses. Allele frequencies for six pig breeds are given.     

Genetic polymorphism of adenosine deaminase and mannose phosphate isomerase in blood of arctic foxes, Alopex lagopus

Blood samples of 70 foxes, including 10 families, have been investigated by horizontal starch gel electrophoresis for the enzymes adenosine deaminase (ADA) and mannose phosphate isomerase (MPI). The observed variation of the enzymes could be explained as a result of one locus with two codominant alleles and one with three respectively. The segregation in the families of the alleles assumed for the two loci is in accordance with this genetic model. The frequency of the two alleles at the Ada locus is about the same and the slowest anodic migrating allele at the Mpi locus is the most frequent.