Density‐dependent and density‐independent growth in a population of juvenile sea‐trout,Salmo trutta,assessed using long‐term data from a small stream in Northwest England

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Size-independent growth in fishes: patterns, models and metrics

A combination of a dynamic energy budget (DEB) model, field data on Atlantic salmon Salmo salar and brown trout Salmo trutta and laboratory data on Atlantic salmon was used to assess the underlying assumptions of three different metrics of growth including specific growth rate (G), standardized mass‐specific growth rate (G_S) and absolute growth rate in length (G_L) in salmonids. Close agreement was found between predictions of the DEB model and the assumptions of linear growth in length and parabolic growth in mass. Field data comparing spring growth rates of age 1+ year and 2+ year Atlantic salmon demonstrated that in all years the larger age 2+ year fish exhibited a significantly lower G, but differences in growth in terms of G_S and G_L depended on the year examined. For brown trout, larger age 2+ year fish also consistently exhibited slower growth rates in terms of G but grew at similar rates as age 1+ year fish in terms of G_S and G_L. Laboratory results revealed that during the age 0+ year (autumn) the divergence in growth between future Atlantic salmon smolts and non‐smolts was similar in terms of all three metrics with smolts displaying higher growth than non‐smolts, however, both G_S and G_L indicated that smolts maintain relatively fast growth into the late autumn where G suggested that both smolts and non‐smolts exhibit a sharp decrease in growth from October to November. During the spring, patterns of growth in length were significantly decoupled from patterns in growth in mass. Smolts maintained relatively fast growth though April in length but not in mass. These results suggest G_S can be a useful alternative to G as a size‐independent measure of growth rate in immature salmonids. In addition, during certain growth stanzas, G_S may be highly correlated with G_L. The decoupling of growth in mass from growth in length over ontogeny, however, may necessitate a combination of metrics to adequately describe variation in growth depending on ontogenetic stage particularly if life histories differ.     

The effect of stocking with 0+ year age‐class Atlantic salmon Salmo salar fry: a case study from the River Bush,Northern Ireland

An enhancement programme based on stocking 0+ year age‐class Atlantic salmon Salmo salar, conducted in the River Bush, Northern Ireland, U.K. over the period 1996–2005, was reviewed with reference to the performance and biological characteristics of wild fish. Wild ova to 0+ year fry (summer) survival was c. 8% with subsequent wild 0+ year fry‐to‐smolt survival c. 9%. Stocked unfed 0+ year juveniles gave c. 1% survival to smolt whilst fed 0+ year S. salar stocked in late summer exhibited survival at c. 5%. Stocking with unfed and fed fry contributed to increased smolt production and helped attain local management objectives between 2001 and 2005. Significant differences in biological characteristics were observed between wild and stocked‐origin fish. Wild‐smolt cohorts were dominated by 2+ year age‐class fish on the River Bush whilst smolts originating from fed fry mostly comprised younger 1+ year individuals. The mean mass of 1+ year smolts derived from stocked fed fry was significantly lower than that of wild 1+ year smolts, although these differences were not evident between older age classes. Differences in run timing between wild smolts and smolts derived from stocked fry were also apparent with the stocked‐origin fish tending to run earlier than wild fish. Although the stocking exercise was useful in terms of maximizing freshwater production, concerns over the quality of stocked‐origin recruits and the long term consequences for productivity are highlighted.     

The relationship between smolt density and fry density in salmonids

If the Ricker stock–recruitment model describes the relationship between egg density and survivor density at different stages of the life cycle, then the relationship between smolt density and fry density is not simple. Using data from a long-term study (1966–1990) of a sea trout population Salmo trutta , the relationship between density of potential smolts and fry densities in late M ay/early June or late August/early September is shown to be a reflexed curve with zero origin, so that there are two smolt densities for each fry density. A linear relationship is obtained only when the exponential parameter in the Ricker model is constant for the different life Stages.     

Compensatory growth offsets poor condition in native trout populations

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Temperature requirements of Atlantic salmon Salmo salar, brown trout Salmo trutta and Arctic charr Salvelinus alpinus: predicting the effects of climate change

Atlantic salmon Salmo salar, brown trout Salmo trutta (including the anadromous form, sea trout) and Arctic charr Salvelinus alpinus (including anadromous fish) provide important commercial and sports fisheries in Western Europe. As water temperature increases as a result of climate change, quantitative information on the thermal requirements of these three species is essential so that potential problems can be anticipated by those responsible for the conservation and sustainable management of the fisheries and the maintenance of biodiversity in freshwater ecosystems. Part I compares the temperature limits for survival, feeding and growth. Salmo salar has the highest temperature tolerance, followed by S. trutta and finally S. alpinus. For all three species, the temperature tolerance for alevins is slightly lower than that for parr and smolts, and the eggs have the lowest tolerance; this being the most vulnerable life stage to any temperature increase, especially for eggs of S. alpinus in shallow water. There was little evidence to support local thermal adaptation, except in very cold rivers (mean annual temperature <6·5° C). Part II illustrates the importance of developing predictive models, using data from a long-term study (1967-2000) of a juvenile anadromous S. trutta population. Individual-based models predicted the emergence period for the fry. Mean values over 34 years revealed a large variation in the timing of emergence with c. 2 months between extreme values. The emergence time correlated significantly with the North Atlantic Oscillation Index, indicating that interannual variations in emergence were linked to more general changes in climate. Mean stream temperatures increased significantly in winter and spring at a rate of 0·37° C per decade, but not in summer and autumn, and led to an increase in the mean mass of pre-smolts. A growth model for S. trutta was validated by growth data from the long-term study and predicted growth under possible future conditions. Small increases (<2·5° C) in winter and spring would be beneficial for growth with 1 year-old smolts being more common. Water temperatures would have to increase by c. 4° C in winter and spring, and 3° C in summer and autumn before they had a marked negative effect on trout growth.     

Temperature‐Dependent Growth and Life Cycle of Nemoura sichuanensis (Plecoptera: Nemouridae) in a Chinese Mountain Stream

Plecoptera constitute a numerically and ecologically significant component in mountain streams all over the world, but little is known of their life cycles in Asia. The life cycle of Nemoura sichuanensis and its relationship to water temperature was investigated during a 4‐year study in a headwater stream (known as the Jiuchong torrent) of the Xiangxi River in Central China. Size structure histograms suggest that the life cycle was univoltine, and the relationships between the growth of Nemoura sichuanensis, physiological time, and effective accumulated water temperature were described using logistic regressions. The growth pattern was generally similar within year classes but growth rates did vary between year‐classes. Our field data suggest a critical thermal threshold for emergence in Nemoura sichuanensis, that was close to 9 °C. The total number of physiological days required for completing larval development was 250 days. The effective accumulated water temperature was 2500 degree‐days in the field. Development during the life cycle increased somewhat linearly with the physiological time and the effective accumulated water temperature, but some non‐linear relationships were best developed by logistic equations. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Growth, survival and production of juvenile salmon and trout in a Scottish stream, 1966–75

Investigations of the growth, survival and production of young salmon Salmo salar , brown trout and sea trout S. trutta in sections of a stream in Scotland were made during 1966–75. At the end of the growing season, in autumn, the size of the 0+ salmon ranged from a mean weight of 1.12 g in 1966 to 2.82 g in 1973, and the size of the 0+ trout ranged from a mean of 2.20 g in 1966 to 3.56 g (68.0 mm) in 1974. Growth rates of 0+ salmon between July to September were similar from year to year, as was the case with the 0+ trout. The greater size attained in their first year by trout, resulted from the longer feeding season, provided by earlier emergence of fry and ability to continue growing in colder weather in autumn. The lengths attained by 0+ salmon and 0+ trout in September were related to the population densities of 0+ salmon and the number of days above 0° C from 1 December. There was no discernible relationship between lengths of 0+ trout and the population densities of 0+ trout. Salmon and trout lost weight during the winter, which was made up by April. The densities of 0+ salmon in June varied between 2–12m ^–2. Rates of decrease of the population densities in their first year were related to their densities at the beginning of the season, and, more closely, to the densities of salmon and trout combined. At the end of the second year's growth there were between 0.06 and 0.25 salmon m ^–2. Size of the trout populations varied less from year to year than those of salmon. The life of a year class of salmon and trout could be divided into several stages characterized by different rates of decrease of the population.     

The growth of juvenile Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L., in a Scottish river system subject to cooling-water discharge

The River Fiddich, a tributary of the R. Spey in north-east Scotland, is a spawning river for both Atlantic salmon and brown trout. Warm cooling water effluent is discharged from several distilleries at different points in the lower reaches and raises the temperature of the river 1–3°C above ambient for most of the year. Salmon and trout grow more rapidly in this region than further upstream, and juvenile salmon generally migrate a year earlier, as 2 + smolts. Available data were too few to determine whether there was a similar difference for trout. Similar studies on the R. Dullan, a tributary of the Fiddich, and on the Cromdale Burn in the same area, confirmed that the growth rate of fish is faster downstream from distillery discharge points. It is suggested that increased invertebrate production may influence the growth rate.     

Production of salmon and trout in a stream in Scotland

For three years the population size, rates of growth, standing stock, production and yield of all year classes of salmon and trout within three sections of a stream in Scotland were studied. Total salmon production as fresh weight per m² was 6.5 g in 1966, 10.6 g in 1967 and 11.1 g in 1968, and total trout production was 10.3g in 1966, 12.3 g in 1967 and 7.7g in 1968. Fish of 0+ and 1+ year old provided usually more than 90% of the total annual salmon production and 80 % of the annual trout production. Yield of salmon smolts (about 9 cm or longer after 2 years growth) per m² was 0.10 in 1966, 0.22 in 1967 and 0.15 in 1968. The smolt yield by weight was 29 % of the production of the 1966 year class of salmon and 16% of the 1967 year class. Numbers of trout of 9 cm or longer produced in the same time were higher and their weight was 60% of the total production of the 1966 year class and 32% of the 1967 year class.     

The distribution of salmonids in upland streams in relation to depth and gradient

The distribution of various age classes of salmon and trout was assessed in upland streams by electrofishing. Water depths and site gradients were measured and correlated to fish densities. The fry of both species were significantly more abundant in shallow water; up to 75·3% of salmon fry and 72·2% of trout fry were captured in sites of mean depth < 20 cm. Older trout were found mainly in the deeper areas, with a maximum of 7·4% captured in sites < 20 cm mean depth. Yearling fish were found in all the depth-ranges sampled, but with a tendency for higher numbers in mid-range depths. There were similar correlations in the abundance of each age class with the actual areas of shallow, mid-range and deep water habitat available within sites. Correlations of fish density with gradient indicated that trout were limited in their distribution to areas of lower flow, whereas salmon were not. Since depth and gradient were significantly negatively correlated, there was an apparent preference of trout for slightly deeper habitats than the equivalent year classes of salmon. The observed habitat segregation is discussed in terms of competition and selection.     

Change in allometric length–weight relationship of Salmo trutta at emergence from the redd

In their life cycle, especially at early life stages, fishes pass through distinct growth stanzas with different length‐weight relationship patterns. Salmonid fish fry emergence from the redd gravel is a crucial moment in their life history. This study presents an ontogenic change in allometric length–weight relationship scaling of sea trout Salmo trutta at the time of emergence from the natural redds in a small lowland stream of western Lithuania. The paper also gives shrinkage rates and correction factors for emergent Salmo trutta fry after protracted formalin preservation.     

Life history of a wide‐ranging deepwater lantern shark in the north‐east Atlantic,Etmopterus spinax (Chondrichthyes: Etmopteridae), with implications for conservation

In this paper, the population biology of the velvet belly lanternshark Etmopterus spinax was studied and life‐history coefficients determined. Age was estimated from sections of the second dorsal spine and validated by marginal increment analysis. Males attained a maximum age of 8 years while 11 year‐old females were found. Several growth models were fitted and compared for both size‐at‐age and mass‐at‐age data, showing that even though this is a small‐sized species, it has a relatively slow growth rate. This species matures late, specifically at 49·6 and 42·5% of the maximum observed ages for males and females, respectively. It has a low fecundity, with a mean ovarian fecundity of 9·94 oocytes and a mean uterine fecundity of 7·59 embryos per reproductive cycle. This species seems to have a long reproductive cycle, and even though no conclusive data were obtained, a 2–3 year cycle is possible. The estimated coefficients indicate that this species has a vulnerable life cycle, typical of deepwater squalid sharks. Given the high fishing pressures that it is suffering in the north‐east Atlantic, this fish may already be facing severe declines or in risk of facing them in the near future.     

Ontogenetic variation in density‐dependent growth of brown trout through habitat competition

1. Density‐dependent growth has been widely reported in freshwater fishes, but the ontogenetic evolution of competition and its subsequent effects on growth through a life span remains unclear. 2. Patterns of competition can be described by integrating population abundance data with habitat‐modelling results. Weighted usable area (WUA; m² WUA ha^?1) curves are obtained for each flow value and are then coupled with demographic data to obtain the occupancy rates (trout m^?2 WUA, the density of a given age class related to its suitable habitat) of the WUA for every age class, year and site. 3. We examined a long‐term data series searching for temporal variation in the influence of habitat occupancy rate on the growth of brown trout Salmo trutta. We tested whether (i) mean cohort mass (mean mass of the cohort during the first 3 years of life) is affected by the occupancy rate experienced across a life span; and (ii) the occupancy rate experienced at different ages influenced mean body size. 4. We observed a consistent negative power relationship between average cohort mass and mean occupancy rate through a life span, indicating that stronger cohorts were related to a reduced growth, with likely consequences for individual fitness. 5. The effects of occupancy rate on size‐at‐age were mainly detected in the size attained at the second year of life, but they were because of the competition at different times. Thus, the level of competition varied through ontogeny, in some of the rivers affecting growth since the first year of life, whereas in most of the rivers the main effects on body size resulted from the competition during the second year of life. 6. Occupancy rate appears more appropriate than density for assessing the occurrence of habitat competition in freshwater fishes, since it encompasses the differences in quantity and quality of suitable habitat for each age class. 7. Our study highlights the importance of density‐dependent growth as a key process in the dynamics of brown trout populations, its temporal variation depending on the temporal changes of density and the variation of competition associated with the habitat capacity for each life stage.     

Early Life-History Consequences of Growth-Hormone Transgenesis in Rainbow Trout Reared in Stream Ecosystem Mesocosms

There is persistent commercial interest in the use of growth modified fishes for shortening production cycles and increasing overall food production, but there is concern over the potential impact that transgenic fishes might have if ever released into nature. To explore the ecological consequences of transgenic fish, we performed two experiments in which the early growth and survival of growth-hormone transgenic rainbow trout (Oncorhynchus mykiss) were assessed in naturalized stream mesocosms that either contained predators or were predator-free. We paid special attention to the survival bottleneck that occurs during the early life-history of salmonids, and conducted experiments at two age classes (first-feeding fry and 60 days post-first-feeding) that lie on either side of the bottleneck. In the late summer, the first-feeding transgenic trout could not match the growth potential of their wild-type siblings when reared in a hydrodynamically complex and oligotrophic environment, irrespective of predation pressure. Furthermore, overall survival of transgenic fry was lower than in wild-type (transgenic = 30% without predators, 8% with predators; wild-type = 81% without predators, 31% with predators). In the experiment with 60-day old fry, we explored the effects of the transgene in different genetic backgrounds (wild versus domesticated). We found no difference in overwinter survival but significantly higher growth by transgenic trout, irrespective of genetic background. We conclude that the high mortality of GH-transgenic trout during first-feeding reflects an inability to sustain the basic metabolic requirements necessary for life in complex, stream environments. However, when older, GH-transgenic fish display a competitive advantage over wild-type fry, and show greater growth and equal survival as wild-type. These results demonstrate how developmental age and time of year can influence the response of genotypes to environmental conditions. We therefore urge caution when extrapolating the results of GH-transgenesis risk assessment studies across multiple life-history or developmental stages.     

Growth, carcass composition and plasma growth hormone levels in cyclically fed rainbow trout

Growth, body composition and plasma growth hormone levels were recorded weekly for 24 weeks in rainbow trout Oncorhynchus mykiss . Underyearling rainbow trout were individually identified using coded tags and placed on either a cyclic feeding regime of 3 weeks of deprivation followed by 3 weeks of feeding or a daily feeding regime. No significant difference was found in standard length and mass among the cyclically fed and daily fed fish at the end of the experiment. For cyclically fed fish, the absolute specific growth rate and condition factor reached a maximum during the last week of refeeding. Cyclically fed fish had a significantly higher moisture and protein content and lower lipid levels relative to fish fed daily. Absolute mass and fat loss in the deprivation phase of the feeding cycle decreased in intensity with subsequent feeding cycles, indicating that the fish were acclimatizing to the feeding regime. It was proposed that this response was an adaptation against possible adverse effects in the adults ( e.g. locomotor performance, bone ossification rates, fat deposition rate, growth rate and age at sexual maturity). Plasma growth hormone concentrations were not affected by cyclic feeding indicating that variations in plasma growth hormone concentration are not the cause of compensatory growth in rainbow trout.     

Interacting effects of temperature and density on individual growth performance in a wild population of brown trout

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Stocking experiment with Atlantic salmon and sea trout parr reared on either live prey or a pellet diet

Survival rate and growth parameters of Atlantic salmon fry and sea trout fry were determined after stocking in the wild. Before release (22 May 2009) into the wild the larvae were reared for 10 weeks in the hatchery in three groups: (i) fry fed on live zooplankton , (ii) fry fed on larvae of live nekton, and (iii) fry fed on prepared pellet food. In autumn (15 September 2010) the fish were caught in the wild; the survival rate and growth parameters of both Atlantic salmon and sea trout were the highest in the zooplankton‐fed group, whilst the pellet‐fed group had the lowest survival rate and growth value parameters. Most effective food for hatchery‐reared fishes to be used as stock was the natural living zooplankton. The general conclusion is that the live diet supplied in the rearing period has a positively impact on fish survival in the wild.     

Anthropogenic influences on population sizes, age and growth of naturalized rainbow trout, Oncorhynchus mykiss, in Kenya

Riverine fishery in Kenya has witnessed profound changes since the 1950s, especially after independence (in 1963) when watersheds on the south‐eastern slopes of Mt Kenya were opened up for settlement. In particular, rainbow trout populations have declined essentially resulting from anthropogenic changes through over‐exploitation and/or degraded habitats. While there still are self‐sustaining rainbow trout populations in this stream among others they are in decline an indication that changes that have occurred on the watershed have altered their sizes, age and growth. The study was carried out in the Sagana, a third‐order stream, which rises at about 4000 m altitude on the south‐eastern slope of Mt Kenya, to obtain and document information on population sizes, age and growth of rainbow trout populations and to relate them with those recorded in the 1950s when trout streams were pristine (with little human influence). Stations were fished bi‐monthly from 1996 to 1998 and later for 6 months in 2002. Information on age and growth was obtained by use of annual marks, tagging fish of known age, and by validating their age using captive fish of known age. The minimum mean back‐calculated length at age for age one rainbow trout was 13.09 cm in upstream station and 15.10 cm for downstream stations. However, there was no significant difference in mean back‐calculated lengths at age for all years between fish in upstream and downstream stations (t‐test, t = −0.01, P = 0.99). Although female fish showed higher mean annual back‐calculated length increments than males, there was no significant difference in mean back‐calculated lengths at age between sexes (t‐test, t = −0.27, P = 0.80). The rate of growth in length was rapid for 1‐year‐old fish and declined in the second and third years. This study observed that most of the fish were small with only a few reaching more than 2 years of age because of overfishing. There are good reasons for optimism about the future of trout populations in this stream but concerted efforts are required to rehabilitate them. If trout populations are to increase, a management strategy is required to reduce fishing pressure and to maintain stream fishery against competing needs for resources in the catchments.     

Passing a seawater challenge test is not indicative of hatchery‐reared Atlantic salmon Salmo salar smolts performing as well at sea as their naturally produced conspecifics

Despite satisfactory reactions to seawater challenge tests indicative of appropriate physiological state, hatchery‐reared Atlantic salmon Salmo salar smolts stocked in the Eira River in Norway between 2001 and 2011 performed less well at sea in terms of growth, age at maturity and survival than smolts of natural origin. The mean rates of return to the river for hatchery‐reared and naturally produced S. salar were 0·98 and 2·35%. In the Eira River, c. 50 000 hatchery‐reared S. salar smolts of local origin were stocked annually to compensate for reduced natural smolt production following regulation for hydroelectric purposes, while a mean of 17 262 smolts were produced naturally in the river. This study demonstrates that, although captive S. salar perform well in seawater challenge tests, hatchery‐reared smolts are not necessarily as adaptable to marine life as their naturally produced counterparts. These findings suggest that production of hatchery‐reared smolts more similar to naturally produced individuals in morphology, physiology and behaviour will be necessary to improve success of hatchery releases. Where possible, supplementary or alternative measures, including habitat restoration, could be implemented to ensure the long‐term viability of wild stocks.