Animal Diversity Web

Geographic Range

Dolly Varden range from the coastal areas of southeast Alaska across the Gulf of Alaska and the Bering Sea into the Beaufort Sea to the Mackenzie River in Canada. Specifically in North America these char range from Washington state to the Arctic coast of Canada. Outside of North America the subspecies Salvelinus malma curilus and Salvelinus malma miyabei can be found along the Pacific coast of Russia and south to Japan and Korea. This species can also thrive in inland steams or lakes around these areas. During the summer Dolly Varden make a feeding migration to the Pacific coast from fresh water steams and rivers. During this migration they typically stay close inshore but can cover several kilometers to feed and find steams to travel upward into and stay for the winter if it is a suitable stream, lake or river.

In Alaska, a northern form Salvelinus malma malma and southern form Salvelinus malma krascheninnikova exist. The two differ in number of vertebrae and number of chromosomes. Southerns have 62–65 vertebrae and 82 chromosomes; northerns have 66-70 vertebrae and 78 chromosomes. They also differ in the number of gill rakers. Northern form Dolly Varden can become much larger than the southern form, although both do make migratory sea-runs if not landlocked. Presumably isolated from one another since the last glaciation, the northern form Dolly Varden range from the north side of the Alaska peninsula northward to the Mackenzie River in Canada and in the Susitna drainage in south-central Alaska. The southern form ranges from southeast Alaska throughout the Gulf of Alaska to the south side of the Alaska Peninsula and the Aleutian and Kodiak islands. ("Dolly Varden (Salvelinus malma)", 2008; Oleinik, et al., 2001; "Southcentral Alaska wild Dolly Varden", 2006)

• Biogeographic Regions
• nearctic
  • native
• oriental
  • native
• pacific ocean
  • native

Habitat

Dolly Varden live primarily in freshwater environments, particularly streams that are often less than one meter wide in the summer. They can also live in lakes, larger rivers, brackish estuaries, marine shorelines and the open ocean occasionally. In streams, populations are much denser in areas with substantial coarse woody debris. This debris offers them protection from current and cover to ambush prey. This habitat increases their efficiency because they use less energy swimming in current as well. Dolly Varden are an anadromous species which means they migrate between fresh and salt water. During the spring, they migrate to the coasts of oceans or brackish estuaries to feed and spawn. In rivers they are typically found around half a meter in depth, and tend to stay around the bottom half of the water column in deep rivers. (Baxter, et al., 1997; Dolloff, 1986; "Dolly Varden (Salvelinus malma)", 2008)

• Habitat Regions
• temperate
• saltwater or marine
• freshwater

• Aquatic Biomes
• lakes and ponds
• rivers and streams
• coastal
• brackish water

• Average depth

  0.5 m

  1.64 ft

Physical Description

Freshwater adult Dolly Varden generally are a blueish-olive green color on their dorsal side. They have very small lightly colored spots, less than 1 mm in diameter, all over their body and especially around their dorsal fin. After a sea-run, they are typically silvery with a faint green sheen overlain with light orange spots. After a couple of weeks in freshwater, the silvery color changes into a blue, green or brown, depending on environment, with dark orange or red spots. Their average length is from 25 centimeters to 70 centimeters. Their average weight typically ranges from 0.25 to 9 kg. The largest Dolly Varden weighed in at 9.46 kilograms and was 101.6 cm long and was caught by a fly fishermen using a imitation leech lure. The northern form can weigh up to 9 kilograms. The southern form can weigh up to 5.5 kilograms, but most commonly around 1-2 kilograms.

Males change in appearance markedly as spawning season begins. Their bellies change to a red, black and or white with black operculum, bright orange to red spots and bright orange and black fins with a distinct bright snow white around the leading edge of their anal and pectoral fins. Males also develop a kype or a strongly-hooked jaw. Females change in color in the same ways as males only to a lesser degree, and they do not develop a kype.

Juveniles are 2 mm long at hatching and are silvery in appearance. Juveniles vary in color once they grow past 1-2 cm long depending on the environment they inhabit; in lakes and non-glacial streams, they are greenish-olive, and in glacial streams they are pale silver to grey. They begin to take on the appearance of the adults when they reach about 7-10 cm long. Some Dolly Varden that are land locked in small head streams less than 1 m wide are dwarfed and fully mature around 6 cm and grow no larger than 15 cm long.

Dolly Varden are char that are commonly mistaken as trout. Char are primarily darker-colored fish with lighter-colored spots, whereas trout are lighter-colored fish with darker spots. This species is also commonly mistaken as Arctic char, Salvelinus alpinus. The two species are separated genetically, but look strikingly similar and are found in similar geographic ranges. (Baxter, et al., 1997; "Dolly Varden (Salvelinus malma)", 2008; "Southcentral Alaska wild Dolly Varden", 2006)

• Other Physical Features
• bilateral symmetry

• Sexual Dimorphism
• male larger
• sexes colored or patterned differently
• male more colorful
• sexes shaped differently

• Range mass

  0.25 to 9.46 kg

  0.55 to 20.84 lb

• Range length

  25 to 101.6 cm

  9.84 to 40.00 in

Development

Dolly Varden spawn and create nests in stream gravel sediment called redd. The alevins (newly spawned char) hatch in early spring and emerge from the sediment when they are 2 mm. At this stage, they use slow-moving water to feed. Dolly Varden exhibit indeterminate growth throughout their lives. Often the juveniles will travel upstream into spring-fed areas or smaller tributaries if available. The deeper spring areas provide a habitat for over-wintering in their first year. Many juveniles will not survive icing conditions in some streams during colder winters unless they migrate into these spring areas. After two to four years, Dolly Varden normally move into larger streams or rivers further downstream where bigger prey is present. On average male and female Dolly Varden become sexually mature at age three and grow at the same rate. They continue to grow and may spend the rest of their lives in the bigger stream or river if it is productive. Many stay in the river and migrate to lakes to over-winter. Non-migratory or resident Dolly Varden typically live from eight to ten years and typically reach 26 cm long. Less commonly, some adults will migrate to the ocean if accessible from the river. Sea-run Dolly Varden usually grow to 38-56 cm and live up to 19 years. In general sea-run Dolly Varden are less common than resident. On average adults weigh from 0.5-2 kg depending on location and migratory patterns. ("Dolly Varden (Salvelinus malma)", 2008; Haas, 1988)

• Development - Life Cycle
• indeterminate growth

Reproduction

Mating occurs once a year in the fall between September and November. Some Dolly Varden may skip 1 to 2 years of spawning due to the physical taxation of spawning. Males and females both may mate with several different individuals during spawning season in order to increase chance of a successful spawn. Males compete with one another over breeding females. This competition can lead to a few males mating with the majority of females. Females primarily compete over the most suitable spots in which to create their redds. Females find shallow, slow moving water areas with a gravel or pebble sediment. They use their tail to move the rocks and create a small indention, roughly 15 cm in diameter. Females will stay on their nests while males come to the redds to mate. In the act of spawning, a single male swims around the nest and rubs against the female occasionally. The male deposits sperm over the redd and fertilizes the eggs the female has laid there. (; Jonsson, et al., 1984)

• Mating System
• polygynandrous (promiscuous)

Spawning happens typically once a year between September and November. Dolly Varden return to their steam of origin to spawn. Spawning is difficult physically on the fish as males and females do not eat while they are on their redds. Females often compete among each other directly for locations and males physically fight for females to breed or to keep other males away from the redd. A individual Dolly Varden typically will not spawn more than three times in their lifespan, male or female, especially male. Often they will skip 1 to 2 years between spawning and spawn depending on the food availability that year, especially in the northern form. Females lay from 600 to 6,000 eggs (2,500 to 10,000 in the northern form) in the redd depending on her size. There is no gestation period as Dolly Varden exhibit external fertilization of the eggs.

The incubation period is 4-5 months but can take 3-6 months, depending on water temperature. Birth mass is 0.01 to 0.1 g, averaging 0.05 g. The alevins will also spend 1 to 2 months in the stream-bed gravel after hatching and consume nutrients from their embryonic egg sac during this time and emerge from the stream-bed gravel when this resource has been exhausted. They come out of the stream-bed typically from March to May but can come out as late as June in the northern form.

The alevins spend 2 to 4 years in the stream in which they hatched before venturing further into larger rivers or the ocean in which they grow rapidly due to the increased availability of prey. Both sexes reach sexual maturity in about 3 years (range 2-4 years). ("Dolly Varden (Salvelinus malma)", 2008; Jonsson, et al., 1984; "Southcentral Alaska wild Dolly Varden", 2006; "The Coastal Forests and Mountains Ecoregion of Southeastern Alaska and the Tongass National Forest", 2007)

• Key Reproductive Features
• iteroparous
• seasonal breeding
• sexual
• fertilization
  • external
• broadcast (group) spawning
• oviparous

• Breeding interval

  Once a year, may skip 1-2 years between spawns

• Breeding season

  September-November

• Range number of offspring

  600 to 10,000

• Range time to hatching

  3 to 6 months

• Average time to hatching

  4-5 months

• Range time to independence

  0 to 0 minutes

• Range age at sexual or reproductive maturity (female)

  2 to 4 years

• Range age at sexual or reproductive maturity (male)

  2 to 4 years

Females will build the redds and once the nest is fertilized by the male, the parental investment becomes non-existent. The female will force any exposed eggs into crevices in the stream bed and cover the redd with gravel with her tail to protect her eggs. After this, she leaves and finds food and protection to recuperate from the spawn. Males will also find food and protection immediately post-spawning, as they are typically weak due to physical fighting and several days or weeks spent guarding the redd prior to spawning. Once the eggs have been fertilized, both parents leave and parental investment ceases. ("Dolly Varden (Salvelinus malma)", 2008; "Southcentral Alaska wild Dolly Varden", 2006; "Southeast Alaska steelhead and Dolly Varden management", 2008)

• Parental Investment
• male parental care
• female parental care
• pre-fertilization
  • provisioning
  • protecting
    • male
    • female

Lifespan/Longevity

Sea-run Dolly Varden have a longer lifespan than resident Dolly Varden. Sea-run Dolly Varden typically live between 10-20 years, the oldest Dolly Varden studied was 22 years old. Freshwater Dolly Varden usually have a maximum lifespan of 10 years and on average live from 6-8 years. Dolly Varden are not stocked and therefore have not been studied in captivity. (Dolloff, 1993; "Dolly Varden (Salvelinus malma)", 2008)

• Range lifespan
  Status: wild

  22 (high) years

• Typical lifespan
  Status: wild

  6 to 20 years

• Average lifespan
  Status: wild

  10 years

Behavior

Dolly Varden that are landlocked in small streams, roughly less than 1 meter wide without easy access to the sea, are dwarfed in this type of habitat. Some have been observed to stay in a freshwater environment their whole lives if the habitat is productive and suitable conditions throughout the year instead of making a feeding sea-run. They can grow up to 60 centimeters in these habitats although sea-run Dolly Varden typically get larger.

During spawning Dolly Varden behave much differently than the rest of the year. Their main focus shifts from eating and growing to breeding. This makes them forage much less and spend from 1-2 weeks looking for mates or creating redds. Typically after spawning Dolly Varden lose much of their weight due to eating much less. Post-spawn is a rebound and they forage much more to recover from the lack of eating.

Dolly Varden spend most of their daylight hours sitting and waiting behind objects that disrupt current in a stream, such as rocks or logs in the stream. This is a way to cope with the current and forage efficiently. Dolly Varden wait in these areas for prey moving down river in the fast-moving water and will strike prey and then return to their cover.

Although they forage during the daytime, Pavlov et al. (2015) reported that the young-of-the-year migration for the Russian subspecies is purely nocturnal. ("Dolly Varden (Salvelinus malma)", 2008; Pavlov, et al., 2015)

• Key Behaviors
• natatorial
• diurnal
• nocturnal
• motile
• migratory
• sedentary
• social

• Range territory size

  1 to 2 m^2

Home Range

Dolly Varden have a home range in slow moving water like eddies or pools on the stream, and typically stay in 1-2 square meters. Typically more than 2 Dolly Varden will live in a single eddie, pool, or slow-moving water spot. On a daily basis Dolly Varden will not venture more than 100 m from their home, unless migrating. ("Dolly Varden (Salvelinus malma)", 2008)

Communication and Perception

Like most fish species, Dolly Varden use lateral lines to detect vibration or disruptions in the water to sense prey, which are used primarily after rain when the water is too murky or muddy to use visual cues. Most of the time they use vision to see and target prey in clear water. Schutz (1967) found that Dolly Varden could see within a radius of 5 meters in clear water. Extremely turbid water can make it more difficult for them to see and sense prey. However, once prey have been detected, water turbidity does not disrupt their capture. Dolly Varden use tactile senses when spawning and to detect current. They also use chemical senses like smell and taste when hunting or eating. (Schutz, 1967)

• Communication Channels
• visual
• tactile

• Perception Channels
• visual
• tactile
• vibrations
• chemical

Food Habits

Dolly Varden are opportunistic feeders whose feeding behavior and diet depends primarily on their habitat and season. In lakes, they have been observed eating plankton when there are no smaller fish on which to prey. Adults in rivers and streams typically hunt near the faster-moving water and eat small fish or insects. Often when they are found in eddies and deeper pools, they consume crustaceans, salmonids' eggs and other small fish. They will also feed on the surface of the water, striking winged insects that fall into the water. Fly fishermen tie "flies" to mimic winged insects as lures often when fishing for Dolly Varden. They will also consume any larvae of mayflies and midges.

Northern form adult Dolly Varden make feeding sea-runs every 9 or 20 months, depending on food availability, where they may follow pink salmon Oncorhynchus gorbuscha into rivers to feed on their eggs. They primarily scavenge these eggs rather than attack them as direct predators. They mainly consume the eggs and fry that drift away downstream that would most likely not survive.

Juveniles hatch in slow-moving fresh water and initially consume the nutrients from their soft embryotic egg shell. Once they have exhausted that resource, they feed in slow-moving water, eating worms, mayfly larvae and juvenile insects that inhabit the sediment often under stones and in mud. Often times juveniles forage very close to the bank rather than in the middle of the stream or river. ("Dolly Varden (Salvelinus malma)", 2008; Hume and Northcote, 1985; Langeland, et al., 1991; "Southcentral Alaska wild Dolly Varden", 2006)

• Primary Diet
• carnivore
  • piscivore
  • eats eggs
  • insectivore
  • eats non-insect arthropods
  • molluscivore
  • vermivore
  • scavenger

• Animal Foods
• fish
• eggs
• insects
• mollusks
• terrestrial worms
• aquatic or marine worms
• aquatic crustaceans
• other marine invertebrates

Predation

Dolly Varden are prey of northern river otters Lontra canadensis. In southeastern Alaska, during the late spring, Dolloff (1993) reported northern river otters eating hundreds of fish in a day and thousands over a six-week period. Many of these fish are Dolly Varden. Other fish in streams or rivers also can prey on juvenile Dolly Varden.

They can also be prey of some birds such as white-tailed eagles Haliaeetus albicilla to the subspecies of Dolly Varden that resides in Russia and Japan. Adult Dolly Varden avoid predation due to their dark greenish-blue camouflaged dorsal side, which matches the sediment or rocks from above. (Dolloff, 1993)

• Anti-predator Adaptations
• cryptic

• Known Predators

  • northern river otters (Lontra canadensis)
  • white-tailed eagles (Haliaeetus albicilla)

Ecosystem Roles

Dolly Varden are the apex predator of their freshwater habitats. While they are in the saltwater however they are vulnerable to bigger predators and fall into a larger food web than in freshwater. Dolly Varden are also vulnerable to many parasites. The protozoan parasites include Trichodina truttae and Tripartiella californica. The trematode parasites include Aonurus. Brachyphallus crenatus, Buphalopsis gracilescens, Bucephalopsis ozarkii, Crepidostomum cooperi, Crepidostomim farionis, Derogenes varicus, Discocotyle salmonis, Genarches mulleri, Hemiurus levinseni, Lecithaster gibbosus, Lecithaster salmonis, Prosorhynchus crucibulum, Tetraconchus alaskensis and Tetraconchus borealis. Cestodes include Cyathocephalus truncates, Eubothrium crassum, Eubothrium salvelini, Nybeliniasurmincola, Pelichnibothrium speciosum, Phyllobothrium, Proteocephalus salmonidicola and Scolex pleuronectis. Nematode parasites include Ascarophis malmae, Bulbodacnitis globose, Contracaecum aduncum, Contracaecum, Cucullanus laevis, Cystidicola farionis, Dacnitis truttae, Eustrongylides, Hepaticola bakeri, Metabronema canadense, Metabronema salvelini, Philonema oncorhynchi, Rhabdochona amago and Rhabdochona salvelini. Acanthocephalan parasites include Echinorhynchus coregoni, Echinorhynchus leidyi, Neoechinorhynchus rutill, Neoechinorhynchus and Rhadinorhynchoides mijagawai. Crustacean parasites include Lepeophtheirus salmonis, Salmincola bicauliculata, Salmincola edwardsii, Salmincola falculata, Salmincola gibber, Salmincola siscowet and Salmincola smirnova. (Hoffman, 1967)

Economic Importance for Humans: Positive

Dolly Varden are one of Alaska's most sought-after game fish. There are massive tourism industries for recreational fishing in Alaska. Dolly Varden add to this industry tremendously by attracting anglers from around the world to come to Alaska to catch them. Sea-run Dolly Varden are harvested every year on their migration to the sea for food by some locals. In the mid 1900's, one record states before Alaska was a state that 86,182 kg were harvested in a single year. However since the 1960's the harvest of Dolly Varden has dropped drastically in order to preserve their population and fuel tourism. ("Dolly Varden (Salvelinus malma)", 2008; "Southcentral Alaska wild Dolly Varden", 2006; "Southeast Alaska steelhead and Dolly Varden management", 2008; "The Coastal Forests and Mountains Ecoregion of Southeastern Alaska and the Tongass National Forest", 2007)

• Positive Impacts
• food
• ecotourism

Economic Importance for Humans: Negative

There are no negative economic impacts of Dolly Varden char on humans.

Conservation Status

The IUCN Red List and CITES have not evaluated Dolly Varden's conservation status. According the State of Michigan, Dolly Varden have no special status. On the US Federal list, Dolly Varden are marked as "Proposed Similarity of Appearance (Threatened)" meaning they are similar in appearance to bull trout Salvelinus confluentus that are threatened.

Over-fishing can cause serious damage to the adult population and will consequently harm the spawn and future generations of Dolly Varden. No serious natural threats are known to harm Dolly Varden. Catch-and-release is a fishing practice that is strongly encouraged by Alaska's government and strict laws have been in place to prevent over-fishing since 1959. Alaska is the only state to have explicit conservation about fishing in its constitution. They operate on a strict "sustainable yield" harvest of wildlife including Dolly Varden. These laws are enforced by the Alaska Department of Fish and Game. The Alaska Department of Fish and Game along with a variety of federal land management entities are responsible for managing and conserving Alaska's Dolly Varden. ("Dolly Varden (Salvelinus malma)", 2008; "Southcentral Alaska wild Dolly Varden", 2006)

• IUCN Red List

  Not Evaluated

• US Federal List

  Threatened

• CITES

  No special status

• State of Michigan List

  No special status

Other Comments

Dolly Varden are classified as char, although commonly referred to as trout. One distinguishing factor between trout and char is their difference in the spots on their dorsal side. Char have light spots such as white or yellow on a dark body while trout have dark spots (brown to black) on a light body. ("Dolly Varden (Salvelinus malma)", 2008)

Contributors

Will Mann (author), Radford University, Layne DiBuono (editor), Radford University, Lindsey Lee (editor), Radford University, Kioshi Lettsome (editor), Radford University, Karen Powers (editor), Radford University, Tanya Dewey (editor), University of Michigan-Ann Arbor.

References

Alaska Department of Fish and Game. Dolly Varden (Salvelinus malma). None. Alaska Department of Fish and Game: Wildlife Notebook Series. 2008. Accessed September 06, 2018 at http://www.adfg.alaska.gov/static/education/wns/dolly_varden.pdf.

Alaska Department of Fish and Game. Southcentral Alaska wild Dolly Varden. None. Anchorage, Alaska: Southcentral Region, Alaska Department of Fish and Game, Division of Sport Fish. 2006. Accessed September 06, 2018 at https://web.archive.org/web/20140201193236/http:/www.planitalaska.com/files/dolly.pdf.

Division of Sport and Commercial Fisheries. Southeast Alaska steelhead and Dolly Varden management. 8-21. Anchorage, Alaska: Alaska Department of Fish and Game. 2008. Accessed September 06, 2018 at http://www.sf.adfg.state.ak.us/FedAidpdfs/sp08-21.pdf.

The Nature Conservancy. The Coastal Forests and Mountains Ecoregion of Southeastern Alaska and the Tongass National Forest. None. Anchorage, Alaska: The Nature Conservancy. 2007. Accessed September 11, 2018 at https://www.conservationgateway.org/ConservationByGeography/NorthAmerica/UnitedStates/alaska/seak/era/cfm/Documents/1_Introduction.pdf.

Armstrong, R., M. Hermans. 2006. Dolly Varden (Salvelinus malma). Pp. 1-5 in Southeast Alaska Conservation Assessment. Ketchikan, AK: The Nature Conservancy. Accessed September 30, 2018 at https://www.conservationgateway.org/ConservationByGeography/NorthAmerica/UnitedStates/alaska/seak/era/cfm/Documents/8.8_DollyVarden.pdf.

Baxter, J., E. Taylor, R. Devlin, J. Hagen, D. McPhail. 1997. Evidence for natural hybridization between Dolly Varden (Salvelinus malma) and bull trout (Salvelinus confluentus) in a northcentral British Columbia watershed. Canadian Journal of Fisheries and Aquatic Sciences, 54/2: 421-429.

DeCicco, F. 2005. Dolly Varden: Beautiful and misunderstood Dolly Varden's reputation as varmint undeserved. Alaska Fish and Wildlife News, May: 1.

Dolloff, A. 1986. Effects of stream cleaning on juvenile coho salmon and Dolly Varden in southeast Alaska. Transactions of the American Fisheries Society, 115/5: 743-755.

Dolloff, A. 1993. Predation by river otters (Lutra canadensis) on juvenile coho salmon (Oncorhynchus kisutch) and Dolly Varden (Salvelinus malma) in southeast Alaska. Canadian Journal of Fisheries and Aquatic Sciences, 50/2: 312-315.

Haas, G. 1988. The systematics, zoogeography and evolution of Dolly Varden and bull trout in British Columbia. Vancouver, British Columbia: The University of British Columbia.

Henderson, M., T. Northcote. 2011. Visual prey detection and foraging in sympatric cutthroat trout (Salmo clarki clarki) and Dolly Varden (Salvelinus malma). Canadian Journal of Fisheries and Aquatic Sciences, 42/4: 785-790.

Hoffman, G. 1967. Parasites of North American Freshwater Fishes. Los Angeles, CA: University of California Press.

Hume, M., T. Northcote. 1985. Intial changes in use of space and food by experimentally segregated populations of Dolly Varden (Salvelinus malma) and cutthroat trout (Salmo clarki). Canadian Journal of Fisheries and Aquatic Sciences, 45/1: 101-109.

Jonsson, B., K. Hindar, T. Northcote. 1984. Optimal age at sexual maturity of sympatric and experimentally allopatric cutthroat trout and Dolly Varden charr. Oecologia, 61/3: 319-325.

Langeland, A., J. L'abee-lund, B. Jonsson, N. Jonsson. 1991. Resource partitioning and niche shift in arctic charr Salvelinus alpinus and brown trout Salmo trutta. Journal of Animal Ecology, 60/3: 895-912.

Oleinik, A., L. Skurikhina, V. Brykov, J. Wenburg. 2005. Differentiation of Dolly Varden char Salvelinus malma from Asia and North America inferred from PCR–RFLP analysis of mitochondrial DNA. Genetika, 41/5: 626-634.

Oleinik, A., L. Skurikhina, V. Brykov, S. Frolov, I. Chereshnev. 2001. Divergence of mitochondrial DNA of two Dolly Varden trout (Salvelinus malma; Salmonidae: Pisces) subspecies. Doklady Biological Sciences, 376/6: 99-101.

Pavlov, D., E. Kirillova, P. Kirillov. 2015. Migration of young of the year of Dolly Varden char Salvelinus malma (Walbaum) from the spawning tributary in the basin of Lake Kuril’skoye (Southern Kamchatka). Inland Water Biology, 8/1: 65-74.

Reist, J., G. Low, J. Johnson, D. McDowell. 2002. Range extension of bull trout, Salvelinus confluentus, to the central Northwest Territories, with notes on identification and distribution of Dolly Varden, Salvelinus malma, in the western Canadian Arctic. Arctic, 55/1: 70-76.

Schutz, D. 1967. An Experiemental Study of Feeding Behavior and Interaction of Coastal Cutthroat Trout (Salmo clarki clarki) and Dolly Varden (Salvelinus malma) (Master's Thesis). British Columbia, Canada: University of British Columbia.

Takami, T., F. Kitano, S. Nakano. 1997. High water temperature influences on foraging responses and thermal deaths of Dolly Varden (Salvelinus malma) and white-spotted charr (S. leucomaenis) in a laboratory. Fisheries Science, 63/1: 6-8.

Taylor, E., E. Lowery, A. Lilliestrale, A. Elzs, T. Quinn. 2008. Genetic analysis of sympatric char populations in western Alaska: Arctic char (Salvelinus alpinus) and Dolly Varden (Salvelinus malma) are not two sides of the same coin. Journal of Evolutionary Biology, 21/6: 1609-1625.

Taylor, E., Z. Redenbach, A. Costello, S. Pollard, C. Pacas. 2001. Nested analysis of genetic diversity in northwestern North American char, Dolly Varden (Salvelinus malma) and bull trout (Salvelinus confluentus). Canadian Journal of Fisheries and Aquatic Sciences, 58/2: 406-420.