Biogeographic freshwater fish pattern legacy revealed despite rapid socio‐economic changes in China

Understanding drivers of freshwater fish assemblages is critically important for biodiversity conservation strategies, especially in rapidly developing countries, which often have environmental protections lagging behind economic development. The influences of natural and human factors in structuring fish assemblages and their relative contributions are likely to change given the increasing magnitude of human activities. To discriminate natural and human drivers of fish diversity and assemblage patterns in developing countries with rapid socio‐economic development, a dataset of 908 freshwater fish species and 13 metrics including three categories of both natural (i.e., biogeographic) and human drivers (i.e., economic growth, inland fisheries) in China were analysed with machine learning algorithms (i.e., self‐organizing map, random forest). Here, we found that biogeographic drivers explained 21.8% of the observed fish assemblage patterns in China and remained stronger predictors when compared to human drivers (i.e., 15.6%, respectively). Freshwater fish species richness was positively correlated to rainfall, air temperature, surface water area and inland fisheries production but negatively correlated with urbanization. In addition, the strong structuring effects of climatic variables on Chinese fish richness patterns suggested that the fish assemblages could be particularly vulnerable to climate change. Our results showed that natural biogeographic factors still dominate in driving freshwater fish assemblage patterns despite increased human disturbances on aquatic ecosystems in a rapidly developing country. These findings consequently suggested that we should consider both natural (e.g., climate) and human (e.g., urbanization, inland fisheries) factors when establishing aquatic conservation strategies and priorities for developing countries that are experiencing rapid socio‐economic changes.     

Analysis of the trophy sport fishery for the speckled peacock bass in the Rio Negro River, Brazil

Abstract  The middle portion of the Rio Negro River in Brazil near the equator supports a popular recreational sport fishery for speckled peacock bass, Cichla temensis (Humboldt). The objective of this study was to determine the effect of fishing mortality on this population. Fish were collected from sport-fishing ( n  = 72) and commercial ( n  = 103) catches and otoliths were aged to estimate longevity, growth and natural mortality. Recreational anglers in this region seek to catch, then release, larger speckled peacock bass; and fish larger than 62 cm standard length (SL) (about 4.5 kg) served as a bench mark to assess the potential impact of subsistence and commercial harvest on the abundance of larger fish in the sport fishery. Time of opaque band formation on otoliths generally coincided with the dry season (November to April); these bands appeared to form once per year, but formation was highly variable. Speckled peacock bass grew to 62 cm SL on average in 6.4 years, but some fish obtained this size in 4–5 years. Maximum age was 9 years, but most fish were less than 7 years. Instantaneous annual natural mortality ( M ) estimated from maximum size, longevity and growth ranged from 0.19 to 0.44. Simulation modelling predicted that exploitation rates of fish >25 cm SL similar to the estimated natural mortality rates would reduce the abundance of fish >62 cm by 67–89% compared with no harvest. Even modest exploitation rates of 5% and 10% would result in approximately 30–50% reduction, respectively, of these larger fish. Abundance of large speckled peacock bass that sustains the sport fishery is susceptible to low rates of exploitation in this remote region of Brazil.     

Short-term mortality of Australian bass, Macquaria novemaculeata, after catch-and-release angling

Abstract  Australian bass, Macquaria novemaculeata Steindachner, is angled in large numbers from coastal impoundments and rivers throughout south-eastern Australia. Many are released in the belief that most survive with few negative short-term impacts. This assumption was investigated in three experiments involving a range of conventional angling scenarios. A total of 313 Australian bass were angled and then released along with appropriate controls into either land-based tanks or floating cages, where they were monitored for up to 5 days. No controls died. Short-term mortalities of angled fish were low (0–6%) and attributed to the effects of bait type, hook location and fish size ( P  <   0.05). Specifically, fish that ingested hooks or were caught with natural baits were more likely to die than those that were mouth-hooked or caught on lures. Mortality negatively correlated with fish size. The results supported previous assertions that Australian bass is a resilient, hardy species that should tolerate short-term impacts associated with catch-and-release angling. However, other sublethal physiological impacts that could contribute towards delayed mortalities or reduced reproductive output warrant further investigation, particularly in relation to native populations in coastal rivers that form gauntlet fisheries during their annual spawning season.     

Spring home ranges of white bass in irrigation reservoirs of the Republican River Basin, Nebraska

Abstract –  Fishery biologists have documented small home ranges, relative to available habitat, for many littoral freshwater fishes. Home ranges for pelagic species, such as white bass Morone chrysops , are generally not well described, yet are thought to be large. We studied white bass movement using acoustic telemetry in two irrigation reservoirs of the Republican River basin in south-western Nebraska. Acoustic transmitters were implanted in fall of the previous year and tracking occurred a minimum of once per week throughout spring (mid-March to May) 2007 and 2008. Linear home ranges were calculated from observed locations of individual fish. Twelve of the twenty-seven tagged fish with at least five locations exhibited localised home ranges throughout the spring whereas the remaining fish exhibited home ranges extending across large portions of each reservoir. Home range size was not correlated with fish size or condition.     

Relationship between freshwater input to the coastal zone and the historical landings of the benthic/demersal fish Eleginops maclovinus in central-south Chile

The ecosystem off central-south Chile is one of the most productive marine systems in the world, providing approximately 4% of worldwide fish captures. We analysed the effect of freshwater input (river runoff and rainfall) to the coastal zone in central-south Chile (36°00'–37°30' lat. S) on the landings of the Róbalo ( Eleginops maclovinus ) using the following data: Róbalo annual landing statistics, monthly mean and annual mean runoff of Itata (1975–91) and Bio-Bio rivers (1975–94), and cumulative monthly rainfall and annual mean rainfall (1975–94). Time series were smoothed using three-point equally weighted running means. To analyse the relationship between Róbalo's landing and river runoff, as well as rainfall, Pearson's product-moment correlation analyses were conducted at relevant lag times. The statistical significance of correlation coefficients was calculated taking into account intraseries autocorrelation. Our results showed that Róbalo's landings were significantly correlated to freshwater input to the coastal zone in central-south Chile. Thus, significant correlation coefficients were found between annual Róbalo's landings and: (i) annual mean runoff of the Itata ( r =−0.94) and Bio-Bio ( r =−0.80) rivers lagged 3 and 4 years, respectively, and (ii) annual mean rainfall lagged 4 years ( r =−0.75). The lag time at which highest correlation coefficients between annual mean environmental and annual landing time series took place (i.e. 3–4 years) corresponded to the age at which Róbalo reaches the minimum exploitation size. Accordingly, we hypothesize that river runoff and rainfall strongly influence the survival rate of Róbalo's early stages during the first year of life.     

The response of fish larvae to decadal changes in environmental forcing factors off the Oregon coast

We conducted a statistical analysis to characterize the influence of large‐scale and local environmental factors on presence‐absence, concentration, and assemblage structure of larval fish within the northern California Current (NCC) ecosystem, based on samples collected at two nearshore stations along the Newport Hydrographic line off the central Oregon coast. Data from 1996 to 2005 were compared with historical data from the 1970s and 1980s to evaluate pseudo‐decadal, annual, and seasonal variability. Our results indicate that the most abundant taxa from 1996 to 2005 differ from those of earlier decades. Concentrations of the dominant taxa and total larvae were generally greater in the winter/spring than summer/fall season. Using generalized additive modeling, variations in presence‐absence and concentration of taxa were compared to climate indices such as the Pacific Decadal Oscillation, Northern Oscillation Index, and the multivariate ENSO index and local environmental factors, such as upwelling, Ekman transport, and wind stress curl. Significant relationships were found for various combinations of environmental variables with lag periods ranging from 0 to 7 months. We found that the large‐scale climate indices explained more of the variance in larval fish concentration and diversity than did the more local factors. Our results indicate that readily available oceanographic and climate indices can explain variations in the dominant ichthyoplankton taxa in the NCC. However, variation in response among taxa to the environmental metrics suggests additional unknown factors not included in the analysis likely contributed to the observed distribution patterns and larval fish community structure in the NCC.     

Temperature influence on the fish assemblage structure in a large lowland river, the lower Oder River, Germany

Abstract –  Water temperatures were expected to structure river fish assemblages by promoting: (i) offspring's growth and densities in warm years, (ii) spring spawning fish when temperatures increase early and (iii) improved growth and overwintering condition from elongated vegetation periods. Favourable temperature conditions in their first year should improve growth and survival in consecutive years. In the lower Oder River, Germany, fish have been sampled annually, 0+ in July since 1997, and older fish in November since 1998. This data set was analysed to test the hypotheses mentioned above. The water temperature ranged between −0.4 and 25.1 °C during the study period, resulting in substantial variations of the 29 temperature metrics derived from daily means. Cumulative degree-days until catch, in spring and during March, as well as minimum and mean temperatures in March and spring explained 63% of the variability observed. The mean total length of the most frequent 0+ fish differed significantly between years related to temperature. Increased length growth at higher temperatures was a general tendency. Total length and density of 0+ fish were significantly inversely correlated, but the potential influence of temperature on species densities was not significant. The mean fish densities varied not significantly between years. No indication was found that higher total length in the first year of life resulted in an improved growth in subsequent years.
For future research and management it has to be expected that temperature impacts become increasingly significant in regulated river systems. With decreasing hydrodynamics the significance of temperature effects will increase.     

Diet overlap between native bigmouth sleepers (Gobiomorus dormitor) and introduced predatory fishes in a Puerto Rico reservoir

Abstract –  Native fish species coexist with introduced species in Puerto Rico's freshwater systems, yet competition between these species has not been evaluated. We examined the extent of diet overlap between native bigmouth sleepers Gobiomorus dormitor and introduced largemouth bass Micropterus salmoides and peacock bass Cichla ocellaris in a Puerto Rico reservoir. Bigmouth sleepers and largemouth bass exhibited an ontogenetic shift in feeding habits, whereas peacock bass were exclusively piscivorous at all sizes collected in this study. Biologically significant diet overlap was observed between large bigmouth sleepers and largemouth bass, but not between large bigmouth sleepers and peacock bass, or between large largemouth bass and adult peacock bass. No significant diet overlap in any species combination was observed in small or medium size classes. Better understanding of the ecology of these coexisting predators should lead to improved conservation of bigmouth sleepers, and improved fisheries management for all three predatory species.     

Replacement of fishmeal with rice protein concentrate in practical diets for European sea bass Dicentrarchus labrax reared at winter temperatures

As no information is available regarding the efficacy of using rice protein concentrate (RPC) to replace fishmeal (FM) in diets for European sea bass (Dicentrarchus labrax), a trial was conducted to determine the optimum inclusion for European sea bass juveniles. Diets were formulated to replace 25%, 50% and 75% of fishmeal with RPC. Additional high level inclusion diets (50% and 75%) were supplemented with lysine and methionine to determine if these were limiting factors. Fish (18.0 ± 0.06 g) were fed the experimental diets for 12 weeks. The final mean weight and specific growth rate (SGR) were significantly higher in fish fed the control diet (diet FM) and low level RPC (diet RPC25) than in fish fed the other diets. The supplementation of the amino acids had a positive effect on growth, elevating the final weight of the high level RPC (RPC75 + AA) group above that of the non‐supplemented group (RPC75). However, this was still not comparable to fish fed FM and RPC25 diets. No significant differences in body composition were observed. A digestibility study demonstrated an inverse relationship between elevating inclusion levels of RPC and apparent digestibility coefficients (ADCs) for dry matter and lipid, with significantly lower values than the control being observed with RPC inclusions above 25%. The mesenteric fat index followed this trend. Circulating leucocyte levels, leucocyte ratios and serum lysozyme activity remained unaffected by dietary treatment. However, compared with the control group, fish fed RPC75, RPC50 + AA RPC75 + AA displayed significantly lower haematocrit values. The present study demonstrates that the inclusion of RPC at 140 g kg^?1 (effectively replacing 25% of FM content) does not compromise European sea bass growth performance, body quality or basic haematological parameters. It is suggested that future studies including higher levels test the efficacy of additional amino acid supplementation (e.g. tryptophan).     

Influence of hatch duration and individual daily growth rates on size structure of age-0 smallmouth bass cohorts in two glacial lakes

Abstract –  We assessed hatch dates and daily growth rates of age-0 smallmouth bass from two glacial lakes over a 3-year period. Hatching durations (19–27 days among years) of smallmouth bass occurred from late May until late June. Mean daily growth rate of age-0 smallmouth bass ranged from 0.56 to 1.56 mm·day⁻¹. Correlation analysis indicated that hatch date had little effect on daily growth rates. Hatch date was significant in explaining variation in total length (TL) of age-0 smallmouth bass at time of capture in only three of six cases and hatch date never explained >50% of the variation in bass length. Daily growth rate significantly explained variation in smallmouth bass TL at time of capture in all six models, accounting for 31–86% of the variability in bass length. Our findings suggest that size structure of age-0 cohorts in some populations may be more strongly regulated by variation in individual daily growth rate than by hatch timing.     

Growth rate and mortality of Australian bass, Macquaria novemaculeata, in four freshwater impoundments in south-eastern Queensland, Australia

Abstract  Growth and total mortality of Australian bass, Macquaria novemaculeata Steindachner, were studied in four freshwater impoundments using data from a cooperative angler tagging programme. Growth was modelled using the computer program GROTAG. Annual growth increments varied among impoundments, ranging from 5.0 to 7.8 cm yr⁻¹ among fish 20-cm total length (TL) and 1.7 to 4.9 cm yr⁻¹ among fish 30 cm TL, but exceeded previous estimates for the species. Annual total mortality rates, based on the decrease in recapture rate with time, ranged from 0.03 to 0.07 and were lower than for other comparable species, presumably because of the prevalence of catch and release fishing in the impoundments studied.     

A study on relationships between large‐scale climate indices and estimates of North Pacific albacore tuna productivity

A logistic production model was used to examine potential relationships between three climate indices, the North Pacific Gyre Oscillation (NPGO), the Pacific Decadal Oscillation (PDO), and the Multivariate El Niño‐Southern Oscillation Index (MEI), and productivity estimates of the North Pacific albacore tuna (Thunnus alalunga) population. Catch and standardized catch‐per‐unit‐effort data from three longline fisheries (Japan, US, and Taiwan) were used in the model. The climate indices were incorporated into the model by correlating time‐varying intrinsic population growth rate (r_y) of the production model with the annual mean value for each index. The estimated probability that the NPGO is positively correlated with stock productivity, as measured by r_y, was 0.99, and the calculated probability that MEI is negatively correlated with the productivity was 0.95. The time lag for these correlations is 4 yr, which is consistent with the timing of recruitment to the Japan longline fishery. The PDO did not seem to have any detectable relationship with stock productivity. However, it remains uncertain if there is a conclusive linkage between the albacore productivity and the NPGO or the MEI index, because model fit to the data is about the same as that of a base model which does not use any climate index and assumes a time‐invariant r.     

Climate‐induced seasonal changes in smallmouth bass growth rate potential at the southern range extent

Temperature increases due to climate change over the coming century will likely affect smallmouth bass (Micropterus dolomieu) growth in lotic systems at the southern extent of their native range. However, the thermal response of a stream to warming climate conditions could be affected by the flow regime of each stream, mitigating the effects on smallmouth bass populations. We developed bioenergetics models to compare change in smallmouth bass growth rate potential (GRP) from present to future projected monthly stream temperatures across two flow regimes: runoff and groundwater‐dominated. Seasonal differences in GRP between stream types were then compared. The models were developed for fourteen streams within the Ozark–Ouachita Interior Highlands in Arkansas, Oklahoma and Missouri, USA, which contain smallmouth bass. In our simulations, smallmouth bass mean GRP during summer months decreased by 0.005 g g^?1 day^?1 in runoff streams and 0.002 g g^?1 day^?1 in groundwater streams by the end of century. Mean GRP during winter, fall and early spring increased under future climate conditions within both stream types (e.g., 0.00019 g g^?1 day^?1 in runoff and 0.0014 g g^?1 day^?1 in groundwater streams in spring months). We found significant differences in change in GRP between runoff and groundwater streams in three seasons in end‐of‐century simulations (spring, summer and fall). Potential differences in stream temperature across flow regimes could be an important habitat component to consider when investigating effects of climate change as fishes from various flow regimes that are relatively close geographically could be affected differently by warming climate conditions.     

Compensatory growth response of European sea bass (Dicentrarchus labrax L.) under cycled starvation and restricted feeding rate

The compensatory growth response of the European sea bass (Dicentrarchus labrax) that faced cycling starvation and restricted ration was assessed. Juveniles (10.5 g) were stocked into 15 tanks at a density of 25 fish per tank. Five different feeding regimes were tested on triplicate groups of fish: C_Satiation: control fed for 60 days without deprivation, C_Restricted: 25% restricted feeding, S‐R: 1 day starvation then 4 days C_Restricted feeding, R‐F: 1 day C_Restricted feeding then 4 days C_Satiation feeding, and, finally, S‐F: 1 day starvation then 4 days C_Satiation feeding. The specific growth rate of fish in the C_Satiation (2.5 ± 0.06% day^?1), S‐F (2.5 ± 0.11% day^?1) and R‐F (2.4 ± 0.18% day^?1) were significantly higher than that of C_Restricted (2.2 ± 0.05% day^?1) or S‐R (2.0 ± 0.01% day^?1). Fish in S‐F group were able to achieve catching up with the C_Satiation. There was no significant difference in feed conversion rates but R‐F and S‐F consumed approximately 34% more feed than C_Satiation following the first re‐feeding day. Although, the highest lipid content was observed in C_Satiation (14.4%), S‐R (33.3% dry matter) had the highest water content compared with the C_Satiation and C_Restricted (37.8% and 36.9% dry matter respectively). In conclusion, it may be concluded that sea bass has rapid response to cycling starvation/re‐feeding and that a 25% restricted feeding ratio is insufficient to invoke a compensatory growth response in sea bass.     

摄食不同饵料的大口黑鲈肠道菌群分析

为了探究投喂冰鲜杂鱼和人工配合饲料对大口黑鲈肠道菌群的影响,本研究采用变性梯度凝胶电泳技术(PCR-DGGE)对摄食不同饵料的2组鲈鱼肠道内容物的细菌群落结构进行了比较分析.结果显示,冰鲜组与饲料组的大口黑鲈肠道菌群组成相似性较低(相似性24％),且冰鲜组肠道菌群多样性高于饲料组.对2组大口黑鲈中共有条带和特异性条带进一步测序分析显示,2组大口黑鲈肠道共有菌有红螺菌、梭菌、保科爱德华菌、嗜麦芽窄食单胞菌和蓝细菌.冰鲜组大口黑鲈肠道中的特异性菌归属为拟杆菌、肉食杆菌、魏斯氏菌和邻单胞菌;饲料组大口黑鲈肠道中的特异性菌归属为黄杆菌、鲸杆菌、柠檬明串珠菌、缺陷短波单胞菌.研究表明,与投喂冰鲜杂鱼的大口黑鲈相比,人工配合饲料降低了大口黑鲈肠道菌群多样性,也抑制了拟杆菌等有益菌在大口黑鲈肠道内的分布.推测饲料组大口黑鲈生长较慢可能与其肠道菌群中拟杆菌与厚壁菌的比值下降有关.研究结果提示,在今后大口黑鲈养殖或者人工配合饲料研制过程中,添加拟杆菌等益生菌也许可促进大口黑鲈的生长和营养吸收.     

Pelagic larval traits of the amphidromous goby Sicyopterus lagocephalus display seasonal variations related to temperature in La Réunion Island

Amphidromous gobies represent a substantial part of freshwater fish diversity throughout islands of the Indo‐Pacific region. They display a marine pelagic phase during several months before recruiting in rivers. Understanding the relationship between larval traits and environmental conditions is a major challenge for the evaluation of a spatial scale of connectivity and populations’ dynamics, especially in a climate change context. In this study, the larval traits of Sicyopterus lagocephalus were examined and related to the sea surface temperature (SST), over three consecutive years in La Réunion Island (Mascarene archipelago). The pelagic larval duration (PLD, range from 96 to 293 days), the size‐at‐recruitment (range from 26.5 to 37 mm TL) and the larval growth rate (range from 0.112 to 0.293 mm·day^?1 TL) varied seasonally depending on hatching date. The larval growth rate was inversely correlated to the fluctuations of PLD and size. Larvae living in high sea water temperatures exhibited a faster growth, shorter PLDs and smaller sizes‐at‐recruitment than those living in cool water temperatures. The instantaneous daily growth was assessed by the study of otolith increment widths. The daily growth was not linear throughout larval life and was positively correlated to SST. We showed high amplitude of PLD periodic fluctuations (170.39 ± 43.75 days) related to temperature (12.8% PLD drop per degree), which could affect dispersal and induce intermittent connectivity between distant populations. This high plasticity of larval traits is likely to be advantageous to respond to the wide range of environmental conditions encountered throughout the species distribution range.     

Prey fish selection by Far Eastern catfish <Emphasis Type="Italic">Silurus asotus</Emphasis> and largemouth bass <Emphasis Type="Italic">Micropterus salmoides</Emphasis>

ABSTRACT:   Prey fish selection by Far Eastern catfish and largemouth bass were examined using bluegill, Japanese dace and crucian carp as food fish. In both aquaria (1.2 m × 0.45 m) and ponds (2.8 m × 1.2 m), bluegill was not preyed on by catfish and bass more than dace and crucian carp. In aquaria, there was no significant difference in the consumption of dace and crucian carp between catfish and bass, but in ponds, catfish preyed on dace and crucian carp more and less than bass, respectively. In the case that only bluegill and catfish were introduced in ponds, catfish consumed 4–15 g of bluegill per day. The introduction of catfish into ponds and lakes for the purpose of eradicating bluegill is considered appropriate for areas with few native fish species.     

Exploitation and length limit evaluation of Alabama bass in a South‐eastern USA reservoir

Alabama bass, Micropterus henshalli Hubbs & Bailey, are the dominant sportfish of Allatoona Reservoir, Georgia, USA, but no population assessment has been conducted. Thus, growth and total annual mortality were estimated in spring 2005, and a tagging study was conducted in 2006 and 2007 to estimate angler exploitation. These data were used with an age‐structured model to assess performance of a 356‐mm minimum length limit (MLL), a 406‐mm MLL and a 330‐ to 406‐mm protected slot length limit (SLL) compared to the present harvest regulation of no MLL. Mean annual exploitation varied from 12 to 22% each year and was generally highest for fish > 330 mm; total annual mortality was 44%. Models predicted a 49–153% increase in numbers of Alabama bass reaching 432 mm, a 22–66% decline in numbers harvested and only moderate declines in yield (5–25%) with the alternative harvest regulations compared to current conditions. The SLL may be an acceptable compromise to allow Allatoona Reservoir anglers to still harvest fish while also improving Alabama bass size structure.     

Covariation in climate, zooplankton biomass and mackerel recruitment in the southern Gulf of St Lawrence

Between 1982 and 1991, an annual survey of stage I egg production of Atlantic mackerel ( Scomber scombrus ) was conducted in June/early July in the southern Gulf of St Lawrence. We investigated the relationship between interannual variability in biomass of zooplankton, determined from the archived survey plankton samples, and mackerel recruitment, estimated from the proportion of three-year-olds in the catch of the commercial fishery. Zooplankton biomass varied by a factor of 2.5, primarily owing to fluctuations in the >1000 μm size fraction. The index of mackerel recruitment fluctuated by a factor of ≈20 and was positively related (linear regression: P  < 0.05; n  = 10) to variations in the zooplankton biomass. Both mackerel recruitment and zooplankton biomass were negatively related (linear regression: P  < 0.05) to RIVSUM, a measure of freshwater discharge from the St Lawrence River system and an index of variability in the region's climate. Three hypotheses are put forward to explain these observations: (1) there is a strong link between interannual variation in abundance of copepod females, which produce prey for mackerel larvae, and larval survival; the exceptional recruitment and subsequent year class resulted from an exceptional production of Calanus finmarchicus nauplii; (2) years of high zooplankton biomass provide better feeding conditions and consequently higher survival of mackerel juveniles; and (3) mackerel recruitment and zooplankton biomass are independently under the control of an underlying physical process, without strong trophic linkage. The first hypothesis is supported by a study of copepod species composition and female abundance conducted for four of the survey years. At the present time, none of these hypotheses can be ruled out.     

Relationships between marine growth and marine survival of one sea winter Atlantic salmon, Salmo salar L., from the River Bush, Northern Ireland

An examination of marine growth/marine survival relationships in Atlantic salmon, Salmo salar L., was carried out, based on scale growth measurements in relation to two indices of marine survival in wild fish from the River Bush, Northern Ireland. The survival of cohorts to the Irish/Northern Irish coast (prefishery) and to fresh water was statistically unrelated to variation in growth from smolt migration to the end of the first winter at sea ( P  > 0.1; – P  > 0.7). Marine growth of 1+ smolts decreased significantly during the period of the study ( P  < 0.05), but growth of 2+ smolts did not change ( P  > 0.05). The variability in marine growth was much less than the variation in natural survival at sea, suggesting that factors instead of or in addition to, growth influence natural survival in the marine environment. Survival to fresh water was not related to survival to the coast ( P  > 0.4), although it was inversely correlated with exploitation rate ( P  < 0.01). These results are discussed in relation to the use of freshwater returns to assess marine survival and the potential for the variation in natural marine mortality to influence total life-cycle variation.