Species Accounts
Salmonidae - Trouts and Salmons - Truites et Saumons
The salmons, trouts, charrs, graylings, whitefishes, ciscoes and their relatives are found in the cool and cold, fresh and salt waters of the Northern Hemisphere. There are about 70 species with 37 reported from Canada, and 9 in the NCR. Salmons are medium to large sized fishes up to about 1.5 m.
These fishes are characterised by having an adipose fin, 11-222 pyloric caeca, the swimbladder is connected to the gut by a duct, the gill membranes are free of the isthmus and extend far forward, the last 3 vertebrae are turned up in the tail skeleton, the eye muscles pass through a deep posterior myodome and attach to trunk muscles and there are 7-20 branchiostegal rays. Scales are cycloid, confined to the body and can be minute (and hard to see at a superficial examination) or moderately large (and fairly obvious at a glance). Pelvic, and sometimes pectoral, axillary scales are present. Pelvic fins are abdominal in position. There are no fin spines and the dorsal fin is at mid-body. The lateral line is obvious. Breeding tubercles develop in the spawning season in some species. Many have strong teeth on both jaws, the tongue and the roof of the mouth. These fishes are tetraploids.
The limits and relationships of species within the family have long been in contention. A number of species exist which have not been named while others have a name but contain more than a single species. The latter are often referred to as a species "complex" since it is difficult to describe the complex observed variation in simple terms of several, named species. Even the correct scientific name to apply to a species can be in dispute or not open to easy resolution. The familiar Rainbow Trout, long known by the scientific name of Salmo gairdneri, is now generally recognised as Oncorhynchus mykiss, a relative of the Pacific salmons rather than salmon and trouts of Europe. The word "salmon" is from the Latin for Atlantic Salmon. Young salmons are difficult to identify as their appearance changes with growth and adults can show colour and other character variations related to locality and season.
The salmon family is extremely important for the sport and commercial fisheries it supports. The commercial salmon landings in British Columbia for 1988 weighed 88,000 tonnes worth $312 million. When combined with the fish processing industry the salmon were worth probably over $1 billion and in addition there is an important sport fishery and farming operations. Many thousands of books and scientific papers have been written about these fishes. Members of the family have been widely introduced elsewhere in the world in cool waters, including the Southern Hemisphere. Certain salmonids are entirely resident in fresh water but many are anadromous, spending their life at sea but running up rivers to spawn and die. Migrations can be over thousands of kilometres and adults return to their stream of birth, a remarkable example of accurate navigation and ability to cross obstacles like falls by jumping. The young develop for several years in fresh water. Young salmons, trouts and charrs are known as fry or alevins, when a few centimetres long they have dark blotches (parr marks) along the flank and are called parr, as they run to the sea they become silvery and are called smolt, an adult male returning early from the sea to freshwater is a grilse, and in the Atlantic Salmon, which can return to spawn several times, the spent fish returning to the sea are called kelt.
A decline in the abundance of Lake Trout, Brook Trout, Cisco and Lake Whitefish in Gatineau Park over the previous hundred years was noted by Rubec (1972; 1975). The causes are overexploitation by commercial fishermen, poaching and angling. Low oxygen levels in the hypolimnia of most Gatineau Park lakes is also major factor limiting salmonids, perhaps resulting from deforestation allowing more nutrients into the lakes coupled with increased human presence and associated wastes.
Bergeron and Brousseau (1982) and Bernatchez and Giroux (2000) map Prosopium cylindraceum (Pennant, 1784) within the NCR but this is not confirmed by specimens.
Cisco / Cisco de lac
Coregonus artedi Lesueur, 1818
Taxonomy
Other common names include Lake Cisco, Lake Herring, Tullibee, Freshwater Herring, Blueback, Sand Herring, Shallowwater Cisco, Grayback Tullibee, Bear Lake Herring, Blueback Tullibee, Herring Salmon, Hareng de lac, Arnaqsleq, Kapisilik and Kaviselik. This species shows considerable variation over its wide range and some variants have been given scientific names, 24 subspecies being described in 1931. The confusion of genetic identity through man-induced changes and extinction of some forms and lack of study makes a resolution of these names difficult. In some lakes there are two forms, a small and a large one with different biology and anatomy. Cisco have been referred to as a Coregonus artedi "complex" which indicates that there is not a single, clearly defined species but a complex of forms. There is also a spring spawning population in Lac des Écorces, to the north of the NCR in the Lièvre River basin which, being reproductively isolated from the normal fall spawning population, can be regarded as a distinct species (see references by M. Hénault and R. Fortin, and Pariseau et al. (1983)). Hybrids with Lake Whitefish are reported for Canada, further confusing some identifications.
Key Characters
This member of the Salmon Family is distinguished from its relatives by having large, silvery scales (94 or less), no parr marks in young, teeth absent from the jaws and roof of the mouth, forked tail fin, 2 flaps of skin between the nostrils, the upper and lower jaws are about equal, giving the front of the head a pointed appearance (and the mouth is not under the snout), and gill rakers 38-64, usually 46-50.
Description
The lower jaw is usually about the same length as the upper jaw which extends back to the anterior eye margin, there is a knob at the lower jaw tip, the premaxilla bone of the upper jaw curves backwards, the body is deepest at the middle and is elongate and rounded in cross section. Dorsal fin rays 9-15, anal rays 10-15, pectoral rays 14-18 and pelvic rays 8-12. Lateral line scales 63-94. Pyloric caeca number about 100. Males have one nuptial tubercle in the centre or near the edge of each scale.
Colour
Overall colour is silvery with pink to purple iridescence. The back may be grey, light brown, bluish, greenish or almost black and the belly is white. The dorsal and caudal fins may be darkly pigmented on the distal half or on the margin. Pelvic and anal fins are milky. The pelvic fins have dark tips. Fins are generally clear with limited pigmentation.
Size
Reaches 57.2 cm total length and 3.63 kg. The Ontario record as of the year 2000 weighed 2.0 kg.
Distribution
Found from eastern Québec, including Ungava Bay and southern Hudson Bay west to northern Alberta and much of the N.W.T., north to Great Slave Lake and the lower Mackenzie River. Also in U.S. states adjacent to the Great Lakes. Specimens were collected from Lac LaPêche in 2003 and are deposited in the Canadian Museum of Nature. A limited number of cisco were introduced to Meach Lake in the late 1960s (Rubec, 1972). Rubec (1972) has it in lakes Lapêche, Philippe, Mousseau and Meach in the late 1800s, absent today, although occasional fish are caught or found dead in Lac Philippe (e.g. in 2003 by A. Martel).
Origin
This species probably entered the NCR from a Mississippian refugium (Turgeon and Bernatchez, 2001) or possibly from a Mississippian or Atlantic refugium (Mandrak and Crossman, 1992).
Habitat
This is a widespread freshwater, lake species also known from some large rivers. Cisco are pelagic in lake midwaters, at 18-53 m in Lake Superior for example, with movements into deeper, cooler water in summer and into shallower waters in fall. There are reports of daily vertical migrations with cisco coming to the surface at night under winter ice to feed on crustaceans. It is rarely found at temperatures above 18°C and its preferred temperature is 13-18°C. Cisco are tolerant of some turbidity.
Age and Growth
Life span is 31 years of age and maturity is attained at 1-6 years depending on locality, later in the north.
Food
Food is mainly plankton and large crustaceans such as opossum shrimps and amphipods but can include aquatic insects and molluscs in shallow waters or terrestrial insects taken at the surface. Eggs of their own and other species are eaten, as well as fry and minnows. A variety of other fishes eat Cisco including Lake Trout, Northern Pike, Yellow Perch, Walleye and Burbot. Lake Trout depend on this species for growth as cisco are found in the deep water of the summer habitat of trout; without cisco, trout feed on small crustaceans and do not reach a large size. Strongly swimming prey is taken by a dart and suck action. Gulping is also used where the fish open and close the mouth 2-3 times per second, taking more than 1 prey at a gulp. Even buried prey can be taken.
Reproduction
Spawning takes place from September to December, earliest in the north, when large schools form, usually at 6°C or less, optimally at 3-4°C. The spawning ground is often gravel shallows at 1-3 m but can occur pelagically at 9-42 m over deep water. Spawning may occur under a thin ice cover and is therefore difficult o observe. Males arrive on the spawning grounds first, 2-5 days before females. As many as 12 males will follow a single female but at spawning time the female is usually accompanied by 2 males. She descends to 15-20 cm above the bottom, leading the males whose heads are level with her anus. Eggs are shed, mostly at night, fall to the bottom and are abandoned. The eggs are slightly adhesive and become attached to the bottom. Ripe females produce up to 29,000, 2.1 mm diameter eggs. These eggs hatch the following spring at ice break-up.
Importance
This cisco was once very abundant and of great commercial importance with as much as 21.8 million kg being taken from Lake Erie in 1918. Localised fisheries still catch this species for food, for fur farms and for dogs in the north. It is sold fresh, frozen or smoked and is very good eating. The total Canadian catch in 1988 was 1400 tonnes. Anglers may catch it on live bait including minnows, artificial lures and flies, and by ice fishing and spearing. Ice fishing is popular in the Outaouais. This species is used as bait for Lake Trout.
Cisco populations are declining. In addition to overfishing and other factors outlined under ciscoes, this species is sensitive to enrichment of lakes since it thrives in infertile waters. Enriched or eutrophic waters have depleted oxygen levels in summer in the deep, cool waters favoured by cisco. The cisco is forced into surface waters where temperatures are too high and summerkills are common.
A fossil of Quaternary age (ca. 10,000 years ago), possibly this species, has been found in a clay nodule from Besserer's Springs (near Green Creek at Hiawatha Park in the NCR) (McAllister et al., 1981; Harington, 1983; McAllister et al., 1987).
Lake Whitefish / Grand corégone
Coregonus clupeaformis (Mitchill, 1818)
Taxonomy
Other common names include Common Whitefish, Sault Whitefish, Eastern Whitefish, Great Lakes Whitefish, Inland Whitefish, Gizzard Fish, Lake Herring, Labrador Whitefish, Sead, Humpback, Buffalo Back, Whitebait, Poisson blanc, Pointu, Pi-kok-tok, Jikuktok, Anahik, Kapihilik, Pikuktuuq, Kakiviatktok, Kavisilik, Anâdlerk, Kakiviartût, Keki-yuak-tuk, Anadleq and Qelaluqaq. This species has been referred to as the Coregonus clupeaformis "complex" because of its wide range in variation, particularly in northwestern North America, which may encompass more than one species. Some lakes, for example in Algonquin Park, Ontario, contain both dwarf and normal forms which differ in biology but not in anatomy, apart from size. In Lake Como, Ontario dwarfs are 170-179 mm and normals 280-289 mm fork length. In the Yukon several lakes have 2 forms of whitefish, a benthic form with low gill raker counts feeding on the bottom and a pelagic form with higher gill raker counts feeding on zooplankton throughout the water column. High raker fish have shorter life spans and mature earlier: they are the more unique member of the 2 forms. Samples from 5 Ontario lakes showed differences in gill raker numbers, pyloric caeca numbers, in size of the tail, dorsal fin and eye, and in biochemistry, evidence of genetic separation since they spawn at the same place and time. Evidence from mitochondrial DNA studies shows that different genetic structures correlate with the Pleistocene glaciations. Hybrids between Lake Whitefish and Cisco exist in the Great Lakes and elsewhere and are known as "mule whitefish". They are bright green in colour and intermediate in anatomical characters.
Key Characters
This member of the Salmon Family is distinguished from its relatives by having large, silvery scales (97 or less), no parr marks in young, teeth absent from the jaws and roof of the mouth, forked tail fin, 2 flaps of skin between the nostrils, the mouth is under the snout, and gill rakers 19-37.
Description
Dorsal fin rays 10-13, anal rays 10-14, pectoral rays 14-17 and pelvic rays 10-12. Lateral line scales 70-97. Pyloric caeca number 140-222. The premaxilla bone of the upper jaw curves backward making the snout rounded and the adjacent maxilla bone is twice or more longer than wide. Nuptial tubercles are found on 3 or more scale rows above the lateral line and 6 rows below, and sparsely on the head. Each scale has 1 central tubercle with smaller tubercles on each or either side on some scales. Large fish develop a hump on the back, particularly in northwestern Canada. Lake whitefish produce more mucus than Ciscoes and feel slipperier.
Colour
Overall colour is silvery with a light to dark brown, greenish or black back and a silvery-white belly. Scales are outlined with pigment in northwestern Canada. Flanks may have a bluish-tinge. Fins vary from clear in the south to black-tipped in the north but may be dusky. Lower fins generally clear but sometimes with dark spots. Faber (1985a) illustrates a larva.
Size
Reaches 91 cm total length and 19.05 kg. The Ontario record as of the year 2000 weighed 6.7 kg.
Distribution
Found from New Brunswick and some Nova Scotia lakes, and Labrador west to British Columbia and throughout the Yukon and N.W.T. including Victoria Island. Also introduced into Newfoundland and parts of British Columbia and Alberta. Also in Alaska and U.S. drainages of the Great Lakes and St. Lawrence River basin.It is rare in the NCR. Small (1883) reported it in many lakes of the Gatineau and Lièvre districts but these may have been north of the NCR limits. Rubec (1975) has it in lakes Meach and Mousseau in the late 1800s, surviving today only in Lake Mousseau. Lake Whitefish have been propagated in Ottawa area hatcheries as early as 1890 (Lasenby et al. 2001).
Origin
This species entered the NCR from a Mississippian refugium or possibly from an Atlantic, Beringian or Missourian refungium (Mandrak and Crossman, 1992).
Habitat
Lake Whitefish are found, appropriately enough, in lakes but are also in large rivers such as the Lac Deschênes section of the Ottawa River. Generally they are found close to the bottom but may be pelagic. They prefer large, cold lakes, depths >12 m for non-spawning periods and 6-14 m for spawning, temperatures of 11.9-17.0°C (preferred temperature is 12.7°C), and less than 6°C for spawning, substrates of gravel, stones and sand for spawning, and a pH greater than 5.4. They can tolerate high levels of total dissolved solids (>3000 mg L-1). Their depth range depends on temperature. In southern Canada they retreat to cooler, deeper waters in summer, down to at least 128 m but in northern Canada summer temperatures are probably cool enough in surface waters.
Age and Growth
Females live longer and mature later than males. Life span is at least 30, perhaps over 50, years. Maturity is attained as late as 8 years in Great Slave and Great Bear lakes but southern populations mature earlier, as early as age 2 for males and 3 for females. Age at first maturity declines with exploitation as growth rate increases. In Lake Winnipeg, age-groups 5, 6 and 7 accounted for 81% of the catch from 1944 to 1948 but by 1969 88% of the catch was age-groups 3 and 4.
Food
Food is various bottom invertebrates, small fishes and fish eggs although some may feed pelagically on plankton or take terrestrial insects fallen on the water surface. Fishes eaten include Johnny Darters, Spottail Minnows, Ninespine Sticklebacks, Rainbow Smelt, Gaspereau and others. Variation in gill raker number and length in any population indicates the main feeding mode. Lake Whitefish are eaten by Lake Trout, Northern Pike, Muskellunge, Walleye, Burbot, Yellow Perch and others. They eat their own eggs.
Reproduction
Spawning takes place over 2-6 weeks from September to January, earlier in the north than the south, when water temperatures fall below 8°C. Some spawning may occur under ice. Northern populations may only spawn at 2-3 year intervals. The fish enter shallow waters, less than 8 m deep, and release eggs over hard, sandy, silty or weedy bottoms. Some fish spawn in deep water however. The spawning process often takes place at night with much splashing and jumping out of the water. Pairs of fish or 1 female and 2 males rise to the surface and shed eggs and sperm. Yellowish eggs are about 2.6 mm in diameter when shed and may exceed 415,000 per female or 27,460 eggs/kg body weight. The eggs hatch in the following spring, usually April or May, and receive no care from the parents. Larvae are 10.5-11.5 mm long at hatching and live in the top metre of lakes. Post-spawning adults move to overwintering sites, such as the delta in the Mackenzie River. Stocks of this whitefish in different lakes vary considerably in diet, growth rate, movement patterns, fecundity, and egg and larval size.
Importance
This is a very important commercial species in Canadian freshwaters although numbers are declining through overfishing, pollution and the depredations of Sea Lampreys. An annual catch in the Great Lakes has been as high as about 8 million kg, caught with gill nets and trapnets. In western Canada gillnets are set through holes in the ice with a "jigger" which bounces the end of the net from an entry hole to another hole a net's length away. Along with Lake Trout, this species is the principal recreational and commercial fish in northern Canada. The total Canadian catch in 1988 was 9600 tonnes worth about $12 million. Whitefish are marketed dressed, fresh or frozen and in such categories as "jumbo", "large" and "medium". They may also be canned and the eggs made into "golden" caviar. They are considered to be better eating than Lake Trout.
Dymond (1939) records catches from 1881 onward in the general vicinity of the NCR but trends cannot be determined as fisheries data is recorded from different areas at different dates. For example, in 1901 the catch was 38,950 lbs (17,683 kg) in the Ottawa River and tributaries, the highest recorded. The Gatineau lakes were the mainstay of the commercial fishery. They also provided winter food for local people, taken on the spawning run in November, while in the summer they were taken by commercial fishermen with nets and spears.
Anglers catch whitefish on hooks baited with a fresh or salted Emerald Shiners or other minnows or by jigging with spoons and colour-beaded hooks. Ice-fishing is popular in the winter.
Cutthroat Trout / Truite fardée
Oncorhynchus clarkii (Richardson, 1836)
Taxonomy
There are two subspecies of Cutthroat Trout, the Coastal Cutthroat Trout (Truite fardée côtière), Oncorhynchus clarkii clarkii (Richardson, 1836), and the West-slope Cutthroat Trout (Truite fardée du flanc de l'ouest), Oncorhynchus clarkii lewisi Suckley, 1874. The subspecies of specimens introduced to the NCR is unknown. Other common names for the Coastal Cutthroat Trout include Red-throated Trout, Clark's Trout, Sea Trout, Short-tailed Trout, Black- spotted Trout and Harvest Trout and for the West-slope Cutthroat Trout include Red-throated Trout, Lake Trout, Short Tailed Trout, Native Trout, Black-spotted Trout, Montana Blackspot and Yellowstone Cutthroat Trout. This species was formerly placed in the genus Salmo.
Key Characters
The two subspecies are most easily separated by distribution in their native range in western Canada. There is much overlap in their characters and they are difficult to separate. The species itself is identified by the low number of principal anal fin rays (8-12), small dark spots without halos on the whole flank, teeth on the head and shaft of the vomer bone in the roof of the mouth and on the back of the tongue, and the two unique red to orange streaks on the underside of the jaw - the "cut throat".
Description
Dorsal fin with 8-11 principal rays, pectoral rays 12-15 and pelvic rays 9-10. Lateral line scales 116-240 and pyloric caeca 24-60. Males develop a slight kype or hooked lower jaw when in breeding condition.
Colour
Coastal Cutthroat Trout:- Colour is extremely variable and both subspecies have been introduced into the other's range. Hybrids with Rainbow and golden trout further confuse colour patterns and complicate identifications. The "cut throat" marks may be absent in sea dwelling fish or recent migrants to fresh water. These fish have more silvery flanks, are more bluish on the back, have yellowish lower flanks and fins, and spots are less evident. Generally the back is dark green to greenish-blue, the upper flank olive-green and the rest of the flank and belly is silvery. The gill covers are pinkish. Flank spots below the lateral line are more numerous anteriorly (cf. west-slope cutthroat). The outline of spots is irregular, not rounded. Spots are present on the dorsal, adipose and caudal fins and the anal, pectoral and pelvic fin bases. Young have 10 oval, grey-violet parr marks along the lateral line covered with small black dots which extend onto the back and tail. The back may have 5 dark ovals and is olive. The leading edge of the dorsal fin is dark. The dorsal and anal fins may have white patches. The adipose fin has a few black spots. "Cut throat" markings develop on fish over 7.6 cm long.
West-slope Cutthroat Trout:- Colour is very variable and is complicated by the introduction of coastal cutthroat trout. Hybrids with rainbow and golden trout further complicate colour patterns and identifications. The body has a yellow-green to olive back and red on the head sides, anterior flank and belly. The red may be present year round or develop in the breeding season. The flanks are sparsely spotted as are the dorsal, adipose and caudal fins. Flank spots below the lateral line are more numerous posteriorly (cf. coastal cutthroat trout). The flanks may have a narrow pink stripe. The roof of the mouth is whitish in spawning males. Young are the same colour as coastal cutthroat trout. Pure stocks are probably rare in Canada because of introductions from other areas and hybridisation with other trouts such as Rainbow Trout.
Size
Reaches 99.1 cm in non-migrating fish, usually smaller in sea run fish. The world, all-tackle record for a "cutthroat trout" weighed 18.59 kg and was caught in Pyramid Lake, Nevada in 1925.
Distribution
The Coastal Cutthroat Trout is found from southeast Alaska south to California including British Columbia as far inland as the Skeena River headwaters and has been introduced elsewhere. The West-slope Cutthroat Trout is found in western Alberta and southeastern British Columbia and has been introduced into Saskatchewan, Manitoba, Ontario and Québec but perhaps is not established. In the U.S.A., it is found south to Montana (and New Mexico as various subspecies) and has been introduced elsewhere. Bergeron and Brousseau (1981) map this species without an exact locality in the NCR but it is not known if it is maintaining itself (Coad, 1986b).
Origin
An exotic species introduced through a human agency.
Habitat
Coastal Cutthroat Trout are found in fresh and salt waters, migrating between them but not extending into the higher reaches of major rivers like the Columbia in British Columbia. Some populations remain in fresh water. This trout prefers smaller streams or those that have long slow reaches before entering the sea. It is also found in small, coastal bog lakes in British Columbia. Their preferred temperature is <20°C.
West-slope Cutthroat Trout are found in lakes and streams up to over 2440 m, favouring headwaters and small tributaries. Some stream populations are permanent residents, others being spawning fish on a migration from a lake.
Age and Growth
Coastal Cutthroat Trout have a maximum life span of 10 years. Sea run trout are not always larger than stream resident fish because they may not spend long in the sea. Males mature earlier than females, as early as 2 years compared to as late as 6 years. Most fish spawn at 2-4 years.
West-slope Cutthroat Trout have a maximum life span of about 10 years. Males mature earlier than females, as early as 2 years compared to as late as 6 years. Most fish spawn at 2-4 years. In Alberta fish are larger and faster growing in the larger streams and growth is generally faster in warmer water. There are also differences in growth between lake and stream dwelling fish, depending in part on the time spent in streams compared to lake residency.
Food
Coastal Cutthroat Trout food is aquatic and terrestrial insects, crustaceans, small fishes and salmon eggs. Migrating salmon are an important food, taken when cutthroats go to sea at the same time, and other fishes eaten include trout, sculpins, flatfishes, rockfishes and sticklebacks.
West-slope Cutthroat Trout food is aquatic and terrestrial insects, crustaceans, frogs, small fishes and their eggs. Fish eaten include trout, sculpins, carps and sticklebacks.
Reproduction
Coastal Cutthroat Trout spawn in January to May in British Columbia after a migration in late autumn and early winter. There may be late runs in December and January in short streams. Small gravel streams are favoured. The female excavates a redd by lying on her flank and lashing her tail. Redds are about 30 cm across and 10-13 cm deep. Males court females with nudges and by quivering. The female lies in the redd with head and tail bent up, the male joins her, they gape, vibrate and release eggs and sperm. The fertilised eggs fall between the gravel. The female dislodges gravel at the upstream rim of the redd to cover the eggs with up to 20 cm of gravel. The spawning pair may have other males sneaking in to shed sperm. Females may dig more than 1 redd and both sexes spawn with more than 1 other fish. Each female may have 2000 or more orange-red, adhesive eggs of 5.1 mm maximum diameter. The red cut throat may be used in aggressive displays. Sea run cutthroat often survive to spawn again, 12% spawning a fourth time in one study. The fry emerge from the gravel in April and can run to the sea in the spring of their second or third year at about 13-20 cm in British Columbia. They live mostly in estuaries or near shore areas for one or more years, re-entering rivers to spawn in the fall or to feed on migrating salmon in the spring. Growth in the sea can be 25 mm per month. There is variation in migration times, sea life span and spawning time between stocks and geographical areas.
West-slope Cutthroat Trout spawn in spring when water temperatures reach 10°C, June-July in Sheep River, Alberta, May-June in the Flathead River of British Columbia, Alberta and Montana. Small gravel streams are favoured. The female excavates a redd by lying on her flank and lashing her tail. Redds are about 30 cm across and 10-13 cm deep. Males court females with nudges and by quivering. The female lies in the redd with head and tail bent up, the male joins her, they gape, vibrate and release eggs and sperm. The fertilised eggs fall between the gravel. The female dislodges gravel at the upstream rim of the redd to cover the eggs with up to 20 cm of gravel. The spawning pair may have other males sneaking in to shed sperm. Females may dig more than 1 redd and both sexes spawn with more than 1 other fish. Each female may have 2000 or more orange-red, adhesive eggs of 5.1 mm maximum diameter. The red cut throat may be used in aggressive displays. Beach and shoal spawning occurs in some lakes. Fry emerge from the gravel in July-August and stay in streams for up to 4 years before migrating to a lake. Repeat spawning occurs in subsequent years and about 18% of a run is fish which have spawned before. About 7% of the run is non-spawners and 75% are first spawners.
Importance
Coastal Cutthroat Trout are an important sport fish caught on flies, spoons and live bait and often leaps when hooked. In the Bella Coola system, B.C., runs of Chum and Pink salmon in April attract cutthroats which are caught using flies that imitate the salmon fry. Federal and provincial authorities are endeavouring to improve tenfold cutthroat populations in southern British Columbia and Vancouver Island by improving stream conditions where these have deteriorated in populated areas. The flesh is orange-red to pink and best when smoked, fried or baked.
West-slope Cutthroat Trout is an important sport fish caught on flies, spoons and live bait although it does not leap as much as some salmons. The flesh is orange-red and best when smoked, fried or baked.
Rainbow Trout / Truite arc-en-ciel
Oncorhynchus mykiss (Walbaum, 1792)
Taxonomy
Other common names include Steelhead Trout, Kamloops Trout, Coast Rainbow Trout, Silver Trout, Half-pounder, Redsides and Pacific Trout. Fish from the upper Columbia, Fraser, Athabasca and Peace River basins are placed in the Redband Rainbow Trout subspecies O. mykiss gairdneri (Richardson, 1836) and those from the coast of British Columbia and the lower Fraser River in the Coastal Rainbow Trout subspecies O. mykiss irideus (Gibbons, 1855). The former subspecies has larger spots, more elliptical parr marks (more rounded in the coastal subspecies), often with additional rows above and below the main parr marks (usually reduced or absent in the coastal subspecies), more yellow and orange tints on the body, a trace of a cut throat mark, and light coloured tips to the dorsal, anal and pelvic fins. The scientific name of the species was formerly Salmo gairdneri.
Key Characters
This species is distinguished by having no red spots on the body but only small dark ones and radiating rows of black spots on the dorsal and caudal fins, no reddish "cutthroat" marks, 100-161 lateral line scales, 8-12 principal anal fin rays, the vomer bone in the roof of the mouth has teeth on its head and shaft, and no teeth at the tongue base.
Description
Dorsal fin principal rays 10-12, pectoral rays 11-17 and pelvic rays 9-10. Pyloric caeca 27-80 and gill rakers 16-22. Breeding males have an elongated snout and the lower jaw is hooked.
Colour
Overall colour is very variable and this is reflected in the common names. Stream fish are darker and more colourful (rainbows) than lighter, silvery lake or sea fish (Kamloops or Steelhead). Some sea-run and lake fish have small orange to red marks below the lower jaw similar to those in Coastal Cutthroat Trout. The back and upper flank are steel-blue, greenish, silvery-olive or even brown, flanks and belly are silvery, grey, white or yellow-green. The side of the head and the flank are characteristically pink. The flank has a broad pink to red or lilac stripe with small black spots. The adipose fin has a black margin and a few spots. Pectoral, pelvic and anal fins may have a few spots and are dusky without any strong markings. Pectoral and pelvic fins are often orange-red. Spawning fish are very dark and the flank stripe is dark red or purple. Breeding males have the roof of the mouth is white. The young have 5-13 dark, oval parr marks centred on the lateral line with the spaces between the marks wider than the marks. There are 5-10 parr marks on the back in front of the dorsal fin. The upper flank has some dark spots. The dorsal fin is tipped white or orange and has a dark leading edge, sometimes broken up into spots. The adipose fin has a black margin. The anal fin has a white or orange tip. Black spots are few to absent on the tail. Some adults in streams do not lose their parr marks. Golden and Palomino trout are genetic variants of Rainbow Trout reared in hatcheries and released into the Great Lakes. Golden Trout (not true Golden Trout, Oncorhynchus aguabonito (Jordan, 1892)) are golden-orange without black spots and Palomino Trout are golden with a few black spots but much less than normal rainbows.
Size
Reaches 122.0 cm and 25.8 kg as sea-run or lake fish but smaller in streams. Fish less than 10 kg are trophy-size in most of Canada. The 25.8 kg fish was from Jewel Lake, British Columbia. The world, all-tackle angling record from Bell Island, Alaska caught in 1970 weighed 19.1 kg. The Ontario record as of the year 2000 weighed 13.2 kg.
Distribution
Found in coastal waters and basins draining to the ocean in British Columbia and extreme southwest Yukon and in the upper Peace and Athabasca river basins. Also from Japan and Alaska to Mexico. Widely introduced outside this natural range in all provinces of Canada and world-wide in suitable waters. Broad areas of introduction are southwestern Québec, southern Ontario, all the Great Lakes and across the southern Prairie provinces but also at various places in the Yukon, N.W.T., the Maritimes and in Newfoundland since 1887. This species is known from trout farms in the NCR (Coad, 1986b), and has been stocked in various waters including Meach Lake and Bernard Lake (subsequently disappeared after a 5 lbs (2.3 kg) one was caught in the former lake), the Rideau River above Hog's Back (a half dozen adults from the Central Canada Exhibition, Ottawa in 1935, 1936 and 1937)(Dymond, 1939), and in the Val-des-Bois and other areas of Québec (Finn, 1964; Dumont et al., 1988). None were caught in a survey of the Rideau River between 1998-2000 (www.rideauvalley.on.ca/programs/rrr/rrr.html, downloaded 15 July 2002). It has been suggested that it could be successfully stocked in the Ottawa River at Ottawa (Hopkins, 1996a).
Origin
A species introduced by man to the NCR.
Habitat
Rainbows are found in rivers or streams where there are pools and riffles. Some live in lakes and are called Kamloops Trout while others run to sea and are called Steelheads in their natural habitat. They can tolerate temperatures up to 24°C, warm for a trout, but prefer temperatures below 20°C and their preferred temperature is 11.3°C. The Great Lakes have introduced Steelhead stocks. Lake populations run up streams to spawn while river fish enter headwater streams.
Age and Growth
Life span varies with habitat, up to 11 years in some lake fish, 8 years in the Great Lakes but only 3-4 years in many streams and small lakes. Growth varies with habitat including such factors as length of sea, stream or lake life, years before spawning, available food supplies, latitude, altitude, temperature regime, competition with other salmonids, and so on. Ageing these fish may be difficult because of the complicated life history pattern of stream and lake residency. Maturity is also variable with habitat. Some males mature at 9 months in fish introduced to warm southern waters and some females only at 8 years, but generally maturity is reached at 3-5 years in Canada with males maturing a year earlier than females.
Food
Food includes plankton, crustaceans, aquatic and terrestrial insects, snails, leeches, worms, molluscs, salmon eggs, and many fishes. The fish eaten enhance growth and the species taken depends on what is available. Rainbows are food for various other Salmon Family members found in the same habitat, most predatory fishes, as well as various birds and some mammals. Lampreys attack this trout.
Reproduction
Repeat spawning can occur for up to 5 years and in the NCR this can occur in streams and lakes. Spawning takes place from March to August but is usually in spring. Great Lakes fish may spawn from late December to late April. Water temperature for spawning usually exceeds 10°C but may be 5-13°C. A female excavates a redd by lying on her side and thrashing her tail. Redd excavation occurs during the day and night and dimensions are usually longer and deeper than the female's body. Females construct several redds and may spawn with several males. A male courts a female by rubbing his snout and body against her, by vibrating, by swimming over her in the redd and by pressing against her. Several males are found around each female but one male is dominant. The spawning act lasts 5-8 seconds with the pair parallel in the redd pressed together, both fish gape, arch and vibrate. Other males may shed sperm. The female covers the eggs with gravel by dislodging it from the upstream end of the redd. Most spawning takes place in the morning and evening and nests may be abandoned during the day. Eggs are orange or pink, 5.0 mm in diameter and up to 12,749 in number. Egg numbers are usually a few hundreds to thousands. The eggs hatch in about 8 weeks and fry generally emerge in June to August from spring spawnings.
Importance
Rainbow Trout are one of the top few sport fishes in North America and of great commercial importance because of the money spent on gear, accommodation, transport, etc. by anglers in pursuit of this fish. Many books and articles have been written on the methods and joy of catching this trout. It may be caught on flies, with lures or with various still-fished or moving baits. It is a strong fighter which bites easily and leaps often, and may "cartwheel" and "tail-walk". Dawn and dusk are the best times to catch rainbows in streams. Trolling an artificial fly is used on large lakes. Baits and small jigs are used in ice fishing. The flesh is excellent eating fresh or smoked and may be red if food is mostly invertebrates or white if food is fishes. The total Canadian catch in 1988 was 3658 tonnes. Farmed fish are sold frozen world-wide, the most important trout in this regard, and ironically some of that found in Canada comes from Japan or Europe. Small put-and-take fisheries are common wherever this species is found or has been introduced. Rainbows have been used extensively as research animals as their requirements are well known and they are readily available from hatcheries. Hatchery fish often have reduced or absent fins and deformed mouths. Introductions of this fish to various waters throughout Canada may have serious consequences for other species.
Atlantic Salmon / Saumon Atlantique
Salmo salar Linnaeus, 1758
Taxonomy
Other common names include Saumon de l'atlantique, Ouananiche, Landlocked Salmon, Lake Atlantic Salmon, Sebago, Kennebec Salmon, Black Salmon, Grayling, Bratan, Saumon d'eau douce, Sâma. Saama, Saamakutaak and Kavisilik.
Key Characters
This species is characterised by having 109-124 lateral line scales, 10-12 principal dorsal fin rays, 8-11 principal anal rays, vomer bone in the roof of the mouth with teeth on its head and shaft, no spots or rows of spots on the caudal fin, 2-3 large spots on the gill cover, upper jaw not usually extending beyond the eye, no red on the flank or adipose fin in adults.
Description
Pectoral rays 14-15 and pelvic rays 9-10. Pyloric caeca 40-74. Spawning males develop a kype, or hooked lower jaw.
Colour
The back is brown, green or blue, flanks are silvery, and the belly is white. Black spots, often x- or y-shaped are found on the upper flank and back and sometimes but not usually on the caudal fin. Spawning males have bronze to dark brown colour and may have red, orange or rust-brown spots on the head and body. The pectoral and caudal fins may darken. Spawning females are grey to purple-blue and may become blackish. After spawning they become very dark and are known as kelts, slinks or black salmon. Young fish have a red spot between each of 8-11 narrow, parr marks. Young fish migrating to sea, known as smolts, racers, grilts or fiddlers, become silvery. Ouananiche have dark brown, bronze or bluish flanks with larger spots than sea-run fish and sometimes light halos around the spots.
Size
Reaches 150.0 cm and 35.9 kg. The world, all-tackle angling record from Norway in 1928 weighed 35.89 kg. The Ontario record as of the year 2000 weighed 11.0 kg. The land-locked salmon (Ouananiche) angling record is 10.31 kg for Lake Lobstick, Newfoundland in 1982.
Distribution
Found from Ungava Bay south to the Gulf of Maine throughout Atlantic Canada including the Grand Bank and over deep ocean east of the Grand Bank, and up the St. Lawrence River, once to Lake Ontario. Rare specimens enter eastern Hudson Bay. Across the Atlantic to Greenland, Iceland and in western Europe. There are a number of naturally land-locked populations, known as Ouananiche. Blais and Legendre (1978) give several localities in Québec where Ouananiche and Atlantic Salmon have been introduced, many close to the NCR, but continued survival is unknown (Coad, 1986b). Only one locality falls within the NCR, at Lac St-Charles east of Brennan Hills and the Gatineau River (45°46'30"N, 75°52'25"E) where 1000 Atlantic Salmon alevins were introduced in 1966 by the Club Byng. Ouananiche were also stocked in Lac Deschênes of the Ottawa River on 31 October 1989, numbering 400 2+ age fish but none were ever caught (Chabot and Caron, 1996). Also introduced to Ontario, Alberta and British Columbia.
It remains uncertain that salmon ever ascended the Ottawa River as far as the NCR. Van Cortlandt (1865b) states "this splendid fish never visits the Ottawa waters, nor, as far as I can learn, ever has done so." He later states "That the true Salmon, Salmon Salar, was wont to visit our tributaries, and this too within a very limited period, does not admit of the shadow of a doubt" (Van Cortlandt, 1867; reprinted 1999, possibly accounting for the spelling "Salmon Salar"). Small (1883) comments that there is a lack of any authentic data that the salmon was ever abundant in the Ottawa River prior to sawdust and mill refuse pollution. This implies, but does not confirm, a presence. He then states that the name Salmon River near Montebello "was retained ..... long after the native fish had abandoned its waters", again implying a presence but without clear confirmation. Early attempts at stocking the Salmon River at Montebello failed. There was a hatchery in Ottawa founded in 1890 that produced Atlantic Salmon (Rodd, 1912) but whether these were ever stocked locally was not stated therein. Egan (1999) mentions salmon making it up the Ottawa as an attraction for settlement of Hull by Philemon Wright in 1800 and Egan (1996b) mentions a run reported by early settlers up to the Chaudière Falls. Salmon (assumed to be Atlantic Salmon but possibly a Pacific species introduced to the the Great Lakes) have been caught at Hawkesbury below the Carillon Dam. If they are Atlantic Salmon then they are presumably strays from a stocking programme or migrants from the Atlantic Ocean (Egan, 1996a; 1996b; 1997).
Origin
This species has its origin in an Atlantic refugium.
Habitat
Atlantic Salmon adults are found in coastal waters and some travel as far as Greenland, a migration of about 2400 km, or even Norway. They home to their birth river to reproduce after 1 or more years at sea. The ability of salmon to home to their birth stream over thousands of kilometres has not been fully explained. The earth's magnetic field, ocean currents or stars may be used to navigate to coastal waters where smell directs them to their stream of birth. Fish returning to freshwater after 1 year are known as grilse and weigh 1.4-2.7 kg. Fish with 2 sea years weigh 2.7-6.8 kg and are known as salmon. Ouananiche are generally smaller than fish which have access to the rich food in the sea. Ouananiche in Gros Morne National Park, Newfoundland have slow growth, a small maximum fork length of 25 cm, low fecundity at a maximum of 268 eggs, and retain juvenile characters. Young salmon or parr spend 2-4 years in fresh water in cool streams and rivers until 12-15 cm long when they run to sea as smolts. In Ungava this is delayed until 4-8 years of age and 18 cm. They may spend 6 or more years in the sea. Landlocked salmon remain in a lake and run into tributaries to spawn. Some Ouananiche may have access to the sea but do not migrate while others are truly landlocked by physical barriers. Males grow faster and are larger than females. Their preferred temperature is 16°C.
Age and Growth
Maximum life span is 11 years as some fish survive spawning and run to sea more than once. Salmon in the sea grow much more rapidly than those in freshwater because of the year-round access to a wide range of foods compared to a 4-month feeding period restricted to smaller, less varied and sparse aquatic insect supplies. Maturity is attained at age 3. Fingerlings of Ouananiche (landlocked Atlantic Salmon) have been introduced to Papin Lake in Pontiac County (76°06'N, 46°09'W) just outside the NCR from 1961 to 1973 (Cuerrier, 1983). Growth was rapid as Pygmy Smelt (Osmerus spectrum) had also been introduced as a forage fish. After three summers some salmon had attained 2.0-2.84 kg with sexual maturity at age 2+. The largest fish caught was over 9.0 kg and the oldest was 8 years. There was no natural reproduction in the absence of suitable spawning streams and salmon are no longer present.
Food
Food of young salmon in freshwater is aquatic and terrestrial insects, crustaceans and fishes and of adults in the sea various crustaceans and a wide variety of fishes such as Capelin, Rainbow Smelt, Gaspereau, Atlantic Herring, Atlantic Cod, sand lances and mackerel. Salmon in lakes prefer Rainbow Smelt. Salmon on the spawning run do not feed and why they readily strike at wet and dry flies is something of a mystery. American Eels, sharks, Swordfish, Pollock, tuna, birds and seals all eat salmon as young or adults.
Reproduction
Although some fish spawn twice, very few survive the arduous migrations and spawning acts to reproduce for a third time. Rarely some fish spawn 8 times. Repeat spawners may comprise as much as 34% of the Grand Cascapedia River, Québec population while in other rivers repeat spawners are as low as 5%. Salmon spawn in October and November after entering estuaries from spring to autumn. Ouananiche in the Outaouias spawn too in October and November at water temperatures around 5C (Thellen, 1994). The time of entry to each river is the same each year. The timing of entry depends in part on distance to travel. Spring runners often run to distant headwater streams and fall runners to lower reaches. The migration is not blocked by rapids or low waterfalls which the salmon leaps spectacularly, sometimes over 3.5 m. The female excavates a redd in gravel bars and riffles by turning on her side and lashing her tail. Redds are up to 5.9 m long and 0.9 m wide and are found in water about 25 cm deep. Non-anadromous salmon in Newfoundland scatter their eggs among boulders while the anadromous salmon in the same stream construct redds. The male drives away competitors particularly grilse. Large male reproductive success depends on successful competition for access to a female as the large male rarely survives spawning. The female enters the redd by backing in and tests it with her anal fin for spawning suitability. She is joined by the male and eggs and sperm are shed as the pair quiver and gape. The male may nudge the female and glide across her. The female uses her tail at the upstream end of the redd to dislodge gravel and bury the eggs as deep as 25.4 cm. Small male parr, only 10-15 cm length, rush in to fertilise eggs during the spawning act. These are known as "sneaky males". In some Canadian waters, such as the Matamek River, Québec, the proportion of grilse has increased from about 70% to about 85-90% and precocious parr (sneaky males) are much more numerous. The capture of large adults at sea by commercial fishermen has shifted reproduction to earlier life history stages. The shift is genetic as grilse matings result in offspring which return to spawn as grilse rather than salmon. The large adults repeat the redd excavation and spawning for a week or more. Eggs are up to 7.0 mm in diameter, orange to amber and number up to 629 per kilogram of female body weight (and some females have over 20,000 eggs). Spawned out fish or kelts may rest in a pool for a few weeks or over winter or run to sea immediately. Eggs hatch in April after overwintering and the young fish, known as alevins, emerge from the gravel in May and June. Small salmon are called fingerlings or underyearlings and parr once the characteristic flank marks develop. Parr retreat into the gravel and under large rocks in the fall when temperatures drop to 9-10°C and re-appear only in spring and early summer when waters warm up. During the summer, most parr are found above the stream bed holding station over a stone.
Importance
Atlantic Salmon are the quintessential sport fish as well as having great commercial importance. Comments on the lore and techniques of catching salmon are superfluous as numerous books have been written. They are easy to catch on the spawning migration using flies. Salmon rivers are a valuable commercial resource, access being controlled by public and private clubs. Their concentration in rivers for spawning make them susceptible to environmental changes and their high seas life to commercial exploitation by nations without an interest in conserving freshwater runs. The coastal fishery in Newfoundland and Labrador for example includes some fish from other Canadian provinces and from Maine. The West Greenland area is very important for maintaining Canadian stocks which are dependent on limitations on this fishery. About 40% of Canadian fish visit the Greenland feeding grounds and some travel as far as the Faroes. The annual North Atlantic catch is about 10,000 tons or 3 million fish. The Canadian catch in 1988 was 3847 tonnes and landings were worth about $4 million. Lake Ontario salmon were fished to extinction by 1890, compounded by construction of dams which blocked spawning streams. In addition to dams throughout its range, various pollutants and acid rain are just a few of many threats to this species. The species is now being farmed in Canada with cage-reared fish in the Bay of Fundy being worth an estimated $16 million in 1987, with figures for more recent years expected to reach about $100 million. Attempts are being made to incorporate the "anti-freeze genes" from winter flounder into salmon to facilitate winter survival in inshore waters and so make cage-culture more effective.
Brown Trout / Truite brune
Salmo trutta Linnaeus, 1758
Taxonomy
Other common names include German Brown Trout, European Brown Trout, Sea Trout, Brownie, Loch Leven Trout, Von Behr Trout, Spotted Trout and Liberty Trout.
Key Characters
This species is a relative of the Atlantic Salmon and is distinguished from it by the upper jaw extending beyond the eye in adults (and below rear half of the eye in young rather than the centre in young), the gill cover has many spots, dorsal fin principal rays usually 10-14, and orange to rusty-red spots are often present on adult flanks.
Description
Principal anal fin rays 9-12, pectoral rays 13-14 and pelvic rays 9-10. Lateral line scales 110-136 and pyloric caeca 30-60. Males develop a hooked lower jaw when spawning and the brown colour becomes more intense and golden.
Colour
Overall colour is a light to golden brown with silvery flanks and a white to yellowish belly. The back, flanks, side of the head and dorsal and adipose fins bear black or dark brown spots, often with a lighter halo of orange, pink or red, and the flank has pink or red spots. Only Brown Trout have both light and black spots on the flanks. The caudal fin may have spots restricted to its upper lobe but lacks the overall radiating spots of Rainbow Trout. The adipose fin is orange to orange-red, the only family member with this colouration. Sea run or lake fish are more silvery, obscuring some of the spots. Spots may be x- or y-shaped. The red flank spots usually have blue halos. Young have 7-14, narrow parr marks and a few red spots along the lateral line. The adipose fin is orange with a light margin. Hybrids with Brook Trout are called Tiger Trout for their distinctive markings
Size
Reaches 1.4 m and reputedly 50 kg. The world, all-tackle angling record from Arkansas in 1992 weighed 18.25 kg. The Ontario record as of the year 2000 weighed 15.6 kg. Brown Trout over 2 kg and 16-24 inches long have been caught at Remic Rapids in Ottawa from smaller stocked fish (see below).
Distribution
Found in all Canadian provinces, except P.E.I., Yukon, Nunavut and Northwest Territories, as an introduced species from Europe and western Asia. Most widely distributed in southern Ontario and Alberta, sporadically elsewhere.
In the NCR, this species has been introduced to the Mississippi River (Anonymous, 1932; Dymond, 1939; the latter noting that none survived there to be caught by anglers despite numbers of 15,000 in 1931, 25,000 in 1932 and 10,000 in 1934 and 1500 15 month old fish in 1937); the Rideau River below Hog's Back in 1937 (1600 yearlings) and above Hog's Back (a half dozen adults from the Central Canada Exhibition in Ottawa); in waters of the Seigniory Club near Montebello, Québec (Dymond, 1939); 3600 in the Rideau River at Mooney's Bay just above Hog's Back in 1941, many eaten by gulls at low water the following spring (newspaper reports; Bebee, 2004); east of Hull (Dumont et al., 1988); the Kemptville area (Lasenby and Kerr, 2001); and the Remic Rapids in the Ottawa River in the late 1980s, in 1997 (15,000 three-inch fish (Hopkins (1997a; c)) and again in 1999-2000 (5000 fish mostly at 15-20 cm and some adults at 50 cm(Hopkins, 2000)) or in another report 10,000 7 inch fish, 500 10 inch fish and 120 fish between 2.5 and 5.5 lbs (www.flyanglersonline.com/features/canada/can131.html, downloaded 3 June 2003). This latter source reports that it was the Province of Québec that stocked rapids west of Ottawa in 1987 with 14,000 fingerlings and 11,000 fingerlings and 20,000 7 inch fish between 1990 and 1994. Chabot and Caron (1996) summarise stocking reports from the Deschênes Rapids which presumably includes "rapids west of Ottawa" mentioned above. They give 14,000 fingerlings in 1987, 5000 1+ age young on 14 May 1990, 10,000 fingerlings on 17 October 1990 (Hopkins (1991b) has 15,000 fish in 1990, most fingerlings but 5000 one-year-olds and 100 two-year-olds), 100 2+ fish on 30 November 199? (date incomplete, ?1991), 5000 1+ fish on 30 April 1992, 5000 1+ fish on 22 April 1993, 5000 1+ fish on 3 May 1994 and 7500 fingerlings on 27 October 1994, as well as 5000 1+ fish at Kettle Island in 1992.
None were caught in a survey of the Rideau River between 1998-2000 (www.rideauvalley.on.ca/programs/rrr/rrr.html, downloaded 15 July 2002) but some are caught in the Ottawa River near Ottawa still.
Several local creeks were proposed as stocking sites for brown trout, namely Findlay or Kelly near Leitrim, Poole in Stittsville and Shield near Osgoode (Hopkins, 1993d) but this may never have happened.
Origin
This species is introduced by man to the NCR.
Habitat
Brown Trout are mostly stream and river dwellers although some are in lakes and ponds. They can tolerate warmer and more turbid waters than Brook Trout and only the Rainbow Trout is more tolerant among salmonids. Brown Trout may survive in areas no longer suitable for Brook Trout because deforestation has increased stream temperatures and agriculture and industry have increased pollution and turbidity. Ideally there should be overhanging and submerged vegetation, coarse gravel and cover such as logs, boulders and undercut banks, growth is better in alkaline waters (20-200 mg L-1, total dissolved solids 800 mg L-1 for culturing this species), a depth of 7-58 cm is needed for spawning, a winter flow of less than 15 cm sec-1 is required, preferred temperatures are 10.0-17.6°C (other reports give 21.1°C and temperatures up to 24°C are tolerated) and for spawning 6.7-8.9°C, dissolved oxygen is optimal at 7-9 mg L-1 and pH range tolerated is 5.0-9.5 although the optimal range is 6.8-7.8.
Age and Growth
Life span is over 18 years with maturity attained at 2-4 years.
Food
Food is aquatic and terrestrial insects, crustaceans, molluscs, various fishes, frogs, salamanders and even small mammals. Most food is taken as drift, the trout positioned in slower water behind a rock and darting out into faster current to seize prey. Predators include other fishes, birds, water-snakes and otters.
Reproduction
Spawning occurs from October to January depending on locality, usually at 7-9°C but as low as 2°C or as high as 14°C. In the Outaouais, reproduction occurs in the second half of October and in November (Thellen, 1994). It usually takes place in gravel stream riffles but may occur on rocky shores of lakes. The female excavates a redd into which the spawning pair deposit eggs and sperm while gaping and quivering over a 4 second period. The female covers the redd with gravel after spawning to protect the eggs. Subsequent spawning occurs, usually after a 10 hour interval. Occasionally a community redd is excavated by a number of fish spawning close together. Eggs are up to 5.0 mm in diameter and each female can contain 20,865. Young may spend about 2 years in their natal stream before going to a lake but some fish remain permanently in streams.
Importance
This trout is a valuable and popular sport fish in Europe, less so in Canada because of its localised introductions. The benefits of stocking Brown Trout include its difficulty of capture (and thus appeal to anglers), its longer contribution to the fishery than Brook Trout and Rainbow Trout, better survival and growth than other trouts, and its tolerance of warmer waters. There is an extensive European literature on angling methods and it is reputed to be a wilier fish than native Brook Trout in Canada. However studies in an Ontario stream compared catchability favourably with native species. They tend to bite best in the late evening or at dawn, especially when large. These trout are caught on worms, crayfish, lures and flies. Brown Trout have white to pink flaky flesh and are excellent eating. Flesh colour changes with age, to pink, and is related to diet.
Arctic Char / Omble chevalier
Salvelinus alpinus (Linnaeus, 1758)
Taxonomy
Other common names include Alpine Char, Silver Char, Hearne's Salmon, Sea Trout, Coppermine River Salmon, European Char, Hudson Bay Salmon, Mountain Char, Arctic Salmon, Blueback Trout, Greenland Char, Québec Red Trout, Omble rouge du Québec, Truite rouge du Québec, Ekaluk, Ilkalupik, Kaloarpok, Ivatarak, Iloraq, Aniaq, Nutidilik, and many others. Char may be spelled charr.
There is considerable variation in characters of Arctic Char around the Northern Hemisphere and even within one lake. The species is known as the Arctic Char "complex" as an acknowledgement that more than one species or subspecies may exist. Landlocked forms in Québec were known as Québec Red Trout or Marston Trout and was given the scientific names marstoni. They are generally recognised now as relict Arctic Char, descendants of an older, anadromous form of char which became landlocked and isolated from later char forms invading eastern North America. An alternative view is that char forms are the result of generalist or specialist life history styles, the former in perturbed environments and the latter in unperturbed environments.
Key Characters
This species is characterised by having small scales (123-152 in the lateral line), few major anal fin rays (8-11, total 11-15), teeth only on the anterior or head end of the vomer bone in the roof of the mouth, a truncate or square-cut caudal fin, and colour pattern.
Description
Teeth are present on the jaws, tongue and the palatine and basibranchial bones. Major dorsal fin rays 10-12 (total 12-16), pectoral rays 14-16 and pelvic rays 9-11. Pyloric caeca 20-75, upper arch gill rakers 7-13, lower arch rakers 12-19. Males develop a kype or hooked lower jaw during spawning in anadromous populations. Anadromous males also develop a dorsal ridge anterior to the dorsal fin and enlarged teeth, characters not found or only poorly-developed in landlocked males.
Colour
Colour varies between landlocked, migratory and spawning fish and between individuals by sex. However Arctic Char lack spots or vermiculations on the dorsal and caudal fins found in its relatives, the Lake Trout and Brook Trout. The pectoral, pelvic and anal fins have a white leading edge, followed by contrasting black and red bands, with the rest of these fins and other fins dusky. Generally non-spawners are an overall silvery colour. A spawning male has a brown, dark green to blue-green or steel blue back, flanks are silvery-blue with white to cream, orange to pink or red spots larger than the pupil and the belly is white or golden to a bright orange-red. Some populations lack spots while in others the spots have a blue halo. This bright spawning colour may be retained year-round in some freshwater populations. Isolated, stream-resident males retain parr marks, have yellow flank spots and are nearly black dorsally. Young Arctic Char have 8-17 oval parr marks along the flank.
Size
Reaches 101.6 cm and 16.0 kg. The world, all-tackle angling record from the Tree River, N.W.T. weighed 14.77 kg and was caught on 30 July 1981 by Jeffrey L. Ward.
Distribution
Found around the Northern Hemisphere including the Arctic coast and islands of Canada, Labrador, and northern Québec, Newfoundland, the Québec north shore and in lakes of southwestern Québec and New Brunswick. Introduced to Alberta. Also in lakes of Maine and New Hampshire in North America. In the NCR they may no longer be present in many of the lakes mapped through overfishing and competition with introduced Smallmouth Bass.
Origin
This char survived in an Atlantic coastal refugium or a Beringian refugium (Mandrak and Crossman, 1992).
Habitat
In the NCR char are found in deep and cool waters of lakes during the summer months. They may approach the surface or shallow water during the cooler days of spring and fall. Elsewhere they may be found in streams and rivers.
Age and Growth
Life span exceeds 40 years and growth varies between localities. Isolated stream resident fish seldom live longer than 10 years however. Landlocked fish tend to be smaller than searun fish of the same age. In Candlestick Pond, Newfoundland landlocked char have slow growth, a short life span of 7 years, small size (to 16.4 cm fork length), maturity at age 3 and low fecundity (104 ova per fish maximum). Maturity can be attained at 1 year for landlocked females in Newfoundland but anadromous fish can mature as late as 10-25 years with ages in between for various populations.
Food
Arctic Char are predators on various crustaceans, insects, snails, clams and fishes, feeding in both salt and fresh water on whatever is available. Amphipods are an important food when at sea near Baffin Island. In a Newfoundland lake Rainbow Smelt were the predominant food. Diet shifts occur depending on which, if any, fish species the char is sympatric with. They are cannibals and are eaten by some birds and seals.
Reproduction
Spawning occurs in August to October in the far north or as late as December-January in southern Québec. Char which spawn do not migrate to the sea but may go downriver as far as estuaries. Females spawn only every 2-3 years in the north but may do so annually in the south. The absence of a sea migration in northern populations which spawn is a major energy drain and prevents fish from spawning 2 years in succession. In the central Canadian Arctic and Arctic islands spawning takes place in lakes as the rivers are completely frozen in winter. Females clear a nest area by turning on their sides and flapping the caudal fin on stream gravel or rocky lake shoals. Pair spawning takes place at about 4°C during the day. Males guard territories and may spawn with more than 1 female. The male circles the female on the redd, quivering as he passes along her flank. The female and male quiver and release the sex products during one of the passes by the male. The female undulates over the eggs to force them into the gravel and dislodges upstream gravel to cover the eggs but only after several spawning acts. A female may construct 10 or more redds but is eventually abandoned by the male when she is spawned out after 4 hours to 3 days. The male will court a new female. Eggs are up to 5.5 mm in diameter when deposited and number up to 9245. The eggs hatch at ice break-up in the following spring. Temperatures over 7.8°C will kill the eggs.
Importance
Arctic Char have long been used as food for humans and for dogs in the north and are caught in nets, traps and by spearing. They are eaten fresh, smoked, salted or dried. The flesh is red, pink or white. The 1988 Canadian catch was 88 tonnes valued at $486,000. They are strong fighters and sought after by anglers. Smaller fish may leap though large ones seldom do so. Char are caught on streamers, spoons and dry flies, which they may follow for long distances before striking. Char populations grow slowly and have a low fecundity so they can easily be fished out. However some char are now being farmed in fresh and salt waters. Char are important research tools for biologists, being the only fish species present in some undisturbed northern lakes. They can be used to test conceptual models since the ecosystem is much less complex than the multi-species lakes of southern Canada.
Brook Trout / Omble de fontaine
Salvelinus fontinalis (Mitchill, 1814)
Taxonomy
Other common names include Brook Char(r), Speckled Trout, Eastern Brook Trout, Brookie, Square-tail, Sea Trout, Salter, Redspotted Trout, Mud Trout, Slob, Coaster, Harness Trout, Native Trout, Mountain Trout, Speck, Whitefin, Truite mouchetée, Truite de mer, Truite de ruisseau, Truite saumonée, Iqaluk Tasirsiutik, Anokik, Anuk, Aanak, Anakleq, Aanaatlik, A Na, I Ha Luk and Âna.
Key Characters
This species is characterised by 109-132 lateral line scales, 8-13 anal fin principal rays, teeth on the head of the vomer bone in the roof of the mouth, truncate caudal fin, pectoral, pelvic and anal fins with a white leading edge followed by contrasting black, red spots ringed with blue on the dark body, dorsal and caudal fins have wavy, dark lines and blotches and the back has dark or light green or cream, worm-track markings (vermiculations).
Description
Dorsal fin principal rays 9-14, pectoral rays 10-15 and pelvic rays 7-10. Pyloric caeca 20-55. Spawning males develop a hooked lower jaw or kype and have larger pectoral and anal fins than females and longer upper jaws.
Colour
The back is olive-green to dark brown or blackish fading to a silvery-white belly. Flanks have a red to yellow tint. Flank spots are pale but there are also small, red spots with blue halos. The pectoral, pelvic and anal fins are yellow, orange, or reddish behind the white and black leading edges. Trout in large lakes are also more silvery than stream resident fish. The jaw tips and the roof of the mouth are blackish. Spawning males are much brighter in overall colour and have an orange-red lower flank and upper belly, bordered below by black on each side which delimits the white belly. Young have 6-12 brown parr marks, the widest equal to eye diameter, and small red, yellow or blue flank spots. The white leading edge to the lower fins is apparent.
Size
Attains a reputed 86.0 cm but most are smaller. The world, all-tackle angling record was caught in the Nipigon River, Ontario in July 1916 by W.J. Cook and weighed 6.57 kg.
Distribution
Found from the shores of Hudson Bay and Labrador south in marine waters to Cape Cod in the east and Georgia in the Appalachian Mountains, west through all the Maritime provinces, Québec and Ontario (except the extreme west) to northeast Manitoba. Widely introduced to western Canadian provinces, the U.S.A., South America, Europe, Asia and Australasia. In the NCR, it is almost entirely restricted to the Québec side although there are some surviving in Kelly's Creek in Ontario. Hopkins (1993c) records trout in the past from Poole Creek in Stittsville but none in a September 1993 survey despite 500 being stocked in the spring, presumably because of warm summer temperatures. Findlay (= Kelly's) Creek in Leitrim was stocked in spring 1993 too. Hopkins (1993c) also reports trout formerly in Shields Creek at Greely. The disappearance of Brook Trout in Gatineau Park lakes is attributed to the introduction of Smallmouth Bass which both ate them and competed with them for the same foods (Rubec, 1972; 1975). Rubec (1975) lists 15 lakes having this species in the late 1800s in Gatineau Park and only 5 of these still retain populations at the time of his survey. Brook Trout have been widely introduced into smaller lakes in Gatineau Park, such as Black Lake which provided sport for anglers in 1972 and 1973, while those introduced to Lusk Lake had limited reproduction (Rubec, 1972).
Origin
Brook Trout could have entered the NCR from either an Atlantic or Mississippian refugium or perhaps both. Mandrak and Crossman (1992) refer to an Atlantic refugium origin.
Habitat
Brook Trout are found in cool waters of streams and lakes, usually at less than 20°C. Their preferred temperature is 16.0°C. Pools, underneath banks, under overhanging bushes or behind rocks are favoured spots. Most specimens from the NCR were captured over stony or gravel bottoms with slow to medium currents. During summer months they retreat to deeper water, to about 8 m, in lakes. Populations in the Great Lakes live and feed mostly in the lake and run up natal streams to spawn. They are known as "coasters".
Age and Growth
Maximum life span is over 20 years but most reach only 5 years. Maturity is attained at 2-3 years, some males at 1 year. Growth is often faster than other trouts and chars. Stunting is common in small streams while sea run fish grow faster than freshwater ones. Optimum growth temperatures are 10-19°C.
Food
Food includes aquatic and terrestrial insects, molluscs, crustaceans, worms, various fishes, frogs, salamanders and even snakes, mice, voles and shrews. Stream-dwelling fish feed heavily on drifting aquatic organisms during spring run-off but in summer as drift decreases surface insects become important. Most feeding occurs in the early morning and late evening although some food is taken throughout the day. Diet shifts in response to competition with other species. Trout feed on large, bottom invertebrates in some Québec lakes but switch to zooplankton when found with Creek Chub. Trout are more aggressive in groups but chub forage successfully in groups. Sea run fish take various marine invertebrates and fishes. Sea run adults in spring and summer eat crustaceans and fish in lower estuarine areas while young are in the upper estuary eating crustaceans and insects. During the fall in the river adults eat little and during winter back in the estuary consumed mostly crustaceans. There is thus a division of food resources between young and adults. Brook Trout are cannibals on their eggs and young and are eaten themselves by other fishes, water snakes, turtles, various birds and otters.
Reproduction
Spawning takes place from August to December, earlier in the north and later in the south. Sea-run trout enter their natal stream in spring and summer even though spawning occurs in fall. Each year they spend 1.5-3 months feeding in the sea. The spawning ground is usually gravelly streams but may be lake shoals if there is some current or spring outflow to keep eggs oxygenated. Spring flows are preferred even in streams. Males arrive on the spawning ground first and defend a territory. Both sexes will rush at other fish entering the redd area. The female cleans a redd of debris by turning on her side and lashing her tail. Redd depth between stones is tested by inserting the anal fin. Redd construction may take up to 2 days with work being carried out both by day and night. Courtship involves gentle pushes, touches and strokes of the female by the male. The female is ready to spawn when she crouches in the redd with her genital area between the stones. The male arches his body and may press the female against the redd bottom, both fish vibrate and eggs and sperm are shed. Accessory or sneaky males may rush in to shed sperm. The female lashes her tail to push eggs into the gravel and then dislodges gravel with her anal fin to cover the eggs to depths as great as 20 cm. Yellow-orange eggs are up to 5.0 mm in diameter and number up to perhaps 17,000 per female although averages range from the low hundreds to a few thousand. Both sexes may spawn again with other fish. The eggs develop over winter, taking 165 days at 2.8°C but only 47 days at 10°C. Temperatures above 11°C will kill the eggs.
Importance
Brook Trout are very popular sport fish in eastern Canada caught on lures, live baits and flies. These trout are easier to catch than Brown Trout and take a wider range of lures. They fight well but are often quite small and do not leap spectacularly like some Salmon Family members do. Trout taken on baited hooks and returned to the water show 14 times the death rate of those taken on flies. Anglers wishing to conserve stocks or in search only of trophy fish are best advised to use flies. Hatcheries stock various waters with this trout, sometimes dropping them into lakes from planes. Restocking in the neighbourhood of the NCR on the Québec side in the 1930s suffered from poaching in quantity of these fish (McCuaig, 1935). The ready availability of this salmonid has made it a useful experimental fish for various physiological, biochemical, toxicological and other studies. Some reared stocks however show deformed or lost fins and distorted mouths. The possibility of "sea-ranching" Brook Trout in Atlantic Canada has been explored by releasing stream trout in estuarine areas to improve angling. In freshwater their preference for cool and clear water makes them susceptible to loss if waters are dammed, channelised and polluted or if banks are eroded deforested and overgrazed.
Lake Trout / Touladi
Salvelinus namaycush (Walbaum, 1792)
Taxonomy
Other common names include Lake Charr; Great Lakes, Forktail, Mackinaw, Salmon, Fat Lake, Grey or Mountain Trout; Laker, Landlocked Salmon, Siscowet, Taque, Togue, Truite de lac, Truite grise, Omble gris, Namaycush, Isok, Ihok, Nauktoq, Näluarryuk, Isuuq, Isuuraaryok, Isuuqiq, Siuktuuk, Sigguayaq, Ilortoq, Ivitaruk, Iqluq, and others. A Lake Trout and Brook Trout artificial hybrid is known as Splake, and is often stocked in lakes.
Key Characters
This species is distinguished by having 116-138 lateral line scales, 8-10 principal anal fin rays, teeth on the anterior end only of the vomer bone in the roof of the mouth, pectoral, pelvic and anal fins with a white leading edge but no black contrasting bar behind, head, flank, dorsal, adipose and caudal fin spots white to cream but never red, and the caudal fin has a deep fork.
Description
Dorsal fin principal rays 8-10, pectoral rays 12-17 and pelvic rays 8-11. Pyloric caeca 81-210 and gill rakers 16-26. Males and females have tiny nuptial tubercles around the anus.
Colour
Overall colour is light to dark green, dark olive, brown, grey or black with some fish very silvery masking spots. The back may have pale grey worm-tracks or vermiculations. The belly is dirty white to yellowish. Pectoral, pelvic and anal fins orange. Breeding males develop a black, flank stripe and more reddish brown body and reddish lower fins. The jaw tips and roof of the mouth become whitish. Young have 5-12 parr marks with the spaces between equal or greater in width. Fins are clear except dark bars develop on the dorsal fin of larger parr. Faber (1985a) illustrates a larva.
Size
Reaches 131.0 cm standard length and 46.3 kg. The world, all-tackle angling record was caught in Great Bear Lake on 8 August 1970 by Larry Daunis and weighed 29.48 kg. The Ontario record as of the year 2000 weighed 28.6 kg. Small (1883) reports fish of close to 30 lbs (13.6 kg) from the NCR near Buckingham and a 24 inch fish is recorded from Lac Paquin (Ottawa Sun, 10 June2004).
Distribution
Found from Labrador and Nova Scotia west to British Columbia and Alaska (but absent from most of the Hudson Bay lowlands, southern Saskatchewan, southeastern Alberta and southwestern B.C.), north to Banks, Victoria and Baffin islands (but rare on the last) and south to Idaho and Pennsylvania. Also introduced in the U.S.A., Europe, South America and New Zealand. It was introduced to Mississippi Lake in 1886 (Brown, 1984). Walker (1905) records the introduction of 30,000 young salmon trout, presumably this species, in "Meaches lake" (Meech or Meach Lake). Rubec (1975) records this species from lakes Lapêche, Philippe, Mousseau and Meach in the late 1800s, surviving in only Meach Lake.
Origin
Dumont (1982) considers populations in lakes of southern Québec to be relicts of anadromous populations living in the Champlain Sea, an arm of the Atlantic Ocean 11,900 years ago. These populations would have dispersed during the first phases of the marine invasion that followed the retreat of the glaciers. McAllister and Coad (1975) consider that this species survived glaciation in either an Atlantic or Mississippian refugium or perhaps both, as do Mandrak and Crossman (1992).
Habitat
Lake Trout are found, naturally enough, in lakes where they are solitary but may also be found in some northern rivers and rarely in brackish water. They prefer larger lakes, >100 ha, deeper than 12-14 m with pH>5.5. Southern populations are found only in deep lakes where they can retreat to cooler depths in summer since they prefer waters about 10°C, range 8-15°C. Distribution of this species is restricted by changes in the environment and interspecific competition, coupled with pollution and introduction of competitors (Dumont, 1982).
Age and Growth
Life span may exceed 53 years with maturity attained at 4-7 years in the south and 8-22 years in northern areas like Great Bear and Great Slave lakes. Growth varies over the wide range of this species, generally being slower in the north. Dymond (1939) notes that this species grow to a larger size when Cisco are present as food.
Food
Food includes plankton, sponges, aquatic insects, terrestrial insects when abundant, crustaceans, fishes including their own species and occasionally small mammals such as mice and shrews, and confused yellow warblers which fell onto the water surface in fog. The diet emphasis varies with habitat and plankton feeders grow more slowly than those which eat mostly fish. In smaller lakes fish may not be readily available in the cooler depths and such plankton as opossum shrimps are the mainstay. Opossum shrimps are 82-95% of the food volume of young (up to age 2) trout in Lake Superior but trout over 40 cm long have a diet which is 94% fish. Ciscoes are the most important fish in Lake Trout diet but sculpins are particularly important to younger trout. A wide variety of other fishes are taken as opportunity permits. Alevins in Heney Lake, north of the NCR, feed on the crustaceans Mysis relicta and Pontoporeia affinis (van Vliet and Qadri, 1970).
Reproduction
Spawning occurs from August to December, principally in October and early November in the Outaouais (Chabot, 1981d), and earlier in the north than the south. Temperatures are about 8-11°C (or 8.9-13.9°C after Thellen (1994), also for the Outaouais) and spawning occurs between 1900 and 2200 hours in the dark. Spawning is triggered, at least in part, by sudden temperature drops, cloud cover and onshore winds. Spawning lasts about 2-38 days at any one site, 7-38, mean 16, days in the Outaouais (Chabot, 1981d). Up to 18,051, 6.0 mm diameter eggs are shed over boulders, rubble, or clam shell beds, falling into crevices in moderately deep (to 60 m) to very shallow (a few centimetres) water. In the Outaouais, spawning occurs in depths less than 1 m and fish have been observed spawning with their backs out of the water (Chabot, 1981d). There is no redd construction unlike other salmons, trouts and chars. The spawning ground is cleaned with body or tail brushes or by rubbing with the snout. Most fish spawn in lakes but rarely river spawning is reported. The female spawns with 1-2 males or a group of males and females spawn together. The male nudges or nips the female, presses against her flank with the vents close together, the male erects his dorsal fin and they gape and quiver. Some populations home to spawning sites and disperse over 160 km, to return again in subsequent years. Spawning intervals for females vary, every third year in Great Bear Lake and every second year in Great Slave Lake. Eggs hatch in February to June depending on latitude and larvae are 15.0-16.0 mm long. Alevins have been noted in Lac David, Gatineau on 4 February (Chabot, 1981d). They leave the gravel at 20.0-25.0 mm as juveniles and swim near the lake bottom.
Importance
Sea Lampreys have caused a major decline in trout of the upper Great Lakes, having gained access via canals. DDT was another major factor in decline of this species, causing embryo death. Lake Trout also accumulate PCBs and other toxic chemicals and in 1976 Wisconsin banned human consumption of this fish from Lake Michigan. Overfishing has also caused a decline and natural populations of Lake Trout are extinct in lakes Ontario, Erie and Michigan and only 2 small, remnant stocks survive in Lake Huron. Lake Trout are now a small part of the Great Lakes sport fishery, having been replaced by introduced Rainbow Trout and Pacific salmons. Stocking of Lake Trout and lamprey control by chemicals have enabled Lake Superior trout to recover to some degree. Lake Trout are an important sport fish taken with flies or lures in cool seasons or by deep trolling in summer. Whole or cut fish can be used as bait when bottom fishing. Northern trout remain in cool, near-surface waters even in summer and deep trolling is not required. They can then be caught by spinning or on bucktails and streamers. Trout are also caught through the ice. The Northwest Territories have several expensive, trophy, sport fisheries requiring float plane transport to reach. They must be carefully managed to avoid over-exploitation by rod-and-line fishing. Catch-and-release is favoured with catch limits and fishery zones. Stockings in some Ontario lakes in efforts to improve catches over the natural return have been shown to be of no help. Limitations on take and shortened fishing seasons are probably the best way to maintain a sport fishery. There is a major planting programme in the Great Lakes with about 9 million yearlings being released annually. The flesh is white, pink or orange-red and is very tasty. Commercial catches in some lakes are taken with gillnets and are particularly important in the Northwest Territories. Lake Trout and Lake Whitefish make up 95% of the total catch there. The total Canadian catch in 1988 was 1050 tonnes.
Dymond (1939) records catches from 1881 onward in the general vicinity of the NCR but trends cannot be determined as fisheries data is recorded from different areas at different dates (and may include Brook Trout). For example, in 1881 the catch was 154,000 lbs (69,916 kg) in the Upper Ottawa and Gatineau lakes division, the highest recorded, in 1896 the catch from the Gatineau lakes was 98,100 lbs (44,537 kg), while in the Ottawa River from Carillon to Pontiac in Québec in 1898 the catch was 650 lbs (295 kg).
A Lake Trout fossil of Quaternary age (ca. 10,000 years ago) assigned to this species has been found in a clay nodule from Besserer's Springs (near Green Creek at Hiawatha Park in the NCR) (Gruchy, 1968; McAllister et al., 1981; Harington, 1972; 1983; McAllister et al., 1987
© Brian W. Coad (www.briancoad.com)