Decline of the Migratory Form in Bull Charr, Salvelinus Confluentus, and Implications for Conservation

Large-bodied, migratory life history forms of bull charr, Salvelinus confluentus, were historically abundant in northwestern North America, but many remaining populations of this now-threatened species presently persist as small-bodied residents isolated in headwater streams. We examined whether the migratory form has been lost from headwater populations of bull charr and their potential for re-establishment. Upstream and downstream movement of bull charr and other salmonids from three tributary populations in the Bitterroot River drainage, Montana, was measured with weirs over a 17-month period. The migratory life history was rare or absent in two tributaries but still present at a low level in a third. In contrast, substantial numbers (n = 1745) of juvenile and adults of other salmonids (brown trout, Salmo trutta, cutthroat trout, Oncorhynchus clarki, and mountain whitefish, Prosopium williamsoni) were captured near tributary mouths, indicating a migratory life history was common in other species. Apparent decline of the migratory life history in bull charr was not directly related to damming suggesting other downstream mortality factors (predation, temperature) also are involved. Isolated, nonmigratory forms have increased risk of extinction, and restoration of the population connectivity via the re-establishment of migratory stocks is an important conservation goal for bull charr recovery. However, the factors governing migratory tendency remain unclear.     

Competitive interactions for foraging microhabitat among introduced brook charr, Salvelinus fontinalis, and native bull charr, S. confluentus, and westslope cutthroat trout, Oncorhynchus clarki lewisi, in a Montana stream

Competitive interactions for foraging microhabitat among introduced brook charr, Salvelinus fontinalis, and native bull charr, S. confluentus, and westslope cutthroat trout, Oncorhynchus clarki lewisi, were studied by species removal experiments in a tributary of the Flathead Lake and River system, northwestern Montana, focusing on brook charr influences on bull charr. When the three species were in sympatry, they interacted with each other, forming a size-structured, mixed-species dominance hierarchy in two stream pools. The influences of interference interactions were examined by measuring changes in five characteristics of foraging microhabitat and behavior, focal point height and velocity, cover use, and foraging rate and distance, after the successive removal of two species. Cutthroat trout removal resulted in increased foraging rates and distances, and decreased cover use for brook charr, but no changes for bull charr. After removal of brook charr from the two-species system, bull charr also increased foraging rates and distances and occupied more exposed positions. Moreover, total fish densities, which had initially decreased owing to the removal experiments, were partly compensated for by subsequent bull charr immigration, implying that competitive interactions with brook charr are an important factor in the mechanisms responsible for the regulation of bull charr densities, at least on a local scale.     

Genetic diversity and population structure of white-spotted charr, Salvelinus leucomaenis, in the Lake Biwa water system inferred from AFLP analysis

The genetic variations—and the time dependence of such variations—of natural populations of the white-spotted charr ,Salvelinus leucomaenis, in the Lake Biwa water system as well as those of a hatchery-reared population were inferred from AFLP. Upon the application of principal coordinate analysis using 118 polymorphic AFLP fragments based on the Jaccard similarity index, specimens of each of six natural local populations from the inlet rivers of Lake Biwa grouped roughly together, suggesting that each local population was genetically differentiated. The hatchery-reared population was shown to be closely related to the local population in the Seri River, suggesting that the Seri River population originated from hatchery-reared charr due to extensive stocking. Furthermore, specimens of the Yasu River grouped in a somewhat different position from the other natural populations, agreeing well with its geographic distance from the other populations. The nucleotide diversities of six natural populations (Harihata River, Ishida River, two reaches of the Takatoki River, Ane River, and Yasu River) in 2002 or 2003 were relatively low (π = 0.067–0.146%) compared with that of the Seri River (0.278%) and the hatchery-reared charr (0.316%). The nucleotide diversity in the five local populations (Ishida River, two reaches of the Takatoki River, Ane River, and Yasu River) remained at a low level from 1994 to 2002/2003, but only the nucleotide diversity in the Harihata River actually decreased. From 1994 to 2002/2003, the nucleotide diversity in the Seri River remained at a higher level among the natural populations from 1994 to 2002/2003; it was enhanced by the artificial release of hatchery-reared charr before 1994. In order to conserve the genetic diversity of the white-spotted charr in the Lake Biwa water system, it is necessary to prevent the stocking of hatchery-reared charr in reaches where hatchery-reared charr have not previously been stocked.     

Arctic charr in Lake Myvatn: The centennial catch record in the light of recent stock estimates

In Lake Myvatn, Arctic charr (Salvelinus alpinus) is economically the most important fish species. It is fast growing fish and its size at maturity is 35–50 cm at the age of 4 to 5 years. Catch statistics that are available for the whole 20th century show considerable long-term variation with the highest catch in the 1920s. The catches after 1970 are about 40% lower than the average from 1930–1969 and the annual catches for the last decade are the lowest. Stock size during the winter fishing season fluctuated considerably between years, with average annual fishing mortality of 83.9%. The Arctic charr population has been monitored annually since 1986, using standard series of gill nets of different mesh sizes. In 1988 one to three-year-old fish were heavily reduced in numbers during the summer months. There are indications of a similar event in 1997. In both years the charr changed its main diet from Cladocera and chironomid midges to three-spined sticklebacks Gasterosteus aculeatus and snails. At the same time its condition deteriorated. The catch in the monitoring fishery in Lake Myvatn correlates with the stock size in the beginning of the following winter fishing season deduced from Leslie's method and can give prospects for the fishery in the successive fishing season. An index of abundance of young charr also correlates with the number of chironomids and cladocerans and also with ducklings that feed on the same food as the charr. Large fluctuations in the Arctic charr population in Lake Myvatn seem to be related to changes in the main food species. The catch records available from Lake Myvatn can to a large extent be used as a measure of changes of the Arctic charr population in the lake for the past century.     

Abundance, Annual Survival, and Recruitment of Unexploited and Exploited Lake Charr, Salvelinus Namaycush, Populations at the Experimental Lakes Area, Northwestern Ontario

We studied abundance, annual survival, and recruitment of nine lake charr, Salvelinus namaycush, populations at the Experimental Lakes Area, Ontario, for periods of 9–24 years. We used the Jolly–Seber mark-recapture method to evaluate abundance and annual survival in all populations, and total catches of individual year classes to evaluate recruitment. Seven populations were unexploited and unaffected by whole-lake experiments. One population was exploited prior to mark-recapture study and another was affected by experimental acidification. Abundance ranged from 8 to 24 fish ha^–1 in the unaffected populations. Annual survival ranged from 78% to 93%yr^–1 in all populations except the exploited population. This population may have been additionally affected by northern pike, Esox lucius, predation. Yearly recruitment was relatively constant in all populations and related to lake charr abundance. Lake charr abundance did not recover in populations affected by exploitation or acidification. Unexploited lake charr populations were characterized by relatively constant abundance, high annual survival, and relatively constant annual recruitment.     

Relationships among tributary length,census population size,and genetic variability of white-spotted charr, <Emphasis Type="Italic">Salvelinus leucomaenis</Emphasis>, in the Lake Biwa water system

The relationships between census population size and tributary length and between haplotype diversity of the mitochondrial DNA and census population size in ten white-spotted charr populations in the Lake Biwa water system and its adjacent basins were investigated. The census population size (number of fish with ≥100 mm in standard length) significantly increased with the tributary length. In the eastern part of the Lake Biwa water system, haplotype diversity increased with the census population size. On other hand, in the western part of the water system and adjacent basins, haplotype diversity was zero irrespective of the census population size. These results suggest that white-spotted charr populations in the eastern and western part of the Lake Biwa water system have undergone different levels of bottlenecks related to the habitat size in the postglacial warming.     

Does global warming threaten the dynamics of Arctic charr in Lake Geneva?

In France, the Arctic charr (Salvelinus alpinus) is native in only two lakes (Lakes Bourget and Geneva), in the most southerly part of its distribution area. It is a profundal morph living at depths of between 30 and 100 m in Lake Geneva. Following considerable stocking of Lake Geneva with juvenile Arctic charr and some good results during the 1980s, catches are currently declining. Several hypotheses have been proposed to explain this decrease, but the potential influence of warming of the lake in the 1990s has not been considered. This article studies the relationship between the strength of the cohorts and the temperature in Lake Geneva from 1992 to 2002, and discusses the various potential causes of the population collapse observed. The findings reveal close correlation between water temperature and the strength of the cohorts. It is concluded that the recent warming of Lake Geneva may have a significant direct or indirect impact on the Arctic charr population.     

Biomass and Production of Lake Charr During the Acidification and PH Recovery of A Small Ontario Lake

The biomass and production of lake charr, Salvelinus namaycush, was studied in Lake 223, a lake that was intentionally acidified to pH 5.0 and then allowed to gradually recover, and in Lake 224, a natural lake of neutral pH. During the first 6 years pH decreased, biomass of Lake 223 lake charr increased, primarily due to high recruitment during the initial years of acidification. Biomass then decreased the final year of acidification. Biomass of Lake 223 lake charr remained low during the subsequent 10 years of pH recovery and 5 additional years after lake pH had returned to neutral pH. Production of lake charr decreased during acidification and increased during pH recovery. Production was still at least an order of magnitude lower at the end of this study than 22 years earlier before lake pH was lowered. The continued low production was caused primarily by low abundance during the years of pH recovery and following years. Production per individual fish in Lake 223 decreased during acidification and quickly increased during pH recovery to values similar to that prior to acidification. Production, biomass, and production per individual varied little between years for lake charr in Lake 224.     

Atypical swimbladders of lake charr, Salvelinus namaycush, from Great Slave Lake, Northwest Territories, Canada

Swimbladder walls of lake charr, Salvelinus namaycush, from Great Slave Lake (GSL), Northwest Territories, Canada, were unusually thick for the species. The thinnest sections of the GSL bladders (mean = 2.44mm, range = 1.1–4.4mm) were significantly thicker (P = 0.001) than lake charr swimbladders collected from two small Province of Ontario lakes (means = 0.65 and 0.92mm), whose populations were assumed to be representative of the species. Variance in wall thickness was also greater in GSL lake charr than in charr from two small lakes (P < 0.02). Within individuals, some of the GSL bladder walls were markedly irregular in thickness, but whether these anomalies exist in situ or were artifacts of preservation remains uncertain. The bulk of the tissue in the thickest sections of the GSL swimbladders was in the tunica serosa (outer layer). The extent of the modification of the GSL swimbladders is extraordinary for northern fishes in postglacial lakes.     

Lake enrichment and the status of Windermere charr, Salvelinus alpinus (L.)

All English populations of Arctic charr, Salvelinus alpinus (L.), are found in the Lake District (northwest England). There are at least four races of charr in Windermere, the largest lake in England; the North and South basins of the lake each contain two distinct races that spawn in autumn and spring respectively. The spring spawners in both basins probably represent less than 15% of the total population in the lake.
Changes in the population density of charr in the lake are described briefly and examined in relation to the trophic status of the lake. Other factors that could possibly affect the charr population are reviewed, especially the influence of climate change.     

Size at maturity of fluvial white-spotted charr, <Emphasis Type="Italic">Salvelinus leucomaenis</Emphasis>, around the Lake Biwa water system varies with habitat size

Size at maturity of fluvial white-spotted charr, Salvelinus leucomaenis, was studied in small headwater tributaries of nine rivers around the Lake Biwa water system, Japan. Threshold size at maturity in both sexes showed significant positive relationships with water discharge, indicating that smaller threshold sizes at maturity of fluvial white-spotted charr occurred in smaller habitats. These results provide a link between size at maturity and habitat size and have important implications for the management of both habitats and white-spotted charr populations.     

Reproduction and development in the dwarf form of Arctic charr Salvelinus alpinus from Lake Davatchan, Transbaikalia

Biological characteristics in the deepwater dwarf form of Arctic charr Salvelinus alpinus from Lake Davatchan, Transbaikalia, are described. In nine from fifteen attempts of artificial insemination of eggs from a female by sperm from a male, developing eggs are obtained. Embryonic and larval development is described at the initial temperature 1.6–3.6°C and subsequent (from early gastrulation to the transition of larvae to exogenous feeding) average temperature 4.4 and 8.6°C. To take into account the data obtained previously (Pavlov et al., 1990, 1993b), the comparison of early ontogeny is conducted between the dwarf and small forms of charr from Lake Davatchan and the charr from Lake Leprindokan (two lakes are separated by the ridge 8 km in width). A substantial similarity in early ontogeny between the dwarf and small forms of charr from Lake Davatchan suggests that genes controlling development of the embryos and prelarvae until the transition of the young to exogenous feeding have not been subjected to natural selection determining a considerable differentiation between two forms and their reproductive isolation. Published in Voprosy Ikhtiologii, 2008, Vol. 48, No. 1, pp. 102–119. The article was translated by the author.     

Morphological characteristics of an introduced threespine stickleback,Gasterosteus aculeatus,from Hasse Lake,Alberta: a first occurrence in the interior plains of North America

Synopsis Threespine stickleback, Gasterosteus aculeatus, are thought to have been introduced into Hasse Lake, Alberta, where they were discovered in 1980. The population has grown in numbers since 1980 and by 1981 was more abundant than the native brook stickleback, Culaea inconstans. This population is the first to be found in the interior plains of North America, and it occurs at a higher elevation than any other Canadian population. The source of this introduced population is unknown. Significant differences among four years were found in several meristic and morphometric characters but no significant trends during 1980–1983 were found. Most characters were well within the range of other populations of the species.     

Speciation and genetic divergence of three species of charr from ancient Lake El'gygytgyn (Chukotka) and their phylogenetic relationships with other representatives of the genus Salvelinus

The diversity of phenotypically different and often reproductively isolated lacustrine forms of charrs of the genus Salvelinus represents a substantial problem for taxonomists and evolutionary biologists. Based on the analysis of variability of ten microsatellite loci and two fragments of mitochondrial DNA (control region and cyt‐b gene), the evolutionary history of three charr species from Lake El'gygytgyn was reconstructed, and phylogenetic relationships between the main representatives of the genus were revealed. Three species from Lake El'gygytgyn were strongly reproductively isolated. Long‐finned charr described previously as Salvethymus svetovidovi, an ancient endemic form in the lake, originated 3.5 Mya (95% Bayesian credible intervals: 1.7, 6.1). Placement of this species in the phylogenetic tree of Salvelinus was not determined strictly, but it should be located in the basal part of the clade Salvelinus alpinus – S. malma species complex. The origin of small‐mouth charr S. elgyticus and Boganida charr S. boganidae in Lake El'gygytgyn was related to allopatric speciation. Their ancestors were represented by two glacial lineages of Taranets charr S. alpinus taranetzi from Asia. In Lake El'gygytgyn, these lineages entered into secondary contact postglacially. A revision of the main phylogenetic groups within the Salvelinus alpinus – S. malma complex is conducted. The Boganida charrs from Lakes El'gygytgyn and Lama (Taimyr) belong to different phylogenetic groups of Arctic charr and should not be regarded as a single species S. boganidae. Using the charrs from Lakes El'gygytgyn and Lama as a case study, we show that a model of sympatric speciation, which seemed more probable based on previous empirical evidence, was rejected by other data.     

The Biology of Arctic Charr, Salvelinus Alpinus, of Gander Lake, A Large, Deep, Oligotrophic Lake in Newfoundland, Canada

Resident Arctic charr, Salvelinus alpinus, are widespread throughout the island of Newfoundland. This study examines aspects of the biology and spatial and temporal distributions of the charr of Gander Lake, the third largest in Newfoundland (surface area = 11320ha, maximum depth = 288m, mean depth = 105.4m). The deepest part of the lake is approximately 258m below sea level. The lake is well oxygenated from the surface to the bottom during all seasons. Sampling was conducted with Lundgren multiple-mesh experimental gillnets and baited hooks. There appears to be two morphs present, based on colour (dark and pale) and certain meristic characteristics. Dark charr were caught mainly in benthic nets (at depths from 1 to 100m inclusive) with only a few pelagic captures. Pale charr were caught only in benthic nets at depths between 20 and 100m inclusive. The maximum depth sampled was 196m, but there was no catch. There was a tendency for dark charr to be found in deeper, cooler water as the upper water column and inshore areas warmed during summer. There was no apparent trend in size of charr with depth sampled. Dark and pale charr both fed on benthic macroinvertebrates; sticklebacks were consumed only by dark charr and the importance of this prey item increased with size of predator. Zooplankton and surface food were not utilised by Gander Lake charr. Results of the study are compared with findings reported for other water bodies in Newfoundland and Labrador, North America, and Europe, particularly Loch Ness which has similarities in morphometry and trophic status to Gander Lake.     

Adaptations to Stochastic Environmental Variations: The Effects of Seasonal Temperatures on the Migratory Window of Svalbard Arctic Charr

The circumpolar Arctic charr, Salvelinus alpinus, is ideal for studying how environmental factors affect life history in fishes. Charr populations demonstrate a tremendous ecological plasticity and adaptations to harsh environments. Arctic charr is the only freshwater fish on Svalbard, including anadromous, resident, and landlocked stocks. Freshwater lake systems on Svalbard are characterized by very low water temperatures, long-term or even permanent ice cover, and low levels of nutrients. Food is thus limited and may lead to growth stagnation and early maturity in Arctic charr. The individual growth pattern may alternatively follow a sigmoid-shaped curve, caused by a shift to either cannibalistic or anadromous (migration to sea) behaviour. In lake systems that include migratory charr, the population may consist of a mixture of parr, postsmolt, and adult migratory individuals, as well as small-sized resident, large-sized resident (cannibals), and large formerly resident individuals transformed to anadromy. Our study in the Lake Dieset watercourse (79°N), Svalbard, demonstrates that the annual water flow in the outlet river is strongly correlated to air temperatures and provides a passage to the sea, allowing the charr access to the nutrient-rich seawater environment, during at most two months each year. During one of the years studied, the youngest and small-sized part of the sea-going stock was prevented from ascending the river and probably suffered mortality during winter. The migratory window of the Arctic charr in Lake Dieset is therefore highly variable among years and thus unpredictable. We hypothesize that in worst case scenarios (cold years, low water discharge), climatic variations may occasionally prevent charr from migrating upstream in Svalbard lake systems in late autumn, resulting in high mortality in the population.     

Identification of the maternal source of young‐of‐the‐year Arctic charr in Lake Hazen,Canada

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Takvatn Through 20 Years: Long-term Effects of an Experimental Mass Removal of Arctic Charr, Salvelinus Alpinus, From a Subarctic Lake

Between 1984 and 1989, the experimental removal of 31 tons (666000 fish) of stunted Arctic charr, Salvelinus alpinus, from Takvatn in northern Norway, had strong effects on the populations of Arctic charr, brown trout, Salmo trutta, and three-spined sticklebacks, Gasterosteus aculeatus. The littoral catch per unit effort (CPUE) of charr had decreased by 90% in 1990 and then increased to about 50% of the initial level by 1994 while the pelagic CPUE had decreased to zero. Growth in both charr and trout greatly improved when the charr density had decreased, and large fish of both species appeared in the catches. These large fish became predators on small charr in the littoral zone. The incidence of trout increased from below 1% to 15% from 1988 to 1999 after a brief peak at 30% in 1992 and 1993. The charr population attained a bimodal size distribution and did not return to the stunted state during the 10 years following the intensive fishing period. The mass removal experiment showed that it is possible to change the structure of a charr population by intensive fishing. Predation on small charr from cannibals and large trout was probably essential for maintaining the new population structure. An increase in the growth of young charr from 1995 to 1997 was related to a high consumption of Daphnia and Eurycercus. Rapid changes in the growth of charr followed the density fluctuations in sticklebacks, which show large annual variations in this system; the rapid changes in charr growth were probably caused by variations in the competition intensity for cladoceran prey between young charr and sticklebacks. Twenty years of data has provided important information, but even more time is needed to follow the long-term trends in northern lakes such as Takvatn.     

The cultural importance and international recognition of the Arctic charr Salvelinus alpinus populations of Windermere,UK

The Arctic charr Salvelinus alpinus populations of Windermere, England’s largest natural lake in the Lake District of north-west England, have been studied since the 1940s. However, the species’ cultural importance has a much longer history and these populations have been fished since 1223. As early as 1660, Arctic charr were exploited in a high-profile commercial net fishery and greatly enjoyed by locals and in a ‘potted’ form by wealthier members of UK society. However, overfishing led to substantially decreased catches and commercial netting was stopped in 1921. Local fishing for Arctic charr persists to the present as a small recreational plumb-line fishery using artificial lures. These fishing activities and resulting catches have long held great cultural interest for the local community and visiting national and international tourists. The cultural importance of the Arctic charr populations of Windermere has recently been the subject of national media interest that culminated in a 2017 documentary film highlighting environmental issues facing the Arctic charr and also celebrating the role of this iconic species in the cultural life of Windermere. In addition, international recognition of the Arctic charr populations of Windermere also contributed to the Lake District becoming a UNESCO World Heritage Site in 2017.

     

Comparative catches and food habits of dolly varden and arctic charrs,Salvelinus malma and S. alpinus,at Karluk,Alaska, in 1939–1941

Synopsis For many years the seriousness of the predations of dolly varden (Salvelinus malma) upon the eggs, alevins and juveniles of Pacific salmons (Oncorhynchus) has been a controversial subject. Over $ 300,000 was spent in Western Alaska from 1920 to 1941 for bounties on dolly varden in the belief that they were serious salmon predators. The writter undertook this study because of an intense desire to evaluate the need and justification for such a bounty system. The incidental examination of stomach contents of charrs which were captured for tagging or for tag recovery purposes from Karluk Lake and Karluk River on Kodiak Island, Alaska, and from Shelikof Strait and Uyak Bay during the summer months of 1939, 1940 and 1941 presented an excellent opportunity to pursue such a project. Stomachs were collected and contents analyzed from 1,992 arctic charr (Salvelinus alpinus) and 2,565 dolly varden charr (Salvelinus malma), taken by beach seine, gill net, fyke net, hook and line and by weir trap from Karluk Lake; from 956 dolly varden taken by seine and weir trap from the Karluk River; and from 462 dolly varden taken by beach seine and commercial salmon traps in Uyak Bay. Among the more than 5,000 charr stomachs examined, only 42 were found to contain salmon smolts, parr or alevin. Examination of 500 stomachs taken from dolly varden charr captured in the lower Karluk River at the height of the red salmon (Oncorhynchus nerka) smolt migration in May of 1939 revealed very little evidence of predation on the smolts in fresh water. There was some evidence that this relationship may have changed after the migrating charr had become adjusted to salt water, although the examination of 460 stomachs of dolly varden captured in salt water revealed nothing particularly incriminating in its role as a salmon predator. The fact that the downstream migration of the dolly varden reached its peak before that of the red salmon smolts was considered significant. Examination of the 4,500 charr stomachs at Karluk Lake in the months from April through September led to the discovery that two distinct species of charrs were present, one, the dolly varden, primarily stream-inhabiting and anadromous, and the other, the arctic charr, primarily lake-inhabiting and nonanadromous. Although red salmon eggs constituted 32% of the total volume of food ingested by both species of charr at Karluk Lake, there was considerable evidence that they were consumed in salvage or scavenger feeding, and there was practically no evidence that they were consumed as a result of predatory feeding. Only five, or one-tenth of one percent, of the thousands of charr stomachs examined at Karluk Lake contained red salmon parr or alevin.A summary of the contrasts in the feeding and migratory habits and physical characteristics of Salvelinus malma and Salvelinus alpinus led to the conclusion that alpinus was a more serious potential salmon predator than malma at Karluk Lake. In view of the fact that alpinus was seldom reached by the predator-control program whereas malma was being systematically destroyed only because it was more easily captured due to its migratory tendencies, it was recommended that the predator control program be terminated in Karluk waters.Published post mortem in shortened and slightly edited version. Reprint requests to the Editor.