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Atlantic Salmon Brown Trout and Arctic Charr: a Review of Aspects Of Ecology of Freshwater Fish 2003: 12: 1–59 Copyright # Blackwell Munksgaard 2003 Printed in Denmark Á All rights reserved Review Atlantic salmon Salmo salar L., brown trout Salmo trutta L. and Arctic charr Salvelinus alpinus (L.): a review of aspects of their life histories Klemetsen A, Amundsen P-A, Dempson JB, Jonsson B, Jonsson A. Klemetsen1, P.-A. Amundsen1,J.B. N, O’Connell MF, Mortensen E. Atlantic salmon Salmo salar L., Dempson2, B. Jonsson3, N. Jonsson3, brown trout Salmo trutta L. and Arctic charr Salvelinus alpinus M. F. O’Connell2, E. Mortensen4 (L.): a review of aspects of their life histories. 1 # Norwegian College of Fishery Science, University of Ecology of Freshwater Fish 2003: 12: 1–59. Blackwell TromsÖ, Breivika, N-9037 TromsÖ, Norway, Munksgaard, 2003 2Fisheries and Oceans Canada, Science, Oceans and Environment Branch, PO Box 5667, St. John’s, Abstract – Among the species in the family Salmonidae, those Newfoundland, Canada A1C 5X1, 3Norwegian represented by the genera Salmo, Salvelinus, and Oncorhynchus Institute of Nature Research, Dronningensgt 13, PO (subfamily Salmoninae) are the most studied. Here, various aspects of Box 736 Sentrum, N-0105 Oslo, Norway, 4ÒrnsÖvej phenotypic and life-history variation of Atlantic salmon Salmo salar L., 15, DK-8600 Silkeborg, Denmark brown trout Salmo trutta L., and Arctic charr Salvelinus alpinus (L.) are reviewed. While many strategies and tactics are commonly used by these species, there are also differences in their ecology and population dynamics that result in a variety of interesting and diverse topics that are challenging for future research. Atlantic salmon display considerable phenotypic plasticity and variability in life-history characters ranging from fully freshwater resident forms, where females can mature at approximately 10 cm in length, to anadromous populations characterised by 3–5 sea-winter (5SW) salmon. Even within simple 1SW populations, 20 or more spawning life-history types can be identified. Juveniles in freshwater can use both fluvial and lacustrine habitats for rearing, and while most smolts migrate to sea during the spring, fall migrations occur in some populations. At sea, some salmon undertake extensive oceanic migrations while other populations stay within the geographical confines of areas such as the Baltic Sea. At the other extreme are those that reside in estuaries and return to freshwater to spawn after spending only a few months at sea. The review of information on the diversity of life-history forms is related to conservation aspects associated with Atlantic salmon populations and current trends in abundance and survival. Brown trout is indigenous to Europe, North Africa and western Asia, but was introduced into at least Key words: Arctic charr; Atlantic salmon; brown 24 countries outside Europe and now has a world-wide distribution. It trout; cannibalism; feeding; freshwater residence; exploits both fresh and salt waters for feeding and spawning (brackish), habitat; life history; migration; phenotypic and populations are often partially migratory. One part of the plasticity; phylogeography; polymorphism; population leaves and feeds elsewhere, while another part stays as research challenges residents. In large, complex systems, the species is polymorphic with A. Klemetsen, Norwegian College of Fishery Science, different size morphs in the various parts of the habitat. Brown trout University of TromsÖ, Breivika, N-9037 TromsÖ, feed close to the surface and near shore, but large individuals may move Norway; e-mail: [email protected] far offshore. The species exhibits ontogenetic niche shifts partly related Dempson and O’Connell reviewed the Atlantic to size and partly to developmental rate. They switch when the amount salmon, Jonsson and Jonsson the brown trout and of surplus energy available for growth becomes small with fast growers Amundsen and Klemetsen the Arctic charr. being younger and smaller fish than slow growers. Brown trout is an Mortensen initiated the review and gave opportunistic carnivore, but individuals specialise at least temporarily constructive criticism to the manuscript. on particular food items; insect larvae are important for the young in Accepted for publication September 22, 2002 1 Klemetsen et al. streams, while littoral epibenthos in lakes and fish are most important for large trout. The sexes differ in resource use and size. Females are more inclined than males to become migratory and feed in pelagic waters. Males exploit running water, near-shore and surface waters more than females. Therefore, females feed more on zooplankton and exhibit a more uniform phenotype than males. The Arctic charr is the northernmost freshwater fish on earth, with a circumpolar distribution in the Holarctic that matches the last glaciation. Recent mtDNA studies indicate that there are five phylogeographic lineages (Atlantic, Arctic, Bering, Siberian and Acadian) that may be of Pleistocene origin. Phenotypic expression and ecology are more variable in charr than in most fish. Weights at maturation range from 3 g to 12 kg. Population differences in morphology and coloration are large and can have some genetic basis. Charr live in streams, at sea and in all habitats of oligotrophic lakes, including very deep areas. Ontogenetic habitat shifts between lacustrine habitats are common. The charr feed on all major prey types of streams, lakes and near-shore marine habitats, but has high niche flexibility in competition. Cannibalism is expressed in several cases, and can be important for developing and maintaining bimodal size distributions. Anadromy is found in the northern part of its range and involves about 40, but sometimes more days in the sea. All charr overwinter in freshwater. Partial migration is common, but the degree of anadromy varies greatly among populations. The food at sea includes zooplankton and pelagic fish, but also epibenthos. Polymorphism and sympatric morphs are much studied. As a prominent fish of glaciated lakes, charr is an important species for studying ecological speciation by the combination of field studies and experiments, particularly in the fields of morphometric heterochrony and comparative behaviour. Un resumen en espan˜ ol se incluye detra´ s del texto principal de este artı´ culo. Introduction across the Atlantic as well as to many other parts of the world (Elliott 1994). The salmon is found in river systems on both sides of the Atlantic and ‘‘There is no group or family of fishes that migrates over most of the northern part of the supplies better materials for the study of the ocean during anadromy (Hansen & Quinn 1998). effects of geographic or physiologic isolation, In contrast to the other two, the charr is a cir- or which presents more curious and interest- cumpolar species. It is also the northernmost of all ing facts in their life histories than do the vari- anadromous and freshwater fishes. ous species of Salmonidae.’’ (Evermann 1925) These three species range among the best-stu- The salmonid subfamily Salmoninae comprises died fish species in the world. The literature on all about 30 species of fish in seven genera (Behnke three is very large, as is evident for instance from 1991; Nelson 1994) of which Salmo and Salvelinus the six volumes of references to the Arctic charr (along with Oncorhyncus) are the best studied. In complex up to 1996 issued by Canada Fisheries the present contribution, we review aspects of the and Oceans (Marshall 1977, 1981; Marshall & biology of Atlantic salmon (Salmo salar), brown Layton 1985; Heuring et al. 1991; Marshall et al. trout (Salmo trutta) and Arctic charr (Salvelinus 1994, 1998). Also, there are several recent books alpinus). For brevity, we will usually refer to them (Balon 1980; Mills 1989, 1993; Shearer 1992; as salmon, trout and charr in the text. The three Elliott 1994), reviews (Johnson 1980; Jonsson & species have wide but different natural distribu- Jonsson 2001) and symposium proceedings tions that overlap extensively in Europe. The (Johnson & Burns 1984; Kawanabe et al. 1989; salmon and trout are both Atlantic species but Klemetsen et al. 1995; Mather et al. 1998; Pre´ vost the trout occurred natively only on the eastern & Chaput 2001; Magnan et al. 2002) that cover side of the ocean. Man has, however, introduced it important parts of the ecology and population 2 Salmonid life histories dynamics of each of the three species considered Furthermore, the structure of populations is made here. We therefore will not necessarily give exten- more complex by the presence of sexually mature sive reviews of all, or even most sides of their parr (e.g., Myers et al. 1986). In most systems, biology. Rather, we focus on aspects that we find mature parr consist of males only, although mat- interesting, diverse, currently topical, and also ure female parr have also been reported in some challenging for future research. As it turned instances (e.g., Power 1969; Prouzet 1981). out, these aspects proved to be quite different The information published on Atlantic salmon for the three species. is immense with an abundance of individual papers, books, and journal supplements, as acknowledged ´ Atlantic salmon, Salmo salar L. recently by Chaput & Prevost (2001), devoted to all aspects of its ecology. In recent years, themes Introduction associated with many publications (e.g., Mills 1993, 2000; Hutchinson 1997; Whoriskey & Whe- Atlantic salmon occur naturally along both east lan 1997; Dempson et al. 1998; Mather et al. 1998; and west coasts of the North Atlantic Ocean Jonsson & Waples 1999; Pre´ vost & Chaput 2001) where it exists in both anadromous and non- often relate, in whole or in part, to the reality that anadromous freshwater resident forms. In the north- numerous salmon populations are in decline. west Atlantic, salmon occur from approximately Here, we present aspects of phenotypic and life- the Connecticut River in the south to Ungava Bay history variation in salmon, in particular high- in the north, while in the Northeast Atlantic, lighting the inherent variability in life-history salmon range from northern Portugal to rivers forms both within and among populations.
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