Assessing the food web impacts of an anadromous Arctic charr introduction to a sub‐Arctic watershed using stable isotopes

Anadromous Arctic charr, Salvelinus alpinus (L.), was introduced to a sub‐Arctic river–lake system near the village of Kujjuuaq, Nunavik, and the stable isotope values and diets of key resident fish species were used to assess changes in feeding patterns. Stable isotope values for most species did not differ significantly between the pre‐ and post‐introduction periods, with observed shifts being within the bounds of expected natural variation. Lake chub, Couesius plumbeus (Agassiz), were the single species to show a difference between study periods, with a small but significant increase in δ¹⁵N. No significant post‐introduction changes were seen in lake trout, Salvelinus namaycush (Walbaum), omnivory or in any of the assessed quantitative food web metrics. Gut contents of major fish species similarly showed significant temporal overlap between the pre‐ and post‐introduction periods, and there was no significant change in species' weight–length relationships. The minor ecological impact was interpreted in relation to the availability of open niches exploitable by ecological generalists such as Arctic charr. The explanation accords with the known habitat and feeding flexibility of Arctic charr and the ecological immaturity of sub‐Arctic lakes known to have driven adaptive variation among Arctic charr. Findings suggest that anadromous Arctic charr may be introduced at moderate densities to other sub‐Arctic watersheds without major negative food web consequences for other resident fish species.     

The suitability of anadromous Arctic charr as host and vector of the monogenean Gyrodactylus salaris

Abstract –  In 2000, the Atlantic salmon ( Salmo salar L.) stock in the river Signaldalselva, North Norway was found to be infected with the monogenean Gyrodactylus salaris (Malmberg). This river system has a multispecies fish community of nine species in total. Low densities of Atlantic salmon parr were found during electrofishing in October (2003) with a prevalence of 94% and a mean abundance of 848 G. salaris. This watercourse also holds a riverine stock of anadromous Arctic charr ( Salvelinus alpinus ), which is not common in Norway. Among the parr of Arctic charr, 64% were infested with an abundance of 23 parasites per fish. The most heavily infected 0+ and 1+ juvenile Arctic charr had 469 and 534 parasites, respectively, indicating that the parasites are able to reproduce on Arctic charr in the wild. There was little variation in the rates of infestation of Arctic charr along the studied stretch of the river, even in areas with very low densities of Atlantic salmon parr. The parr of anadromous Arctic charr seem to be suitable as long-term hosts for G. salaris . Moreover, a few (22%) infested adult sea-running Arctic charr were captured after ascending the river in the autumn, which shows that large fish may also act as carriers of G. salaris . However, no parasites were recorded on potential anadromous Arctic charr before descending during early spring (April 2003 and April 2004). Thus, it is still not clear whether anadromous Arctic charr are able to disperse the parasite between watercourses. So far, it is uncertain whether G. salaris may increase the mortality rates of Arctic charr and thereby be a threat to these unusual occurring riverine anadromous Arctic charr stocks.     

Latitudinal variation in growth and otolith‐inferred field metabolic rates of Canadian young‐of‐the‐year Arctic charr

Countergradient variation (CGV) is defined as genetic variation that counteracts the negative influences of the physical environment, minimising phenotypic variability along an environmental gradient. CGV is thought to have relevance in predicting the response of organisms to climate variability and change. To test the hypothesis that growth rate increased with latitude, consistent with CGV, young‐of‐the‐year (YOY) Arctic charr, Salvelinus alpinus, were examined along a ~27° latitudinal gradient in central and eastern Canada. Growth rates were estimated from fork lengths standardised by the thermal opportunity for growth based on experienced water temperatures derived using otolith oxygen stable isotopes. Results demonstrated patterns consistent with CGV, where northern populations demonstrated faster growth rates. A secondary aim was to test for similar geographical patterns in otolith‐inferred metabolic rates, which reflect the energetic costs of standard metabolic rate (SMR) and other processes such as feeding, locomotion, thermoregulation, reproduction and growth. Results demonstrated a significant, positive relationship between otolith‐inferred metabolic rate and latitude, which may reflect an increase in one, or a combination, of the above‐noted physiological processes. The similar latitudinal pattern in growth and otolith‐inferred metabolic rates suggests greater intake of food per unit of time by northern fish. The phenotypic variation in physiological traits observed here demonstrates the significant adaptability of Arctic charr to different thermal regimes with different growing season lengths. Determining the relative contributions of phenotypic plasticity and genetic variation to the observed latitudinal variation will be critical to predicting the responses of Arctic charr to climate change more accurately.     

Evidence for genetic distinction among sympatric ecotypes of Arctic char (Salvelinus alpinus) in south‐western Alaskan lakes

Resource polymorphism may play an important role in the process of speciation. The Arctic char (Salvelinus alpinus) exhibits great phenotypic and genetic diversity across its range, making it an ideal species for studies of resource polymorphism and divergence. Here, we investigated genetic variation at 11 microsatellite loci among 287 Arctic char from five isolated yet proximate postglacial lakes in south‐western Alaska that were previously examined for resource polymorphism. Significant differences in pairwise F_ST were detected among all lakes (range from 0.05 to 0.28, all P < 0.02). In one lake (Lower Tazimina Lake), we found evidence for two genetic groups of char and for significant differences in the distribution of microsatellite variability among at least two of the three previously described body size morphotypes (‘large’‐, ‘medium’‐, and ‘small’‐bodied char; maximum F_ST = 0.09; differences in admixture proportions). We also found a significant association between genetic admixture proportions and gill raker counts among body size morphs (r = ?0.73, P < 0.001). Our data represent the first record of genetically distinct sympatric morphs of Arctic char in Alaska and provide further evidence that differences in morphology associated with feeding (gill rakers) and growth trajectories reflect niche diversification and promote genetic divergence in Holarctic populations of Arctic char.     

The effect of lake morphometry on thermal habitat use and growth in Arctic charr populations: implications for understanding climate‐change impacts

Oxygen stable isotope temperature reconstruction methods were used to estimate mean experienced summer temperatures from growth zones within individual Arctic charr otoliths sampled from lakes with contrasting morphologies but proximate locations. For either lake, otolith‐estimated temperatures were not significantly related to back‐calculated growth. Fish in the smaller lake evidenced an increase in growth with age related to increasing use of cooler thermal habitats, with the use of thermal habitat possibly governed by predation risks. No relationships between age, growth or temperature were observed in the larger lake. Significant negative effects on back‐calculated growth were observed due to increasing air temperatures in the smaller and shallower lake, possibly owing to warmer surface and littoral waters and a limited amount of preferred cool‐water habitat. A similar relationship was not observed in the larger and deeper lake and indicated that resident Arctic charr were not as vulnerable to the impacts of temperature warming, possibly because of better behavioural thermoregulation opportunities in the cooler, deeper lake. Results provide evidence for differing climate‐influenced growth outcomes among proximately located populations, with outcomes likely to depend on the differences among habitats, including lake size and morphometry which may act to influence fish densities in available preferred thermal habitats.     

Growth and diet of anadromous Arctic charr after their return to freshwater

Abstract – The post smolt of anadromous Arctic charr ( Salvelinus alpinus ) from Lake Storvatn, northern Norway (70°39'48"), continued to feed and grow after their return to freshwater in late summer, but in late autumn (October) both their growth rate and feeding intensity ceased along with the drop in water temperature. In contrast, adult, sexually mature anadromous charr did not appear to feed during their freshwater residency. The stomach fullness of post smolts were similar to that of resident Arctic charr during early autumn, and their diets were similar, being dominated by pelagic prey such as surface insects, Holopedium sp., Bosmina sp. and chironomid pupae. Later in the autumn, the resident charr also included some more benthic prey, in particular gastropods, whereas the post smolts continued to feed predominantly on zooplankton and surface insects, but with a lower stomach fullness than the resident fish. This suggests that after arrival in freshwater, the post smolts continue their pelagic feeding behaviour displayed during the sea-migration.     

Summer survival and activity patterns of estuary feeding anadromous Salmo trutta

There is generally a very poor understanding of how anadromous brown trout, Salmo trutta, use marine ecosystems. In this study, we use acoustic telemetry techniques to test four alternative hypotheses for estuarine coastal habitat use by this species on a population in the Clyde, west-central Scotland. Anadromous brown trout in their second (or more) summer feeding in the marine environment did not show patterns of estuarine use predicted from studies on other populations of this species. They did not (a) use the inner Clyde Estuary as a staging post for onward migration to open coastal waters, nor did they (b) pass through the estuary rapidly nor did they (c) make multiple incursions into freshwater. In contrast, fish utilised a very geographically constrained area of the inner estuary (<8 km in length), within which fish were highly mobile, exhibiting more activity during a flooding tide and exhibited high survivorship compared with reports from other studies. The reasons why estuarine use by anadromous brown trout in the Clyde may differ from that reported elsewhere is discussed.     

Temperature–growth patterns of individually tagged anadromous Arctic charr Salvelinus alpinus in Ungava and Labrador,Canada

Individual measurements of annual, or within‐season growth were determined from tag‐recaptured Arctic charr and examined in relation to summer sea surface temperatures and within‐season capture timing in the Ungava and Labrador regions of Eastern Canada. Differences between two years of growth (2010–2011) were significant for Ungava Bay Arctic charr, with growth being higher in the warmer year. Growth of Labrador Arctic charr did not vary significantly among years (1982–1985). Regional comparisons demonstrated that Ungava Arctic charr had significantly higher annual growth rates and experienced warmer temperatures than Labrador Arctic charr. The higher annual growth of Ungava Bay Arctic charr was attributed to the high sea surface temperatures experienced in 2010–2011 and the localised differences in nearshore productivity as compared to Labrador. Within‐season growth rates of Labrador Arctic charr peaked in June, declined towards August and were negatively correlated with the length of time spent at sea and mean experienced sea surface temperatures. A quadratic model relating growth rate to temperature best explained the pattern of within‐season growth. Collectively, results suggest that increases in water temperature may have profound consequences for Arctic charr growth in the Canadian sub‐Arctic, depending on the responses of local marine productivity to those same temperature increases.     

Otolith size differences during early life of dwarf and cannibal Arctic char (Salvelinus alpinus)

In Arctic and alpine lakes, Arctic char [Salvelinus alpinus (L.)] often form two distinct morphs: invertebrate feeders (‘dwarfs’) and piscivores (‘giant or cannibals’). Here, we test for early life history growth variation in dimorphic Arctic char as a proximate explanation for the observed life history variation between the two forms. Char were sampled in 11 alpine and Arctic Norwegian lakes. Dwarfs (defined as sexually mature char less than 15 cm long; N = 304) had a mean total length of 105 mm, whereas the typical cannibal (body length above 20 cm; N = 153) was 388 mm long. A positive correlation between egg size and otolith hatching ring were ascertained in a separate hatching experiment with brown trout (Salmo trutta) and it is assumed that this relationship also is valid for Arctic char, and otolith size was used as a proxy for length. Otolith hatching ring from Arctic char cannibals was larger (mean ± SD; 187 ± 24 μm) than those from dwarfs (mean ± SD; 164 ± 23 μm). There were only minor size differences between dwarfs and cannibals during the next three years, after which dwarfs usually matured. Two mutually nonexclusive, proximate explanations for the differentiation into separate morphs (dwarfs and cannibals) are therefore maternal effects and/or genetic based differentiation. The high catchability of large piscivorous char and low production in alpine and Arctic lake ecosystems may make these stocks particularly vulnerable to overexploitation.     

Novel insights into the marine phase and river fidelity of anadromous twaite shad Alosa fallax in the UK and Ireland

1. Most research on anadromous fishes has been invested in their freshwater life‐phases, resulting in a relatively sparse understanding of their spatial ecology during marine life‐phases. However, understanding the marine dispersal of anadromous fishes is essential to identify threats and to implement conservation measures that fully encompass their lifecycle.
2. The twaite shad Alosa fallax is an anadromous fish increasingly imperilled across its range due to pollution, harvesting, and impediments to freshwater migration, but little is known about its distribution and movements during its marine life‐phase. Here, the application of acoustic telemetry provided novel insights into the coastal dispersal of twaite shad in the UK and Ireland during 2018–2019, and the freshwater entry of individuals during the 2019 spawning season.
3. Of 73 twaite shad acoustic‐tagged during their upstream migration in the River Severn in May 2018, 58 emigrated from the river. Twelve were subsequently detected 200 km to the south‐west at the Taw–Torridge Estuary between July 2018 and April 2019, where estuarine movements up to 5.8 km inland occurred in summer, winter, and spring. One was subsequently detected in the Munster Blackwater Estuary (Ireland) and then in the River Severn, indicating a minimum movement distance of 950 km. Thirty‐four (59%) of the emigrating individuals from 2018 re‐entered fresh water in the rivers Severn (n = 33) and Wye (n = 2) in April and May 2019.
4. These results suggest year‐round use of estuarine and nearshore habitats by at least a subset of the twaite shad population during their marine phase, providing evidence of potential range overlap between populations that spawn in different areas in the UK and Ireland, which may be facilitated by substantial dispersal. The results also highlight the potential of telemetry for estimating freshwater and marine mortality, and the benefits of sharing detection data across networks.

     

Effect of salinity at constant 10 °C on grow-out of anadromous Arctic charr from Labrador

The goal is to determine the requirements allowing cultured Salvelinus alpinus to thrive in seawater, as they do in the wild. In late-June, eight families of individually identified 1+ year-old charr (mean wt: 427 g) of a domesticated strain derived from the Fraser River population were directly transferred from freshwater (9 °C) to salinities of either 0, 10, 20 and 30 ppt at 10 °C, then on-grown in tanks until December. Cumulative mortality was 16% in 30 ppt salinity, and < 4% in the lower salinities. Repeated measures analysis revealed somatic growth was inhibited by both elevated salinity and sexual maturation. Among immature fish, final mean weight and condition factor in 30 ppt salinity was 490 g and 1.2, compared to nearly 1 kg and > 1.7 in 0 and 10 ppt. In 20 ppt salinity, growth was initially similar to that in ≤ 10 ppt salinity but deteriorated from September onwards. Sexually maturing fish in ≤ 10 ppt salinity attained a final mean weight about 35% less than immatures. Plasma osmolality was only slightly elevated in the 30 ppt salinity, remaining < 340 mOsm kg^− 1. Food intake and conversion were affected by the interaction between salinity and time, being optimum in 0 and 10 ppt treatments. Family effects on final body size were large, but the effect of salinity on growth was independent of family. In conclusion, despite their large body size, direct transfer of this strain from freshwater to seawater does not appear viable for commercial aquaculture.     

Post-release movement and diel activity patterns of hatchery-reared and wild black-spot tuskfish <Emphasis Type="Italic">Choerodon schoenleinii</Emphasis> determined by ultrasonic telemetry

ABSTRACT:   Post-release movement and diel activity patterns of hatchery-reared and wild black-spot tuskfish were examined using ultrasonic telemetry. Five hatchery-reared and four wild fish were released in the sandy bottom of Urasoko Bay, Ishigaki Island, Okinawa, Japan, and monitored using automated monitoring receivers from November 2005 to February 2006. Both hatchery-reared and wild fish tended to stay near the release site for over two weeks, before leaving the release site. Both hatchery-reared and wild tuskfish showed diurnal rhythm intermittently; signals were recorded more frequently in the daytime and less frequently in the nighttime, suggesting that the fish of both origins were active during the day and inactive during the night. These findings indicate that the one-year-old hatchery-reared tuskfish have some consistent behavioral characteristics with those of the wild.     

Lipid dynamics in anadromous Arctic charr, Salvelinus alpinus (L.): seasonal variations in lipid storage depots and lipid class composition

Changes in lipid class composition of tissues of anadromous Arctic charr, Salvelinus alpinus, were studied in relation to seasonal patterns of lipid deposition and lipid mobilisation. Charr of a north Norwegian population were collected in late May when fish were migrating towards the sea, and in mid-July when the fish re-entered fresh water. Some maturing fish captured in mid-July were held in captivity until late September to assess lipid dynamics associated with maturation. The carcass (head, fins, skeleton and skin) accounted for approx. 50% of the total body lipid content when the fish re-entered fresh water from the sea. Triacylglycerols (TAG) comprised 75-80% of the carcass lipid in ascending charr, confirming that the carcass is a major lipid depot. The skeletal muscle also represented a major lipid store: muscle of ascending charr contained 35-40% of the body lipids, with TAG being dominant (approx. 75% of muscle lipid). The gut (gastrointestinal tract and visceral lipid) and liver of ascending charr each held only 4-5%; of the body lipid, TAG being the dominant lipid class in both of these depots (Gut TAG: 60-65%; Liver TAG: 50-55%). Body lipid decreased 30-40% during maturation: lipids were depleted from all depots, but mobilisation of from the carcass and muscle was quantitatively the most important. In mature females, the ovaries held almost 25% of the body lipid, with the proportion of TAG exceeding that of polar lipids (PL)(TAG: 62%; PL 12.4%). In the mature males, the testes accounted for less than 3% of the total lipids. TAG tended to dominate the testis lipids (TAG: 29.5%), but there were also quite high proportions of free fatty acids, cholesterol and polar lipids (FFA: 23.5%; C: 15%; PL 9.7%). Females lost approx. 80% of their body lipid during spawning and overwintering, the majority of this being TAG. Males also lost large amounts of TAG during the winter, but in contrast to the females lipid depletion amounted to only 50-55% of that present at the time of ascent into fresh water.     

Ecological influences on the difference in δ15N and δ13C values between fish tissues: implications for studies of temporal diet variation

Temporal diet changes can be detected by comparing δ¹³C and δ¹⁵N values between tissues with different isotopic turnover times. However, other factors contribute to disparities in δ¹³C and δ¹⁵N signatures between tissues and could confound the interpretation of stable isotope data. We examined the effects of ecological factors on differences in muscle and liver δ¹³C and δ¹⁵N signatures of Arctic charr [Salvelinus alpinus, (L.)] to determine whether their effects were large enough to influence studies of diet change. In addition, we examined whether differences in lipid content accounted for a significant portion of the differences in δ¹³C values between tissues. Results indicated that life history, diet, reproductive status and gender had significant effects on the differences in δ¹³C values between tissues, while gender and diet significantly affected differences in δ¹⁵N values. Differences in % lipid content between tissues also explained approximately 74% of the observed variation in the difference in δ¹³C values between tissues. The differences in δ¹³C values observed between muscle and liver tissues may be partially explained by physiological differences in lipid storage and use between tissues associated with ecological factor effects. These results demonstrate that ecological factors other than a change in diet can influence the differences in δ¹³C and δ¹⁵N signatures between muscle and liver tissues. However, the impact of ecological factors on studies of diet change depends on how the magnitude of the factor effects compared with a biologically meaningful change in diet.     

Winter residency and activity patterns of channel catfish, Ictalurus punctatus (Rafinesque), and common carp, Cyprinus carpio L., in a thermal discharge canal

Radiotelemetry was used to examine the behavioural responses of fish to dynamic environmental and operating conditions in a fossil-generating power station discharge canal on Lake Erie. A series of seven underwater antennas continuously monitored the movements of 10 common carp, Cyprinus carpio L., and 10 channel catfish, Ictalurus punctatus (Rafinesque), in the 550-m discharge canal between 23 December 1997 and 1 May 1998. Common carp residency values were higher (47.71±8.51%) than those of channel catfish (10.15±2.82%). While resident in the discharge canal, channel catfish spent more time in the upper reaches and were more active than carp. Carp spent more time in the lower reaches of the channel, and made more frequent movements between the canal and Lake Erie. Overall residency decreased significantly during the study period. Aside from significant negative correlations between common carp residency and daily mean influent temperature ( r = −0.53), daily discharge mean ( r = −0.30) and minimum water temperature ( r = −0.47), and positive correlations to daily mean cooling water flow ( r = 0.38), the activity and residency of tagged individuals were independent of environmental and operating conditions. There was no relationship between the total length of each fish, and any of the residency or activity indices. The implications of extended fish residency in heated discharges and the utility of radiotelemetry as a monitoring procedure are discussed.     

Lake trout (Salvelinus namaycush) otoliths indicate effects of climate and lake morphology on growth patterns in Arctic lakes

Climate change is occurring rapidly in the Arctic, and an improved understanding of the response of aquatic biota and ecosystems will be important for this data-limited region. Here, we applied biochronology techniques and mixed-effects modelling to assess relationships among growth increments found on lake trout (Salvelinus namaycush) otoliths (N = 49) captured from 13 lakes on the Arctic Coastal Plain of northern Alaska, observed and modelled climate patterns, and individual-level fish and lake characteristics. We found that annual growth varied by year, fish growth slowed significantly as individuals aged, and females grew faster than males. Lake trout had higher growth in flow-through lakes relative to lakes that were perennially or seasonally connected. Annual growth was positively correlated with observed air temperature measurements from a local weather station for the period 1998–2014, but no clear warming trend was evident for this period. Modelled August air temperatures from 1978–2014 predicted lake trout annual growth (root mean squared error = 0.045 mm) and indicated increasing temperatures and annual lake trout growth over the period 1950–2014. This study demonstrated that biochronology techniques can reconstruct recent climate patterns and provide a better understanding of trends in Arctic lake ecosystems under a changing climate.     

Summer habitat use and feeding of juvenile Arctic charr,Salvelinus alpinus,in the Canadian High Arctic

Abstract – Owing to limited knowledge of the habitat use and diet of juvenile Arctic charr from the High Arctic, particularly young‐of‐the‐year (YOY), we assembled data obtained from samples taken in and around Lake Hazen, Nunavut, Canada, to assess juvenile habitat use and feeding. Juvenile charr demonstrated a preference for stream environments, particularly those fed by warm upstream ponds. Charr occupying both stream and nearshore lake habitats were found to feed similarly, with chironomids occurring most frequently in diets. Some older stream‐dwelling charr preyed on smaller, younger Arctic charr. Preferred stream occupancy is likely mediated by physical barriers created mainly by water velocity, and by distance from the lake, lake‐ice dynamics, low water depth, and turbidity. Water velocities resulted in stream habitat segregation by size, with YOY mainly found in low‐velocity pools and back eddies adjacent to stream banks, but not in water velocities >0.1 m·s^?1. Greatest charr densities in streams were found in small, shallow, slow‐flowing side channels, which are highly susceptible to drought. Under predicted climate change scenarios, streams fed by small ponds will be susceptible to intermittent flow conditions, which could result in increased competition among juvenile charr for the remaining stream habitats. In addition, glacier‐fed streams are likely to experience increased flow conditions that will exacerbate physical barriers created by water velocity and further reduce the availability of preferred stream habitat.     

An experimental comparison of cannibalistic response in different Arctic charr (Salvelinus alpinus (L.)) stocks

Abstract– The cannibalistic response of four domesticated Arctic charr stocks were compared experimentally using a radiographic technique. Charr from Arctic Spitsbergen (80°N) consistently exhibited a much higher frequency of cannibalistic response than fish from three charr strains from mainland Norway (70°N). The results indicate that the extent of cannibalistic feeding in Arctic charr is population specific and support the hypothesis that charr from Arctic populations have stronger cannibalistic tendencies than do fish from more temperate regions.