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.     

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.     

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.     

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.     

Tolerance of young allis shad Alosa alosa (Clupeidae) to oxy-thermic stress

The ecology of the young stages of allis shad Alosa alosa is poorly documented, although they can be exposed to many pressures during their freshwater phase and their downstream migration. When passing through systems such as the Gironde-Garonne-Dordogne watershed (GGD, SW France), they can be subjected to high temperatures and low levels of oxygen (hypoxia). The aim of this work is to assess the tolerance of young Alosa alosa at four ages (c. 10, 30, 60 and 85 days old) by challenging them to different temperatures (18, 22, 26 and 28°C) together with decreasing oxygen saturation levels (from 100% to 30%). Survival of the 10-day-old individuals was not influenced by oxy-thermic conditions, but high stress levels were detected and perhaps this age class was too fragile regarding the constraint of the experimental design. Survival at 30 and at 60 days old was negatively influenced by the highest temperatures tested alone (from 26°C and from 28°C, respectively) but no effect was detected at 85 days old up to 28°C. A combined effect of temperature and oxygen level was highlighted, with heat accelerating survival decrease when associated with oxygen level depletion: essentially, survival was critical (<50%) at 30 days old at temperature ≥22°C together with 30% O₂; at 60 days old, at temperature = 28°C with 30% O₂; at 85 days old, at temperature ≥26°C with ≤40% O₂. Tolerance to oxy-thermic pressures appeared to be greater among the migratory ages (60 and 85 days old) than among the 30-day-old group. Based on environmental data recorded in the GGD system and on our experimental results, an exploratory analysis allowed a discussion of the possible impact of past oxy-thermic conditions on the local population dynamics between 2005 and 2018. The oxy-thermic conditions that may affect Alosa alosa at ages when they migrate downstream (60 and 85 days old) were not frequently recorded in this period, except in cases of extreme episodes of heat together with hypoxia that occurred in some years, in summertime in the turbidity maximum zone of the Gironde estuary (particularly in the year 2006). Interestingly, oxy-thermic conditions that are likely to threaten the 30-day-old individuals occurred more frequently in the lower freshwater parts of the GGD system between the years 2005 and 2018. In the context of climate change, a general increase in temperature is predicted, as well as more frequent and severe hypoxic events, therefore we suggest that local Alosa alosa population recruitment could encounter critical oxy-thermic conditions more frequently in the future if no adaptive management of water resources occurs.     

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.     

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     

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。     

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.     

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.     

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.     

Late migration and seawater entry is physiologically disadvantageous for American shad juveniles

Juvenile American shad Alosa sapidissima were subjected to isothermal transfers into sea water (salinity 24)‘early’(1 September; 24° C) and ‘late’(10 November; 10° C) in the autumn migratory season. Early acclimation resulted in a modest osmotic perturbation that recovered rapidly. Haematocrit declined by 14% at 24 h, recovering within 48 h. Plasma osmolality increased by 6% at 4 h, recovering within 8 h. Early acclimation caused a two‐fold increase in gill Na⁺, K⁺‐ATPase activity by 24 h and a four‐fold increase by 4 days. The number of chloride cells on the primary gill filament increased two‐fold by 4 days. Chloride cells on the secondary lamellae rapidly decreased from 22 cells mm^?1 to <2 cells mm^?1 within 4 days. Late acclimation resulted in a severe and protracted osmotic perturbation. Haematocrit levels declined by 23% at 4 days, recovering by 14 days. Plasma osmolality increased by 36% by 48 h, recovering by 4 days. Initial gill Na⁺, K⁺‐ATPase activity was two‐fold greater than in ‘early’ fish and did not change during acclimation. Initial numbers of chloride cells on the primary filament were two‐fold greater than ‘early’ fish and did not increase during acclimation. Initial number of chloride cells on the secondary lamellae was five‐fold greater than ‘early’ fish (116 v. 22 cells mm^?1) and declined to negligible numbers over 14 days. Differences between initial measures for ‘early’ and ‘late’ fish reflect previously described physiological changes associated with migration. These data indicate that late migrants face a greater physiological challenge during seawater acclimation than early migrants. Physiological performance apparently limits the observed duration of autumnal migration.     

Primary and secondary oocyte growth dynamics in anadromous semelparous Allis shad Alosa alosa

We analysed the ovarian dynamics of the anadromous semelparous allis shad Alosa alosa for which our working hypothesis was that mature pre-spawning females would have very low or even exhausted primary growth (PG) oocyte reserves; semelparity has been linked with the depletion of the pool of PG oocytes. To test this hypothesis, the PG oocytes were enumerated, their recruitment pattern to the secondary growth (SG) phase was analysed and their potential replenishment from the pool of oogonia was examined in females caught very close to the Mondego River mouth, in central Portugal and along the river. The development of the SG oocytes was also analysed, the fecundity (batch, total and annual) values were estimated and the intensity of atresia was quantified. Ovarian samples and histological sections were investigated in parallel. A dynamic recruitment pattern of PG oocytes to the SG phase was revealed, where all PG oocytes were recruited and were not replenished by oogonia. Annual fecundity was subject to down-regulation due to atresia prior to spawning and its size was multiple times higher than the size of batch fecundity. Lack of population synchronicity in ovarian development and spawning migration was also observed. This multifaceted analysis of the ovarian dynamics of this species will contribute to management efforts for this critically endangered and economically important fish throughout its geographical distribution. The results reported in this study will also assist in unravelling the complexity of the early processes of oogenesis in fish.     

Changes in haematology during upstream migration to American shad

Heart mass of American shad Alosa sapidissima did not change during migration in the Connecticut River. Spleen mass decreased and there was an increase in available blood haemoglobin (+22%) and haematocrit (+9%). The decreases in spleen somatic index (-29%) and spleen haemoglobin content (-15%) were dependent upon distance travelled upriver and not seasonal migration timing or short-term exercise events such as passage up a fish ladder. There was no effect of migration timing on any of the blood parameters measured, suggesting that any physiological responses during migration were based on distance travelled rather than seasonally variable conditions such as temperature, although blood haemoglobin (+24%) and haematocrit (+21%) increased after passage up a fish ladder. These changes in haematological physiology occurring during upstream migration may increase swimming performance and migratory success in American shad.     

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.     

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.     

Spawning behaviour of Allis shad Alosa alosa: new insights based on imaging sonar data

Spawning behaviour of Alosa alosa was observed by high resolution imaging sonar. Detected clouds of sexual products and micro bubbles served as a potential indicator of spawning activity. Peak spawning time was between 0130 and 0200 hours at night. Increasing detections over three consecutive nights were consistent with sounds of mating events (bulls) assessed in hearing surveys in parallel to the hydro acoustic detection. In 70% of the analysed mating events there were no additional A. alosa joining the event whilst 70% of the mating events showed one or two A. alosa leaving the cloud. In 31% of the analysed mating events, however, three or more A. alosa were leaving the clouds, indicating that matings are not restricted to a pair. Imaging sonar is suitable for monitoring spawning activity and behaviour of anadromous clupeids in their spawning habitats.     

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)

     

Factors related to gizzard shad and the threadfin shad occurrence and abundance in Florida lakes

Gizzard shad Dorosoma cepedianum were collected in 23 and threadfin shad D. petenense were collected in 22 of the 60 Florida lakes sampled. Logistic regression equations were 94% effective for predicting gizzard shad occurrence from chlorophyll and lake surface area, and 84% effective for predicting threadfin shad occurrence from lake surface area and lake volume inhabited (PVI). Occurrence of both shad species was related positively to lake size. In lakes where gizzard shad or threadfin shad were collected, shad density and biomass of both shad species were related positively to chlorophyll. Gizzard shad populations were generally vulnerable to predation in lakes, with the per cent of gizzard shad ≤200mm L _T values exceeding 60% with few exceptions. Effects of gizzard shad and threadfin shad on fish community dynamics may be confined to relatively large (>100 ha) and fertile (chlorophyll >20–30μg l⁻¹) Florida lakes.