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A Review of the Literature on the Upstream Migration of Adult Salmonids

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A Review of the Literature on the Upstream Migration of Adult Salmonids J. Fish Biol. (1969) 1, 85-136 A Review of the Literature on the Upstream Migration of Adult Salmonids J. W. BANKS Department of Zoology, University of Liverpool, Liverpool, England (Received 3 December 2968) CONTENTS I. Introduction .. .. .. 86 11. The homing of salmon and the formation of local races . .. .. 86 111. Methods of observing the upstream migration of salmonid fishes . .. 89 IV. Migration in uncontrolled conditions . .. .. .. .. 91 1. Atlantic salmon Saho salar Linn. .. .. .. .. 91 2. Sea trout Salmo trutta Linn. .. .. .. .. .. 93 3. Brown trout Salmo trutfu Linn. .. .. .. .. .. 93 4. Rainbow trout Salmo gairdneri Richardson . .. .. .. 96 5. Sockeye, red, blueback or Kokanee salmon Oncorhynchus nerka (Walbaum) 97 6. Pink or humpback salmon Oncorhynchus gorbuscha (Walbaum) .. 98 7. Chum or dog salmon Oncorhynchus keta (Walbaum) . .. .. 102 V. Migration under controlled conditions . .. .. .. .. 102 1. Atlantic salmon Saho salar Linn. .. .. .. .. 102 2. Sea trout Salmo trutta Linn. .. .. .. .. *. 112 3. Steelhead, rainbow trout Salmo gairdneri Richardson .. .. 112 4. Sockeye salmon Oncorhynchus nerka (Walbaum) . .. .. 112 5. Chinook or spring salmon Oncorhynchus tshawytscha (Walbaum) .. 114 VI. Discussion .. .. .. .. .. .. .. 114 1. Rate of Flow . *. .. .. .. .. .. 114 2. Problems of flow at dams, diversions and fish passes .. .. 118 3. Temperature . *. .. .. .. .. .. 120 4. Water quality . .. .. .. .. .. .. 123 5. General weather, wind and tide . .. .. .. .. 124 6. Light intensity . .. .. .. .. .. .. 126 VII. The physiology of salmonids and the control of migration .. .. .. 129 VIII. Summary and conclusions . .. .. .. .. .. 130 References . .. .. .. .. .. .. .. 132 85 A 86 J. W. BANKS I. INTRODUCTION The literature on salmonids abounds with instances, both anecdotal and precise, which show that spawning migrations are associated with increased water flows. Doubtless this fact has been a part of fishermen's lore for a very long time. Some early references occur in Day (1887), and even then it was realized that the relationship was not straightforward, other environmental influences being involved. Subsequent work has shown that many factors like water and air temperature, turbid- ity, atmospheric pressure, cloud cover, pH and variations in concentrations of many dissolved ions are associated with the rate of water discharge in a more or less direct fashion. It has been more difficult in individual cases to determine the significance to the salmon of the many possible stimuli available from these covariable factors. Much of the work which has been done has involved the taking of measurements of' only a few of the parameters which are partially covariant with changes in the rate of water discharge, and therefore the conclusions reached in such work involve the assumption that other factors are not important. Where the possible importance of other covariable factors has been acknowledged, it is only rarely that the amount and statistical significance of any covariability has been considered. This is not surprising as the variables are both complex and often difficult to measure. The result is that at the present time we are still only in a position to say that in general the rate of water discharge is an important, if not dominant, stimulus to the upstream migration of salmon, but in particular instances this effect can be profoundly modified by other environmental influences. The nature of these other influences, and their effect, varies widely from place to place, with season, and with different salmonid species. Reviews of the factors controlling the migration and homing of salmon were published by Chidester (1924) and Scheer (1939). More recently, various aspects of the subject have been touched on in reviews by Hoar (1953), Pyefinch (1955), Jones (1959) and Hasler (1965), but no detailed review of this subject seems to have been attempted in recent years. The presentation of the observations on salmonid migra- tion, which forms the bulk of this review, has been made under two headings. First under uncontrolled conditions, and second under controlled conditions. For the sake of convenience all work relating to situations where an artificial vertical ascent has been imposed on the fish will be considered under the second heading. In practice there is little difference between the ascent into a trap on a small weir, and passage through the trap in a counting fence, but this is the most convenient place to draw the line. Under these two headings the work has been presented by species, and where a publication has dealt with more than one these are mentioned separately, unless the reference is a minor one, when it has often been more convenient to place it together with the work on the species with which the paper is primarily concerned. The information which is presented in the sections on migration in uncontrolled and controlled conditions is re-examined in the discussion in a number of general cate- gories of factors affecting salmonid migration. This section also incorporates directly some material from papers, mainly on experimental work, which do not fit easily into either of the two main sections. II. THE HOMING OF SALMON AND THE FORMATION OF LOCAL RACES The genera SaImo and Oncorhynchus are primarily inhabitants of the north temper- ate regions surrounding the Atlantic and Pacific Oceans, respectively. Anadromous UPSTREAM MIGRATION OF ADULT SALMONIDS 87 species migrate into rivers and streams wherever suitable spawning grounds are available, and from which they have not been excluded by natural barriers, artificial obstructions, or by pollution. In this vast area, salmon habitually spawn in cold, well-oxygenated stream waters. The current speed and size of stones in the redds varies with species and size of fish. Sockeye salmon occasionally spawn on the shores of lakes, particularly where ground water seepage is coming up through the gravel, but may also spawn in areas of ground water seepage in streams. Some brown trout and rainbow trout inhabit streams and lakes, but similarly run up to cold headwaters to spawn. Others feed in the sea or estuaries, but again spawn in gravels in the streams. Suitable gravels and conditions of oxygenation are most frequently met with in the headwaters of the river system, but in mountainous areas redds may be found within a very short distance of the head of the tide. As a consequence the range of environments encountered by different individuals, even within the same species, will vary enormously. The spawning migration of some will be entirely in fresh water, others will need to make much greater physiological adjustments in moving in from the sea. The journey in fresh water may be anything from a few hundred yards to more than a thousand miles. It may involve nothing more arduous than a few stickles between pools, or can involve the passage of rapids and waterfalls, with vertical jumps of more than two metres, or fish passes on high dams where maximum swimming speeds of up to 34 m/sec may be needed repeatedly. The stream may be subject to violent fluctuations in discharge, or evenly regulated by dams and lakes. The gradients may be low or steep. Many environmental factors will vary in relation to each other, both throughout the range of the species and probably within the length of the stream. Under these conditions it is not surprising to find that there has been a tendency to develop local ‘ races ’ within each species, especially in those which are anadromous. There has been a good deal of argument about the degree to which salmonids form local ‘ races ’ by persistence in returning to their natal stream. Differences of opinion are now seen to have been the result of too much generalization from particular instances. Ward (1939) seeks to reduce the parent stream theory to a statement that it may be the rule, and that the evidence is insufficient to do more than indicate a general compliance with this practice. Much evidence, both direct and indirect, has, however, been cited in support of the stream homing hypothesis, and its concomitant, the building up of genetically different local races. Menzies (1939) cites an instance in which two neighbouring streams, apparently very similar, have a constant difference in mean length for the salmon returning to them. Dvinin (1952) cites the case of two small rivers in Kamchatka whose mouths are 5 km apart; both formerly had spawn- ing populations of between 5000 and 12,000 chum salmon. One river was fished out many years ago and has never re-established a spawning population, while the population on the other continues to flourish. Jones (1959) described the situation in the River Bann in Ulster where destruction of the spawning grounds of the large fish in a particular reach of the main river has resulted in a steady drop of the average size of fish in the catch; Went (1946) described similar results in the River Shannon, after the hydro-electric scheme. Many years of work on the Fraser River by biologists has shown that different spawning populations of sockeye salmon are maintained in each of the major tributaries, The timing of entry from the sea varies with each tributary’s population and can be quite accurately predicted. In each instance the 88 J. W. BANKS time of entry affords the population just enough time to reach the spawning grounds before the gonads ripen. When abnormal conditions intervene, and delays occur, the effects on spawning success can be disastrous. The situation in the Fraser River was summarized by Thompson (1951). The individuals in each group of migrants are made numerically important by their survival rates and form a recognizable entity known by its home stream to the extent that this stream is itself a unit. The fact of continued favourable survival rates in their particular chain of time and space linked environments gives these groups persistence and individuality. The late race of sockeye into Adams River has maintained its numerical dominance in the cyclic year in which it occurs.
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