The diet of juvenile and adult twaite shad Alosa fallax fallax (lacépède) from the rivers Severn and Wye (Britain)

The diet of juvenile and adult twaite shad was studied at a number of freshwater and estuarine sites in the rivers Severn and Wye and in the coastal waters of Cardigan Bay (West Wales). In the Severn estuary and in the freshwater reaches of the Severn and Wye, adult twaite shad consumed little during their pre-spawning migration. In the Severn Estuary post-spawning adults were present during the summer months where they fed actively on mysids, gammarids and shrimps. Mysids dominated the diet of the adults caught in coastal waters during the autumn. The diet of one year old twaite shad was studied in the Severn Estuary. In May the one year olds from the lower estuary consumed mainly gammarids. During the summer months, this age-class moved into the inner estuary where they fed predominantly on mysids.The larvae and juveniles (age 0⁺) fed mainly on chironomid larvae and pupae and Simuliidae larvae, while in fresh water. In the estuary copepods, cladocerans and mysids dominated the diet of the juveniles. There was some suggestion of an increase in prey size during their period of residency in the estuary. The contrasting distribution of the juveniles in two river systems is discussed in relation to the effects of channelization.     

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.     

Climate change and the green energy paradox: the consequences for twaite shad Alosa fallax from the River Severn, U.K

A stock-recruitment model with a temperature component was used to estimate the effect of an increase in temperature predicted by climate change projections on population persistence and distribution of twaite shad Alosa fallax. An increase of 1 and 2° C above the current mean summer (June to August) water temperature of 17·8° C was estimated to result in a three and six-fold increase in the population, respectively. Climate change is also predicted to result in an earlier commencement to their spawning migration into fresh water. The model was expanded to investigate the effect of any additional mortality that might arise from a tidal power barrage across the Severn Estuary. Turbine mortality was separated into two components: (1) juvenile (pre-maturation) on their out migration during their first year and on their first return to the river to spawn and (2) post-maturation mortality on adults on the repeat spawning component of the population. Under current conditions, decreasing pre-maturation and post-maturation survival by 8% is estimated to result in the stock becoming extinct. It is estimated that an increase in mean summer water temperature of 1° C would mean that survival pre and post-maturation would need to be reduced by c. 10% before the stock becomes extinct. Therefore, climate change is likely to be beneficial to populations of A. fallax within U.K. rivers, increasing survival and thus, population persistence.     

Larval fish production in the tidal River Elbe 1985–1986

In 1985 and 1986 ichthyoplankton composition was studied during seven sampling series at 37 stations along the tidal Elbe and its tributary rivers between Hamburg harbour and the North Sea. The highest larval density was observed in May and April. No, or very few, larvae occurred from late summer to early spring. Smelt (59% on average of all samples) and twaite shad (20%) dominate in the freshwater region. Herring/sprat (17%) dominate in the estuary. All major spawning grounds are located along the southern shore. Smelt spawn further upstream 1 month earlier than twaite shad. Three percent of all larvae found were flounder. Most of them occurred upstream from the estuary in the R. Este and in other tributary rivers. It is not clear whether flounders are able to spawn in freshwater. Attention is drawn to the fact that an unknown but probably very high percentage of the ichthyoplankton is entrained by local power plants.     

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.     

Return of twaite shad <Emphasis Type="Italic">Alosa fallax</Emphasis> (Lacépède, 1803) to the Southern Baltic Sea and the transitional area between the Baltic and North Seas

The status of twaite shad was investigated within the Southern Baltic Sea (ICES subdivision 22–27) and transitional area between the Baltic and North Seas (division IIIa). The following sources of data were analysed: (i) commercial catch statistics and relevant publications, (ii) records from ichthyological museum collections, (iii) records from recent commercial and recreational fisheries, (iv) research fisheries with trawls. A total of 476 records of twaite shad including more than 16 million individuals were obtained for the time between the years 1836 and 2005. About 72.9% of all records originated from commercial catch statistics and publications, whereas 18.9% were received from ichthyological collections. Research fisheries provided 6.3%, and 1.9% of the records were obtained from recent commercial and recreational fisheries. Most records of twaite shad were estimated for subdivisions 24 (45.2%) and 26 (35.5%). From 1836 to 1959, 29.6% of the records date from the period until 1899. 70.4% of the records of twaite shad originate from the twentieth century until 1959. The mean annual catch of twaite shad between 1891 and 1959 amounted to 86,674 kg within subdivisions 24–26 of the Southern Baltic Sea. Catch data show an approximately 20-year-cyclicity of maximum yields and minimum catches, respectively. The maximum annual yield of twaite shad in subdivisions 24–26 (474,700 kg) was registered in 1940, the minimum annual yield was estimated in 1958 (10 kg). In the 1950s, the annual catches of twaite shad declined sharply. Until 1960 twaite shad catches and records originated mainly from the Pommeranian Bay/Pommeranian Coast and adjacent waters including the Szczecin Lagoon (subdivision 24), the Bay of Gdańsk, Vistula Lagoon and Vistula Spit (subdivision 26) and from the Curonian Lagoon and Curonian Spit (subdivision 26). The highest catches of twaite shad originated from the area of Curonian Lagoon/Curonian Spit from 1941 to 1960. Seasonal catches of twaite shad showed maximum values from May to July. From 1960 to 1989, only four records of A. fallax were registered in the Southern Baltic Sea which originated from German coastal waters in subdivisions 22 and 24. A total of 107 records of twaite shad was obtained in subdivisions 20–27 from 1990 to 2005. Most of these recent records originate from a twaite-shad stock in subdivisions 24, 25 and 26. There are indications which suggest a separate stock of twaite shad in subdivisions 20 and 21. Disappearance and recovery of twaite shad stocks were probably caused by the following factors: construction of barriers in rivers with spawning sites of twaite shad; habitat destruction in those rivers as consequence of gravel extraction and reengineering scheme to improve navigation and for flood defence purposes, water pollution in the lagoons of the Southern Baltic and in their tributaries, commercial fishery in the Southern Baltic and climatic variation in the Baltic Sea basin south of the latitude of 60° N. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest editors: S. Dufour, E. Prévost, E. Rochard & P. Williot Fish and diadromy in Europe (ecology, management, conservation)     

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.     

Growth and abundance of O-group herrings, Clupea harengus L., in the Severn Estuary

Collections made from the cooling water intake screens of Power Stations in the Severn Estuary and the Bristol Channel have been used to investigate the biology of O-group herrings in this region. The abundance of herrings at Oldbury, the principal sampling site, was low in most years and in five successive seasons between July 1972 and December 1976, large numbers were caught only between the summer of 1975 and the spring of 1976. The herrings were first observed in July, when many had still not completed metamorphosis, and reached peak numbers in September and October during which period they were clearly growing. Numbers declined in December before rising dramatically in January at which time the length-frequency curves displayed a pronounced bimodality. On the basis of data from other parts of the Bristol Channel and Severn Estuary, it is suggested that this bimodality is due to the influx of some animals from other regions where the growth rate was greater. Although movement is apparently taking place at this time and the numbers decline markedly in subsequent months, the last remnants of the population do not leave the estuary until early May. Post-pelvic scute (K₂) and vertebral counts (VS) of 13.82 and 55.24 respectively, indicate that the Severn Estuary herrings are the product of a spring spawning stock, a view entirely consistent with length and field data.     

Environmental factors as forces structuring the fish community of the Elbe Estuary

Smelt, flounder, eel, three-spined stickleback, twaite shad, ruffe and herring were the major species among 62 forming the fish community of the Elbe estuary. Species richness, species diversity, evenness and total fish biomass decreased in the upstream direction. Total abundance was greatest during summer. Salinity was the most significant physico-chemical factor affecting species richness and total fish biomass, whereas water temperature was the best predictor of total abundance. Marine fish species decreased with decreasing salinity, while the freshwater species roach and ide were absent at salinities > 15‰. Bream, blue bream, white bream, perch and pikeperch occurred more frequently in shallow marginal regions with lower current velocities, while herring and other marine species occurred closer to the deep central regions with high current velocities. Annual and seasonal variations of the community structure mainly reflected in population dynamics of smelt, flounder, twaite shad, three-spined stickleback and eel, especially those of 0-age smelt. Populations of smelt and ruffe and their importance in the fish community increased between 1989 and 1992, while those of twaite shad and eel decreased. At oxygen concentrations <3 mg I¹, smelt and flounder were rare, but eels were at their maximum at 3.5 mg I⁻¹. Changes of species composition and fish abundance were closely related to daytime and tidal cycle.     

Poor water quality constrains the distribution and movements of twaite shad <Emphasis Type="Italic">Alosa fallax</Emphasis><Emphasis Type="Italic">fallax</Emphasis> (Lacépède, 1803) in the watershed of river Scheldt

Worldwide, river fragmentation is primarily responsible for the decline of populations of migrating fish. In particular, anadromous fish species, which necessarily migrate to fresh water to reproduce, are endangered since many are no longer able to reach their natural spawning sites. In addition, pollution of rivers effectively prevents upstream or downstream movements and blocks access to spawning grounds. This article investigates how poor water quality interferes with the life history cycle of twaite shad Alosa fallax fallax (Lacépède, 1803), an anadromous clupeid fish, in the watershed of River Scheldt, a heavily impacted environment in West Europe. We used two models based on known ecological and environmental information to explain past and present twaite shad distribution within the watershed and to make inferences about a future population recovery and juvenile habitat value. We demonstrated that historical spawning areas satisfy water quality conditions necessary to support spawning and successful development of early life history stages of the twaite shad. However, poor water quality conditions just upstream the freshwater–saltwater boundary still act as an effective migration barrier for upstream movement. As a consequence, spawning grounds are inaccessible and the population is dominated by seasonal adults occurring in the lower estuarine part of the watershed. This article provides testable and diagnostic information to the watershed management in that it identifies habitat and water quality requirements needed to support the expected recovery of an endangered anadromous fish population. Guest editors: S. Dufour, E. Prévost, E. Rochard & P. Williot Fish and diadromy in Europe (ecology, management, conservation)     

The effect of the migration of Alosa fallax fallax (Laécpède) into fresh water, on branchial and gut parasites

The prevalence and relative density of branchial and gut parasites taken from Almafullax in the estuary of the River Severn, at the start of the freshwater phase of its spawning migration, were compared with those taken from their spawning grounds in fresh water. Mazocrues alosue showed no significant difference in prevalence between the two habitats, though their relative density did fall significantly. Because of the direct relationship between length and relative density, this difference may be explained by the estuary samples of A. fullux being of fish significantly larger than those from fresh water. Other possible causes are discussed. The prevalence and relative density of M. alosae showed a trend to increase with age of the host. Repeat spawning A. fallax had a significantly greater relative density and prevalence of M. alosue when compared to virgin fish.
Analysis of the sampled population showed that Clavellisu emarginata declined significantly in both prevalence and relative density in fresh water, and this was confined to the younger and first time spawning A. fullux. Thynnuscuris aduncum showed significant reductions in prevalence and relative density of infection in fresh water when compared to the estuarine samples.
The species Hemiurus uppendiculatus and Pomphorhynchus laevis showed no significant difference in prevalence and relative density of infection, and Pronoprymna ventrzcosa no significant difference in prevalence, between the two habitats.     

Spatial Analysis of Twaite Shad,Alosa fallax (Lacepède, 1803), in the Southern North Sea: Application of Non‐Linear Geostatistics as a Tool to Search for Special Areas of Conservation

This study aims to evaluate the suitability of non‐linear geostatistics and indicator kriging (IK) as a tool in environmental impact assessment and nature conservation, in particular to search for potential Special Areas of Conservation (SAC) for the endangered fish species twaite shad, Alosa fallax (Lacepède , 1803) within the German Exclusive Economical Zone (EEZ) of the North Sea. To analyse the spatial distribution of this fish species, data on standardised biomass index (catch per unit effort, c.p.u.e., kg × 30 min^–1) from 1996 to 2001 were used, regarding the third and fourth quarters of each year, respectively. Thereby we assume that the spatial distribution can be described as a time‐invariant process. This assumption is supported by information on annual sampling effort, allocation of hauls and spatial distribution of the positive catches. All indicator variograms obtained for different c.p.u.e. cut‐off values displayed distinct spatial structures, clearly indicating that the indicator variables were spatially autocorrelated. Gaussian models were fitted by least‐squares methods and were evaluated with a goodness‐of‐fit statistic. Subsequently, IK was employed to estimate the probability of exceeding the c.p.u.e. cut‐off values for the twaite shad in the investigation area. These were highest in the Weser‐ and Elbe‐estuary, probably because of migrations of twaite shad to and from estuaries at the time of investigation due to spawning, while within the German EEZ of the North Sea no such areas with increased probabilities could be discerned. Thus, although available data did not allow to identify and implement any SAC in the German EEZ, the methods employed here can be regarded as a promising management tool in biological conservation issues. (© 2004 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Migration behaviour of twaite shad Alosa fallax assessed by otolith Sr:Ca and Ba:Ca profiles

Individual migration behaviour during the juvenile and adult life phase of the anadromous twaite shad Alosa fallax in the Elbe estuary was examined using otolith Sr:Ca and Ba:Ca profiles. Between hatching and the end of the first year of life, juveniles showed two migration patterns. Pattern one exhibited a single downstream migration from fresh water to the sea with no return into fresh water. In contrast, pattern two showed a first migration into the sea, then a return into fresh water and, finally, a second downstream migration into marine water. This first report of migration plasticity for A. fallax points to different exposure times to estuarine threats depending on the migration strategy. In adults, high Sr:Ca and low Ba:Ca in the majority of individuals confirmed prior reports of a primarily marine habitat use. Patterns reflecting spawning migrations were rarely observed on otoliths, possibly due to the short duration of visits to fresh water.     

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.     

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.     

The evolution of the fine sediment regime of the Severn Estuary and Bristol Channel

Fine sediment forms tidal flats along most of the Severn Estuary coastline but subtidal deposits are localized, mainly confined to Newport Deep and Bridgwater Bay. The estuary is experiencing a period of sea level rise which has led to coastal mud erosion in recent centuries. Sediment exchanges with the sea and river inputs are negligible compared to the large-scale exchanges within the estuary itself. Measurements of tidal flat level changes, archaeological finds and anthropogenic chemical concentrations confirm that mud flat erosion is long term and typical of the entire estuary. Sea level rise and coast erosion would normally lead to much of this sediment being redeposited inshore at the landward limit of the alluvium. Other than in narrow salt marsh strips, such deposition is prevented in the Severn by the river walls. In cores of subtidal muddy sediment, the number and thickness of sand layers decreases upwards. The top metre or so is entirely mud and exhibits thousands of layers of varying thickness and origin. Radiochemical analysis confirms that mud is accumulating in the subtidal zone of Newport Deep and the seaward periphery of the Bridgwater Bay mud patch and is increasing in proportion to sand. Circumstantial evidence strongly suggests that for at least the last 600 years the dominant pattern has been one of erosion of fine sediment from the coastal margin and its accumulation in subtidal sinks.     

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.     

Intertidal zonation of animals and plants on rocky shores in the Bristol Channel and Severn Estuary–the northern shores

The vertical distribution of common species of macroalgae and fauna on rocky shores extending through the length of the estuary is described. In the west, the shores are fully marine and exposed to Atlantic waves. In an eastward direction, greater shelter and an increasing tidal range occur along a gradient of salinity and turbidity. The flora and fauna change along this gradient. No single biological feature signals a transition from the Bristol Channel to the Severn Estuary but four arbitrary regions are recognized: a 'marine' section eastwards to Swansea, a 'transitional' section between Swansea and Cardiff, an 'estuarine' section up to Newnham, and the tidal river Severn.     

Threadfin shad, Dorosoma petenense Günther, mortality: causes and ecological implications in a South-eastern United States reservoir

Cold stress was identified as an important factor influencing both reservoir-wide mortality and impingement of threadfin shad, Dorosoma petenense , during the period October 1976 to April 1977 in Watts Bar Reservoir, Tennessee. Relative numbers and size frequency of impinged threadfin shad were similar to the relative numbers and size frequency of shad preyed upon by sauger, Stizostedion canadense , and skipjack herring, Alosa chrysochloris . This relationship implies that the factor mainly responsible for impingement, low temperature, also influences prey vulnerability. Threadfin shad made up 99% of the combined diet of sauger and skipjack herring from November until the threadfin shad disappeared in January. These predators did not readily switch to alternative prey in the short term, but by the next autumn 25–100% of the diet was alternative prey. Reappearance of threadfin shad the year following mass mortality and ability of the predators to vary their diet emphasize the resilient nature of some predator-prey systems.     

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.