Aspects of anadromous Allis shad (Alosa alosa Linnaeus) and Twaite shad (Alosa fallax Lacépède) biology in four Irish Special Areas of Conservation (SACs): status, spawning indications and implications for conservation designation

Four Special Areas of Conservation (SACs) have been designated in the Republic of Ireland for Twaite shad, Alosa fallax (Lacépède), based on recent historical information on spawning activity—the estuarine reaches of the Rivers Munster Blackwater, Suir, Barrow–Nore and Slaney. The spawning status of some of these populations is considered to have declined substantially in recent years, and no known spawning areas of Allis shad, Alosa alosa (L.), are known for Ireland. Data are presented confirming the presence of both Allis shad and Twaite shad in all four SACs. The majority of the material has come as by-catch from estuarine commercial salmon netsmen. Angling is the principal source of material from the R. Barrow. Examination of scale and gonad material of Allis shad indicates some degree of readiness for spawning as well as spawning having taken place. Gonadosomatic index (GSI) data from the R. Barrow for Twaite shad in the May spawning period (2000–2002) had mean GSIs of 13.7–15% with maxima in the range of 21–28%. GSI values of 18% and 22% have been recorded in the July period for individual Allis shad from the upper estuaries of the R. Slaney and R. Suir, respectively, with both fish displaying gonad maturity stage V or VI (after Pina, 2000). Low GSI values have been recorded from other Allis shad in the June–July period displaying gonad condition varying from stage III (immature) to stage VIII (spent). These findings point to a possibility that Allis shad do engage in spawning activity, whether mono-specifically or with Twaite shad. The possibility of the latter is supported by gill raker count data presented. These show clearly defined value ranges for Twaite and for Allis as well as a series of fish with intermediate counts. The apparently low population levels of Twaite shad within the SACs identify a need to examine what measures might be undertaken to ensure the favourable conservation status of this species. Submitted for oral presentation to Fish Diadromy in Europe conference, 27.1.2005. Guest editors: S. Dufour, E. Prévost, E. Rochard & P. Williot Fish and diadromy in Europe (ecology, management, conservation)     

Genetic variations in Clupea pallasii herring from Sea of Okhotsk based on microsatellite markers

The genetic variations among spawning groups of herrings from different spawning grounds of the northwestern part of the Sea of Okhotsk was assessed using ten microsatellite loci. All loci were polymorphic with the expected heterozygosity estimates varying at different loci in the range of 0.7–95.0% (with a mean of 68.5%). The degree of genetic differentiation displayed by the herrings from the Sea of Okhotsk was not statistically significant (θ = 0.74%). The level of pairwise genetic differentiation F _ST varied in the range of 0.002–0.014, nor was it statistically significant in all comparison pairs between the herring samples.     

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.     

Mercury contamination and life history traits of Allis shad <Emphasis Type="Italic">Alosa alosa</Emphasis> (Linnaeus, 1758) and Twaite shad <Emphasis Type="Italic">Alosa fallax</Emphasis> (Lacépède, 1803) in the Gironde estuary (South West France)

Mercury concentration [Hg] was assessed in 20 adult Allis shad Alosa alosa (54–59 cm) and 20 adult Twaite shad Alosa fallax (36–44 cm) collected during their spawning migration in the Dordogne and the Garonne rivers (France). [Hg] was measured in the gills, dorsal muscle, liver and kidney. Twaite shad exhibited higher [Hg] than Allis shad. Median [Hg] values were [Hg]_Gills = 0.33 μg g⁻¹ (dw), [Hg]_Muscle = 1.22 μg g⁻¹, [Hg]_Liver = 1.99 μg g⁻¹, [Hg]_Kidney = 1.93 μg g⁻¹ for Twaite shad and [Hg]_Gills = 0.06 μg g⁻¹, [Hg]_Muscle = 0.20 μg g⁻¹, [Hg]_Liver = 1.18 μg g⁻¹, [Hg]_Kidney = 1.08 μg g⁻¹ for Allis shad. In order to understand such differences, we investigated some life history traits of the two species: migratory history, age (3–6 years), size at age (an expression of growth) and the number of spawning events (0–2 events). The difference in estuarine residence time between the juveniles of both species, which was assumed to influence [Hg], was investigated using otolith Sr:Ca ratio. The microchemical analysis revealed a significant difference in the residence time of juveniles in the estuary (medians are 21 d and 10 d for Twaite shad and Allis shad, respectively) but this residence time seems too short to influence [Hg] (Allis shad: r _spearman < 0.128; Twaite shad: r _spearman < −0.340). As both species show the same age structure, the influence of age on [Hg] was negligible. The literature shows that the differences in growth and in the number of spawning events reported in our study are in favour of a higher [Hg] for Twaite shad than for Allis shad. Although trophic status was not investigated here, the literature reveals that it is another factor that could produce higher [Hg] in Twaite shad, since its diet includes higher trophic levels than Allis shad. Guest editors: S. Dufour, E. Prévost, E. Rochard & P. Williot Fish and diadromy in Europe (ecology, management, conservation)     

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 Effects of Disease-Induced Juvenile Mortality on the Transient and Asymptotic Population Dynamics of Chinook Salmon (Oncorhynchus tshawytscha)

The effects of an increased disease mortality rate on the transient and asymptotic dynamics of Chinook salmon (Oncorhynchus tshawytscha) were investigated. Disease-induced mortality of juvenile salmon has become a serious concern in recent years. However, the overall effects of disease mortality on the asymptotic and transient dynamics of adult spawning abundance are still largely unknown. We explored various scenarios with regard to the density-dependent process, the distribution of survivorship over the juvenile phase, the disease mortality rate, and the infusion of stray hatchery fish. Our results suggest that the sensitivity to the disease mortality rate of the equilibrium adult spawning abundance and resilience (asymptotic return rate toward this equilibrium following a small perturbation) varied widely and differently depending on the scenario. The resilience and coefficient of variation of adult spawning abundance following a large perturbation were consistent with each other under the scenarios investigated. We conclude that the increase in disease mortality likely has an effect on fishery yield under a fluctuating environment, not only because the mean equilibrium adult spawning abundance has likely been reduced, but also because the resilience has likely decreased and the variance in adult spawning abundance has likely increased. We also infer the importance of incorporating finer-scale spatiotemporal information into population models and demonstrate a means for doing so within a matrix population modeling framework.     

Impact of larval Dorosoma predation on Daphnia parvula dynamics

• 1 In reservoirs of the south-eastern United States, Daphnia typically decline in abundance during the summer months. Current understanding suggests that adult shad (Dorosoma spp.), the dominant planktivore in these reservoirs, plays a limited role in Daphnia midsummer declines. The impact of shad larvae is beginning to be documented.
• 2 To examine the impact of larval shad on Daphnia seasonal dynamics, larval dynamics were studied over two seasons and stomach contents were analysed during one season.
• 3 Initial Daphnia panwla declines correlated with abundance of shad larvae. During the decline, shad stomach contents consisted of 97—100%D. parvula. After D. parvula declined, Bosmina and Diaphanosoma accounted for up to 97% of larval diets. When shad diets included other cladocerans, D. parvula abundance increased in the reservoir.
• 4 When this predation pattern was incorporated into a predictive model, larval shad predation accounted for an initial, temporary decline in D. parvula abundance. However, when other crustaceans were incorporated into the shad diet, D. parvula populations increased.
• 5 These results suggest that larval shad have a temporary, strong influence on the structure and abundance of D. parvula populations in south-eastern reservoirs of the United States.

     

Effects of gizzard shad on benthic communities in reservoirs

Effects of gizzard shad Dorosoma cepedianum on benthic communities in a large southern reservoir (Lake Texoma, U.S.A.) were examined during two field enclosure and exclosure experiments in which enclosures were stocked at high and low densities in 1998 and 1999, respectively. In both years, chironomid abundance significantly increased in treatments that excluded large fishes from foraging on sediments. Mean abundance of chironomids and ostracods were significantly higher ( P  < 0·05) in exclosures than enclosures stocked with gizzard shad at 1140–1210 kg ha⁻¹. In 1999, benthic invertebrate abundances did not differ ( P  > 0·08) between exclosure and enclosures stocked at 175–213 kg ha⁻¹. Per cent organic matter, algal abundance and abundance of other macroinvertebrates in sediments did not differ significantly among treatments in either year. Although chironomid abundance was reduced in gizzard shad enclosures in 1998, food habits from this and other studies showed that adult gizzard shad in Lake Texoma only consumed detritus and algae. It is likely that high sedimentation rates in Lake Texoma limit the ability of gizzard shad to regulate algae and detritus in benthic sediments. Thus, it is concluded that disturbance of benthic sediments by gizzard shad caused the observed reduction in chironomid abundance, rather than through consumption or competition for resources.     

Demographic Structure,Sex Ratio and Growth Rates of Southern Bluefin Tuna (Thunnus maccoyii) on the Spawning Ground

The demographics of the southern bluefin tuna (SBT) Thunnus maccoyii spawning stock were examined through a large-scale monitoring program of the Indonesian longline catch on the spawning ground between 1995 and 2012. The size and age structure of the spawning population has undergone significant changes since monitoring began. There has been a reduction in the relative abundance of larger/older SBT in the catch since the early 2000s, and a corresponding decrease in mean length and age, but there was no evidence of a significant truncation of the age distribution. Pulses of young SBT appear in the catches in the early- and mid-2000s and may be the first evidence of increased recruitment into the spawning stock since 1995. Fish in these two recruitment pulses were spawned around 1991 and 1997. Size-related variations in sex ratio were also observed with female bias for fish less than 170 cm FL and male bias for fish greater than 170 cm FL. This trend of increasing proportion of males with size above 170 cm FL is likely to be related to sexual dimorphism in growth rates as male length-at-age is greater than that for females after age 10 years. Mean length-at-age of fish aged 8–10 years was greater for both males and females on the spawning ground than off the spawning ground, suggesting that size may be the dominant factor determining timing of maturation in SBT. In addition to these direct results, the data and samples from this program have been central to the assessment and management of this internationally harvested stock.     

Piscivore enhancement effects on food webs depend on planktivore body size and species composition in replicated whole lake experiments

Recent syntheses of trophic cascade and biomanipulation research have suggested that the effects of piscivores on planktivorous fish populations are reduced, when planktivores are capable of outgrowing predator gape limitation and in systems with complex food web interactions. These hypotheses, however, have not been tested in long-term, whole-lake, experiments where processes such as fish recruitment and compensatory food web responses may be important. We conducted a replicated whole-lake experiment to test for the effects of supplemental piscivore introductions on food webs of eutrophic lakes dominated by deep-bodied planktivores. Responses to piscivore enhancement were compared between lakes differing in food web structure due to the presence of omnivorous gizzard shad (Dorosoma cepedianum). A significant decrease in the relative abundance of juvenile planktivorous fish, and an increase in total benthic macroinvertebrate density was observed in lakes containing mainly bluegills (Lepomus machrochirus). In contrast, lakes containing gizzard shad exhibited no significant responses to piscivore manipulation. Our results support the hypothesis that food webs in lakes dominated by deep-bodied planktivorous fish species respond weakly to piscivore enhancement. In addition, our findings support the hypothesis that cascading trophic interactions are weaker in lake ecosystems with more complex food web interactions such as those containing gizzard shad.     

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.     

An analysis of temporal variability in abundance, diversity and growth rates within the coastal ichthyoplankton assemblage of South Georgia (sub-Antarctic)

The study of spatial and temporal distribution and diversity of ichthyoplankton (fish eggs and larvae) can provide fisheries-independent information on the population dynamics and recruitment processes of marine fish species. Ichthyoplankton studies in the Southern Ocean have to date been largely constrained to the summer months. We analysed ichthyoplankton data collected from a year round, long term (2002–2008), plankton trawl sampling programme in a large fjord system (Cumberland Bay) at South Georgia, sub-Antarctic (54.25°S, 36.5°W) to assess temporal changes in larval fish diversity and abundance. Larvae of 22 species, representing nine families, were identified although three, Krefftichthys anderssoni (Myctophidae), Lepidonotothen nudifrons/Trematomus hansoni (Nototheniidae) and Champsocephalus gunnari (Channichthyidae), dominated abundance in all years. Significant seasonal and interannual differences in the larval fish assemblage were revealed by multivariate analyses. Estimates of larval growth are provided for five abundant species. Considerable inter-specific differences in relative larval growth rate were recorded but interannual variability within species was small. However, in the commercially important C. gunnari, multiple larval cohorts, representing a protracted spawning season, were observed to grow at different rates, and this may be related to temperature and/or food availability. A comparison with historical growth data from South Georgia suggests there has been little change in growth rate for the main species over the last three decades.     

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.     

Ecological investigations of the zooplankton community of Balsfjorden,northern Norway: Population structure and breeding biology of the chaetognath Sagitta elegans Verrill

The population dynamics of the chaetognath Sagitta elegans Verrill has been followed in Balsfjorden in 1976 and 1977. Seasonal variation in abundance, length-frequency distribution, growth in total length, and maturity stages are presented and discussed in relation to changes in hydrography.An annual generation of S. elegans was found, with a protracted and more or less continuous breeding season from May until October during 1977. The 1976 year-class consisted of two distinct length groups, both of which participated in the 1977 spawning. This spawning gave rise to possibly four sub-populations during 1977. The variation in numbers of sub-populations produced during the spawning season in 1976 and 1977 is discussed in relation to the hydrographical conditions in Balsfjorden. From November 1976 to March 1977 the abundance of S. elegans varied between 1 and 8 ind. · m^?3. The lowest value was recorded in May (0.9 ind. · m^?3). From September to December 1977 the population abundance was ≈2 ind. · m^?3.     

Physical forces experienced by echinoid eggs in the oviduct during spawning: comparison of the geminate pair Echinometra vanbrunti and Echinometra lucunter

During spawning, eggs of echinoids are released from the gonad and traverse a narrow oviduct-gonopore complex before being released into the water surrounding the spawning female. As the eggs traverse the oviduct-gonopore complex, a velocity gradient develops within the fluid formed by the eggs. This velocity gradient imposes a shear stress on the eggs that ranges from a maximal value at the walls of the oviduct-gonopore complex and declines linearly to zero at the center of the complex. In addition to shear stress, the eggs of some echinoids also experience strain. Strain is imposed on the eggs in situations where the diameter of the egg is greater than the diameter of the oviduct-gonopore complex. Both shear stress and strain have the potential to damage echinoid eggs during spawning and are likely selective pressures acting on the physical properties of protective extracellular layers, which surround the eggs of these organisms. The amounts of shear stress and strain experienced by eggs of a geminate pair of echinoids, Echinometra vanbrunti and E. lucunter, are calculated. Calculations of shear stress based on the mean spawning rates and dimensions of the oviduct-gonopore complexes of specimens collected during this study ranged up to 30 Pa for E. vanbrunti and up to 17 Pa for E. lucunter. To obtain conservative estimates of shear stress, these calculations were repeated incorporating a spawning rate that was reduced by one order of magnitude below the mean rate measured for both species. These calculations yielded shear stresses that ranged up to 13 and 6 Pa for E. vanbrunti and E. lucunter, respectively. The diameter of the oviduct-gonopore complex of echinoids increases linearly with the diameter of the tests of the adults. Because the diameter of the oviduct-gonopore complex influences the amount of shear stress experienced by eggs of echinoids, shear stress calculations were repeated incorporating the full range of potential diameters of the oviduct-gonopore complexes from data in the literature. Using the mean spawning rates obtained from each species, the amount of shear stress experienced by eggs ranged up to 32 Pa for E. vanbrunti and up to 398 Pa for E. lucunter. When the mean spawning rates were reduced by one order of magnitude, the amount of shear stress experienced by E. vanbrunti eggs ranged up to 14 Pa and up to 140 Pa for the eggs of E. lucunter. The eggs of E. vanbrunti do not experience strain during spawning because the diameter of the its eggs is smaller than the diameter of its oviduct-gonopore complex at all stages of adult growth. In contrast, the diameter of the eggs of E. lucunter is larger than the diameter of the oviduct-gonopore complex at small adult sizes and the eggs of this species experience true strains ranging up to 1.2. These results indicate that there are substantial differences in the amounts of physical forces that the eggs of echinoids experience during spawning both within species at different stages of adult growth and between species. Different amounts of shear stress and strain experienced by eggs among echinoid species may select for differences in the mechanical properties of protective extracellular layers that surround them.     

Potential changes to the biology and challenges to the management of invasive sea lamprey Petromyzon marinus in the Laurentian Great Lakes due to climate change

Control programs are implemented to mitigate the damage caused by invasive species worldwide. In the highly invaded Great Lakes, the climate is expected to become warmer with more extreme weather and variable precipitation, resulting in shorter iced‐over periods and variable tributary flows as well as changes to pH and river hydrology and hydrogeomorphology. We review how climate change influences physiology, behavior, and demography of a damaging invasive species, sea lamprey (Petromyzon marinus), in the Great Lakes, and the consequences for sea lamprey control efforts. Sea lamprey control relies on surveys to monitor abundance of larval sea lamprey in Great Lakes tributaries. The abundance of parasitic, juvenile sea lampreys in the lakes is calculated by surveying wounding rates on lake trout (Salvelinus namaycush), and trap surveys are used to enumerate adult spawning runs. Chemical control using lampricides (i.e., lamprey pesticides) to target larval sea lamprey and barriers to prevent adult lamprey from reaching spawning grounds are the most important tools used for sea lamprey population control. We describe how climate change could affect larval survival in rivers, growth and maturation in lakes, phenology and the spawning migration as adults return to rivers, and the overall abundance and distribution of sea lamprey in the Great Lakes. Our review suggests that Great Lakes sea lamprey may benefit from climate change with longer growing seasons, more rapid growth, and greater access to spawning habitat, but uncertainties remain about the future availability and suitability of larval habitats. Consideration of the biology of invasive species and adaptation of the timing, intensity, and frequency of control efforts is critical to the management of biological invasions in a changing world, such as sea lamprey in the Great Lakes.     

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.     

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.     

Population genetics of pike,genus <Emphasis Type="Italic">Esox</Emphasis> (Actinopterygii,Esocidae), in Northern Italy: evidence for mosaic distribution of native,exotic and introgressed populations

Biomanipulation, or management actions aimed to structure biological communities to achieve certain goals, has often been used in the restoration of aquatic ecosystems. In 2000, The Nature Conservancy acquired the Emiquon Preserve, which included two former Illinois River floodplain lakes, to restore these ecosystems. Restoration included stocking to establish a native fish community commensurate with historical records. Largemouth bass (Micropterus salmoides, bass) were also introduced to control poor water clarity and invasive common carp (Cyprinus carpio, carp). We summarized fish community characteristics and tested whether bass contributed to water clarity maintenance and limited carp during 2007–2014. The fish community was dominated by species stocked in greatest abundance, 13 of 32 species initially stocked have not been collected, and species diversity increased. No carp were observed in bass diets, water clarity declined significantly, and carp relative abundance increased. Increasing water levels during 2008–2009 diffused bass predation potential upon zooplanktivorous fishes and carp and weakened potential trophic cascading interactions. Our findings suggest that water level management, greater stocking of piscivores to maintain predator densities, prevention of gizzard shad (Dorosoma cepedianum) introduction, and/or a more diverse fish community including other native piscivores may be required to achieve long-term restoration goals.