Migratory behaviour and survival rates of wild northern Atlantic salmon Salmo salar post‐smolts: effects of environmental factors

To study smolt behaviour and survival of a northern Atlantic salmon Salmo salar population during river descent, sea entry and fjord migration, 120 wild S. salar were tagged with acoustic tags and registered at four automatic listening station arrays in the mouth of the north Norwegian River Alta and throughout the Alta Fjord. An estimated 75% of the post‐smolts survived from the river mouth, through the estuary and the first 17 km of the fjord. Survival rates in the fjord varied with fork length (L_F), and ranged from 97·0 to 99·5% km^?1. On average, the post‐smolts spent 1·5 days (36 h, range 11–365 h) travelling from the river mouth to the last fjord array, 31 km from the river mouth. The migratory speed was slower (1·8 L_F s^?1) in the first 4 km after sea entry compared with the next 27 km (3·0 L_F s^?1). Post‐smolts entered the fjord more often during the high or ebbing tide (70%). There was no clear diurnal migration pattern within the river and fjord, but most of the post‐smolts entered the fjord at night (66%, 2000–0800 hours), despite the 24 h daylight at this latitude. The tidal cycle, wind‐induced currents and the smolts' own movements seemed to influence migratory speeds and routes in different parts of the fjord. A large variation in migration patterns, both in the river and fjord, might indicate that individuals in stochastic estuarine and marine environments are exposed to highly variable selection regimes, resulting in different responses to environmental factors on both temporal and spatial scales. Post‐smolts in the northern Alta Fjord had similar early marine survival rates to those observed previously in southern fjords; however, fjord residency in the north was shorter.     

Riverine and near coastal migration performance of hatchery brown trout Salmo trutta

To study migration performance and return rates of hatchery brown trout Salmo trutta smolts the first 5 months after release, 50 fish in each year (fork length, L_F, 158–288 mm) were in two subsequent years tagged with acoustic transmitters and recorded by automatic listening stations in the River Nidelva (central Norway), its estuary and in the marine environment. More than half of the smolts became anadromous migrants (52% in 2011 and 70% in 2012). The fish spent longer time in the estuary than in the marine environment and the results suggest that migratory behaviour of S. trutta smolts is not only restricted to be resident or anadrome–lacustrine, but that there is also an intermediary strategy of estuarine feeding. There were no differences in L_F or mass between groups of smolts with different migration patterns. Return rates from the sea within the first 5 months after release were in both years 16%. Median progression rate in the river was 0·090 L_F s^?1 but decreased significantly as the smolts entered the estuary (0·015 L_F s^?1). The long residential time in the estuary may increase the risk of negative effects of anthropogenic activities in estuaries, such as harbours and industrial development, and special attention should be given to evaluate effects of such activities.     

Influence of sea temperature and initial marine feeding on survival of Atlantic salmon Salmo salar post-smolts from the Rivers Orkla and Hals, Norway

The abundance of returning adult Atlantic salmon Salmo salar, in the River Orkla in mid‐norway (1 sea‐winter, SW, fish) and River Hals in north Norway (1–3 SW fish), was tested against the early marine feeding and the seawater temperature experienced by their corresponding year classes of post‐smolts immediately after entry into the Trondheimsfjord (Orkla smolts, 22 years of data) and Altafjord (Hals smolts, 17 years of data). In both river–fjord systems, there was a significant positive correlation between the abundance of returning S. salar and the mean seawater temperature at the time of smolts descending to the sea. The number of 1SW fish reported caught in River Orkla was positively correlated to the proportion of fish larvae in the post‐smolt stomachs in Trondheimsfjord. The abundance of returning S.salar was, however, neither correlated to forage ratio (R_F) nor other prey groups in post‐smolt stomachs in the two fjord systems. In the Altafjord, the post‐smolts fed mainly on pelagic fish larva (70–98%) and had a stable R_F (0·009–0·023) over the 6 years analysed. In the Trondheimsfjord, however, there was a higher variation in R_F (0·003–0·036), and pelagic fish larvae were dominant prey in only two (50 and 91%) of the 8 years analysed. These 2 years also showed the highest return rates of S. salar in River Orkla. These results demonstrate that the thermal conditions experienced by post‐smolts during their early sea migration may be crucial for the subsequent return rate of adults after 1–3 years at sea. Pelagic marine fish larvae seem to be the preferred initial prey for S. salar post‐smolts. As the annual variation in abundance of fish larvae is related to seawater temperature, it is proposed that seawater temperature at sea entry and the subsequent abundance of returning adult S. salar may be indirectly linked through variation in annual availability of pelagic fish larvae or other suitable food items in the early post‐smolt phase.     

Does size matter? A test of size‐specific mortality in Atlantic salmon Salmo salar smolts tagged with acoustic transmitters

Mortality rates of wild Atlantic salmon Salmo salar smolts implanted with acoustic transmitters were assessed to determine if mortality was size dependent. The routinely accepted, but widely debated, ‘2% transmitter mass: body mass’ rule in biotelemetry was tested by extending the transmitter burden up to 12·7% of body mass in small [mean fork length (L_F) 138·3 mm, range 115–168 mm] downstream migrating S. salar smolts. Over the short timescale of emigration (range 11·9–44·5 days) through the lower river and estuary, mortality was not related to S. salar size, nor was a relationship found between mortality probability and transmitter mass: body mass or transmitter length: L_F ratios. This study provides further evidence that smolt migration studies can deviate from the ‘2% rule’ of thumb, to more appropriate study‐specific measures, which enables the use of fishes representative of the body size in natural populations without undue effects.     

Maximum sustainable speed, energetics and swimming kinematics of a tropical carangid fish, the green jack Caranx caballus

Maximum sustained swimming speeds, swimming energetics and swimming kinematics were measured in the green jack Caranx caballus (Teleostei: Carangidae) using a 41 l temperature‐controlled, Brett‐type swimming‐tunnel respirometer. In individual C. caballus [mean ±s.d. of 22·1 ± 2·2 cm fork length (L_F), 190 ± 61 g, n = 11] at 27·2 ± 0·7° C, mean critical speed (U_crit) was 102·5 ± 13·7 cm s^?1 or 4·6 ± 0·9 L_F s^?1. The maximum speed that was maintained for a 30 min period while swimming steadily using the slow, oxidative locomotor muscle (U_max,c) was 99·4 ± 14·4 cm s^?1 or 4·5 ± 0·9 L_F s^?1. Oxygen consumption rate (M in mg O₂ min^?1) increased with swimming speed and with fish mass, but mass‐specific M (mg O₂ kg^?1 h^?1) as a function of relative speed (L_F s^?1) did not vary significantly with fish size. Mean standard metabolic rate (R_S) was 170 ± 38 mg O₂ kg^?1 h^?1, and the mean ratio of M at U_max,c to R_S, an estimate of factorial aerobic scope, was 3·6 ± 1·0. The optimal speed (U_opt), at which the gross cost of transport was a minimum of 2·14 J kg^?1 m^?1, was 3·8 L_F s^?1. In a subset of the fish studied (19·7–22·7 cm L_F, 106–164 g, n = 5), the swimming kinematic variables of tailbeat frequency, yaw and stride length all increased significantly with swimming speed but not fish size, whereas tailbeat amplitude varied significantly with speed, fish mass and L_F. The mean propulsive wavelength was 86·7 ± 5·6 %L_F or 73·7 ± 5·2 %L_T. Mean ±s.d . yaw and tailbeat amplitude values, calculated from lateral displacement of each intervertebral joint during a complete tailbeat cycle in three C. caballus (19·7, 21·6 and 22·7 cm L_F; 23·4, 25·3 and 26·4 cm L_T), were 4·6 ± 0·1 and 17·1 ± 2·2 %L_T, respectively. Overall, the sustained swimming performance, energetics, kinematics, lateral displacement and intervertebral bending angles measured in C. caballus were similar to those of other active ectothermic fishes that have been studied, and C. caballus was more similar to the chub mackerel Scomber japonicus than to the kawakawa tuna Euthynnus affinis.     

The critical swimming speed of Iberian barbel Barbus bocagei in relation to size and sex

The swimming capacity of Barbus bocagei was measured with the critical swimming speed (U_crit) standard test in a modified Bla?ka‐type swim tunnel. Sixty B. bocagei were tested and they exhibited a mean ±s .d . U_crit of 0·81 ± 0·11 m s^?1 or 3·1 ± 0·86 total lengths per second (L_T s^?1). Sex had no effect on U_crit but significant differences were found between the swimming performance of fish with distinct sizes.     

Empirical standard mass equation for Salmo marmoratus

Total length (L_T) (range 24–1000 mm; mean ±s.e . = 170·21 ± 0·36 mm) and mass (W) (range 0·10–9590 g; mean ±s.e . = 76·03 ± 0·87 g) of 36 460 specimens of marble trout Salmo marmoratus were used to compute a standard mass (W_s) equation for this species by means of the empirical percentile (EmP) method. The EmP W_s equation calculated was: log₁₀W_s = ?5·208 + 3·202 log₁₀L_T? 0·046 (log₁₀L_T)² (L_T range 90–570 mm) and it is valid throughout the species' area of distribution across Europe.     

Estuarine survival and migratory behaviour of Atlantic salmon Salmo salar smolts

To estimate mortality rates, assess the spatio‐temporal dynamics of natural mortality and examine migratory behaviour during the fresh to saltwater transition, 185 wild Atlantic salmon Salmo salar smolts were implanted with coded acoustic transmitters. Seaward migration of tagged S. salar from four river systems in an area of Nova Scotia, Canada known as the Southern Upland was monitored using fixed receivers and active telemetry over 3 years. Cumulative survival through the river, inner estuary, outer estuary and bay habitats averaged 59·6% (range = 39·4–73·5%). When standardized to distance travelled, survival rates followed two patterns: (1) constant rates of survival independent of habitat or (2) low survival most frequently associated with inner estuary habitats. In rivers where survival was independent of habitat, residency periods were also independent of habitat, post‐smolts exhibited few upstream movements, took a more direct route to the ocean and reached the ocean rapidly. Alternatively, in rivers where survival was habitat specific, residency was also habitat specific with overall increased residency, more frequent upstream movements and delayed arrival to the open ocean. The sudden disappearance of most (75–100%) smolts and post‐smolts assumed dead during the course of this study warrants further examination into the role of avian predators as a mortality vector.     

Ontogeny of critical and prolonged swimming performance for the larvae of six Australian freshwater fish species

Critical (<30 min) and prolonged (>60 min) swimming speeds in laboratory chambers were determined for larvae of six species of Australian freshwater fishes: trout cod Maccullochella macquariensis, Murray cod Maccullochella peelii, golden perch Macquaria ambigua, silver perch Bidyanus bidyanus, carp gudgeon Hypseleotris spp. and Murray River rainbowfish Melanotaenia fluviatilis. Developmental stage (preflexion, flexion, postflexion and metalarva) better explained swimming ability than did length, size or age (days after hatch). Critical speed increased with larval development, and metalarvae were the fastest swimmers for all species. Maccullochella macquariensis larvae had the highest critical [maximum absolute 46·4 cm s^?1 and 44·6 relative body lengths (L_B) s^?1] and prolonged (maximum 15·4 cm s^?1, 15·6 L_B s^?1) swimming speeds and B. bidyanus larvae the lowest critical (minimum 0·1 cm s^?1, 0·3 L_B s^?1) and prolonged swimming speeds (minimum 1·1 cm s^?1, 1·0 L_B s^?1). Prolonged swimming trials determined that the larvae of some species could not swim for 60 min at any speed, whereas the larvae of the best swimming species, M. macquariensis, could swim for 60 min at 44% of the critical speed. The swimming performance of species with precocial life‐history strategies, with well‐developed larvae at hatch, was comparatively better and potentially had greater ability to influence their dispersal by actively swimming than species with altricial life‐history strategies, with poorly developed larvae at hatch.     

Fjord migration and survival of wild and hatchery-reared Atlantic salmon and wild brown trout post-smolts

The behaviour of wild (n = 43, mean L_T = 152 mm) and hatchery-reared (n = 71, mean L_T = 198 mm) Atlantic salmon and wild anadromous brown trout (n = 34, mean L_T = 171 mm) post-smolts with acoustic transmitters was compared in a Norwegian fjord system. There was no difference in survival between wild and hatchery reared salmon from release in the river mouth to passing receiver sites 9.5 km and 37.0 km from the release site. Mortality approached 65% during the first 37 km of the marine migration for both groups. There was no difference between wild and hatchery-reared salmon either in time from release to first recording at 9.5 km (mean 135 and 80 h), or in the rate of movement through the fjord (mean 0.53 and 0.56 bl s⁻¹). Hatchery-reared salmon reached the 37 km site sooner after release than the wild salmon (mean 168 and 450 h), but rate of movement in terms of body lengths per second did not differ (mean 0.56 and 0.77 bl s⁻¹). The brown trout remained a longer period in the inner part of the fjord system, with much slower rates of movement during the first 9.5 km (mean 0.06 bl s⁻¹).     

Effects of surgically implanted acoustic transmitters >2% of body mass on the swimming performance, survival and growth of juvenile sockeye and Chinook salmon

The influence of surgical implantation of an acoustic transmitter on the swimming performance, growth and survival of juvenile sockeye salmon Oncorhynchus nerka and Chinook salmon Oncorhynchus tshawytscha was examined. The transmitter had a mass of 0·7 g in air while sockeye salmon had a mass of 7·0–16·0 g and Chinook salmon had a mass of 6·7–23·1 g (a transmitter burden of 4·5–10·3% for sockeye salmon and 3·1–10·7% for Chinook salmon). Mean critical swimming speeds (U_crit) for Chinook salmon ranged from 47·5 to 51·2 cm s^?1 [4·34–4·69 body lengths (fork length, L_F) s^?1] and did not differ among tagged, untagged and sham‐tagged groups. Tagged sockeye salmon, however, did have lower U_crit than control or sham fish. The mean U_crit for tagged sockeye salmon was 46·1 cm s^?1 (4·1 L_F s^?1), which was c. 5% less than the mean U_crit for control and sham fish (both groups were 48·6 cm s^?1 or 4·3 L_F s^?1). A laboratory evaluation determined that there was no difference in L_F or mass among treatments (control, sham or tag) either at the start or at the end of the test period, suggesting that implantation did not negatively influence the growth of either species. None of the sockeye salmon held under laboratory conditions died from the influence of surgical implantation of transmitters. In contrast, this study found that the 21 day survival differed between tagged and control groups of Chinook salmon, although this result may have been confounded by the poor health of Chinook salmon treatment groups.     

Short‐term survival and movements of Atlantic sharpnose sharks captured by hook‐and‐line in the north‐east Gulf of Mexico

Ultrasonic telemetry was used to compare post‐release survival and movements of Atlantic sharpnose sharks Rhizoprionodon terraenovae in a coastal area of the north‐east Gulf of Mexico. Ten fish were caught with standardized hook‐and‐line gear during June to October 1999. Atlantic sharpnose sharks were continuously tracked after release for periods of 0·75 to 5·90 h and their positions recorded at a median interval of 9 min. Individual rate of movement was the mean of all distance and time measurements for each fish. Mean ± s.e . individual rate of movement was 0·45 ± 0·06 total lengths per second (L_T s^?1) and ranged from 0·28 to 0·92 L_T s^?1 over all fish. Movement patterns did not differ between jaw and internally hooked Atlantic sharpnose sharks. Individual rate of movement was inversely correlated with bottom water temperature at capture (r² = 0·52, P ≤ 0·05). No consistent direction in movement was detected for Atlantic sharpnose sharks after release, except that they avoided movement towards shallower areas. Capture‐release survival was high (90%), with only one fish not surviving, i.e. this particular fish stopped movement for a period of 10 min. Total rate of movement was total distance over total time (m min^?1) for each Atlantic sharpnose shark. Mean total rate of movement was significantly higher immediately after release at 21·5 m min^?1 over the first 1·5 h of tracking, then decreased to 11·2 m min^?1 over 1·5–6 h, and 7·7 m min^?1 over 3–6 h (P ≤ 0·002), which suggested initial post‐release stress but quick recovery from capture. Thus, high survival (90%) and quick recovery indicate that the practice of catch‐and‐release would be a viable method to reduce capture mortality for R. terraenovae.     

Slow growth of the overexploited milk shark Rhizoprionodon acutus affects its sustainability in West Africa

Age and growth of Rhizoprionodon acutus were estimated from vertebrae age bands. From December 2009 to November 2010, 423 R. acutus between 37 and 112 cm total length (L_T) were sampled along the Senegalese coast. Marginal increment ratio was used to check annual band deposition. Three growth models were adjusted to the length at age and compared using Akaike's information criterion. The Gompertz growth model with estimated size at birth appeared to be the best and resulted in growth parameters of L_∞ = 139·55 (L_T) and K = 0·17 year^?1 for females and L_∞ = 126·52 (L_T) and K = 0·18 year^?1 for males. The largest female and male examined were 8 and 9 years old, but the majority was between 1 and 3 years old. Ages at maturity estimated were 5·8 and 4·8 years for females and males, respectively. These results suggest that R. acutus is a slow‐growing species, which render the species particularly vulnerable to heavy fishery exploitation. The growth parameters estimated in this study are crucial for stock assessments and for demographic analyses to evaluate the sustainability of commercial harvests.     

Age, growth, and sexual maturity of the yellownose skate Dipturus chilensis in the south‐eastern Pacific

The age and growth parameters of Dipturus chilensis were estimated by counting growth rings from thin sections of vertebral centra from 400 fish (246 females and 154 males), ranging from 23 to 124 cm total length (L_T), and backcalculating fish lengths at previous ages. Marginal increment analysis lent support to the hypothesis of annual deposition of band‐pairs, which formed during the winter months. The oldest female D. chilensis aged in this study was 21 years and 117 cm L_T, whereas the oldest male was 18 years and 93 cm L_T. A 4·7% index of average per cent error (I_APE) suggested that this is a precise method for calculating the age of D. chilensis. Observed L_T were lower than backcalculated L_T, which implies the influence of Lee's phenomenon. The von Bertalanffy growth equations, based on mean length‐at‐age data, were estimated as L_t = 128·3 (1 ? e^{?0·112 (t + 0·514)}) for females and L_t = 107·8 (1 ? e^{?0·134 (t + 0·862)}) for males where t is age (years). Growth was significantly different between sexes: females reached a larger adult size. Ages and lengths at 50% maturity were estimated at 14 years of age and 106 cm L_T for females and 11 years of age and 86 cm L_T for males. At c. 14 years, there was a decline in growth rates in females. The factor most likely responsible for this was sexual maturity, which caused a discontinuity in growth of female fish. These results show that this species is slow‐growing, long‐lived, relatively large and of delayed maturity, characteristics that make it vulnerable to exploitation.     

Evidence of changing migratory patterns of wild Atlantic salmon Salmo salar smolts in the River Bush,Northern Ireland,and possible associations with climate change

The migration patterns, timing and biological characteristics of wild Atlantic salmon Salmo salar smolts in the River Bush, Northern Ireland, were examined over the period 1978–2008. A distinct change in the timing of the smolt run was detected with progressively earlier emigration periods evident across the time series. The shift in run timing ranged from 3·6 to 4·8 days 10 years^?1 for a range of standard migratory audit points. The timing of smolt emigration has been linked to ambient river temperature patterns. Distinct seasonal patterns were evident for biological characteristics of River Bush smolts with mean age and fork length decreasing throughout the emigration period. Marine survival patterns in 1 sea winter River Bush S. salar were strongly influenced by the run timing of the preceding smolt year such that later emigrating cohorts demonstrated increased survival. Possible mechanisms for this relationship based on local climatic variation have been explored, including the effect of potential thermal mismatch between freshwater and marine environments.     

Critical swimming speeds of wild‐caught sand‐smelt Atherina presbyter larvae

Swimming abilities of wild‐caught sand‐smelt Atherina presbyter larvae were assessed as critical swimming speed (U_crit) throughout ontogeny. The mean U_crit increased with size, ranging from 3·6 to 18·7 cm s^?1, over the size range of 6·6–21·0 mm L_T. This indicates that at hatching A. presbyter larvae, far from being passive floaters, are already capable of active behaviours, which may influence their dispersal patterns.     

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.     

Migration and survival of Atlantic salmon Salmo salar smolts in a large natural lake

An investigation with acoustic telemetry of the passage of Salmo salar smolts through a large natural lake found heavy mortality occurred at the river‐to‐lake confluences (mean 31.2% km^?1), but was lower in the main body of the lake (mean 2.4% km^?1). Predation was a significant pressure on emigrating smolts as tagged pike Esox lucius aggregated at river‐to‐lake confluences during the peak of the smolt run. Tagged smolts mainly emmigrated into the lake in the late evening after dusk, possibly as a predator‐avoidance behaviour.     

Manipulation of the energetic state of Atlantic salmon Salmo salar juveniles and the effect on migration speed

Atlantic salmon Salmo salar smolts were produced with similar energetic states as wild S. salar and the effect of low energetic state on smolt migration was tested. The total energetic state of the fish (body lipids and proteins) in the spring was correlated with Fulton's condition factor (K). Fish at a low energetic state swam slower but migrated further than fish at a higher energetic state when tested in two experimental streams. During a period of starvation throughout the winter and spring, fish conserved their body‐lipid reserves at 1·5% by using more protein as an energy source and the metabolic shift occurred between 3·5 and 1·5% body lipids. An energetic state of approximately 3·5 kJ g^?1 (K ≈ 0·65) appeared to be the critical limit for survival.