Habitat Requirements for Lake Rearing Sockeye Salmon Populations General Life History The distribution of lake rearing sockeye salmon populations is based on the requirement of a nursery lake for the freshwater rearing of juveniles. The largest populations in British Columbia are located in the Fraser and Skeena river drainages where tributary systems are often connected to large lakes. Other large populations exist in the Nass River system in northern BC, the Somass River system on the west coast of Vancouver Island and in both Rivers and Smith inlets in central BC. Smaller stocks spawn in systems with nursery lakes throughout the province. The majority of mature adult sockeye return to their river of origin to spawn during the summer or autumn as four year old fish that have spent two years in the ocean. The exceptions to this pattern include small fish (primarily males referred to as jacks) that return to spawn after one year of ocean residence and those fish that remain in the ocean for a third year. Migration distances from the ocean to freshwater spawning grounds range from a few kilometers to over 1,000 km. The initiation of the spawning migration for each unique stock is timed to capitalize on ideal conditions in the freshwater habitat that will ensure appropriate egg development. In most cases, stock specific migration timing is remarkably consistent between years. When mature sockeye salmon reach the spawning grounds, the female selects a nest site and begins digging a pit referred to as a redd, where eggs will be deposited. Most spawning areas are located in rivers or streams that are tributary to lakes; however, some stocks spawn in lake outlet streams while others spawn along the lake shore. The digging process removes sand, silt and fine gravel from the nest site creating a favourable environment for incubation of the eggs. Once the nest is complete, the female deposits the eggs which are fertilized by one or more males and then moves to the area immediately upstream of the nest and begins digging another pit. The material removed by this digging action covers the fertilized eggs to protect them from predation and from being washed away by the scouring action of the river or stream. This process may be repeated several times resulting in multiple nests containing eggs from one female. The length of time required for the eggs to incubate depends to a large extent on water temperature. In general, the lower the water temperature, the longer the incubation period required. Upon hatching, the juvenile sockeye (called alevins) move downward in the gravel varying distances depending on gravel size. At this point, the young fish have an attached yolk sac that provides the required nutrition. Towards the end of incubation, alevins move up through the gravel to emerge as fry. This occurs after nightfall between April and early June and generally coincides with the complete absorption of the yolk sac. In streams that are tributary to lakes, fry swim downstream toward the nursery lake often moving passively with the current. In lake outlet spawning areas, newly emerged fry move toward the stream bank where flow is reduced and swim upstream against the current toward the nursery lake. Timing of lake entry by fry is determined by many factors that ensure juveniles encounter the best possible conditions for feeding, predator avoidance and growth. When juvenile sockeye enter the lake they spend the first few months close to the shoreline where they feed on larval and adult insects. Once they have moved to deeper offshore waters, juveniles feed on a variety of zooplankton species that are plentiful in the upper water column. In this environment, they exhibit a complex pattern of daily vertical migration that varies between lakes and changes seasonally. In general, they occupy surface waters at night moving down to deeper waters during the day. This pattern of movement serves to maximize feeding opportunities while minimizing exposure to predators. It also ensures that they receive the greatest benefit from food ingested by controlling metabolism and maximizing growth. In the vast majority of cases, juvenile sockeye from lake rearing populations in BC leave the nursery lake after one year of residence. In some less productive lake environments, two years of lake rearing occur before migration to the ocean. Juvenile sockeye typically leave the nursery lake as water temperatures increase in the spring. A process called smoltification begins to occur in which both physical and behavioral changes take place in preparation for life at sea. These include becoming more silver in colour, slimmer and more streamlined in shape and undergoing the physiological changes required for tolerance to seawater. During the downstream migration, the smolts travel in schools and migrate actively (faster than the current in slow moving water). Most movement takes place at night in order to reduce the risk of exposure to predators. Sockeye smolts exhibit a variety of behaviors upon reaching saltwater. While some may reside for a short period in the river estuary or in nearshore waters adjacent to the river mouth most disperse into coastal waters more quickly. Movement to offshore areas begins with a general migration to the northwest along the coast. During this time, sockeye feed on a variety of organisms preferring zooplankton, insects and fish larvae. By July, they tend to from a band approximately 40 km offshore that extends from southwest coast of Vancouver Island to the Gulf of Alaska. Movement continues in a northwesterly direction at a rate of approximately 18 kilometers per day. Once in the Gulf of Alaska, sockeye begin to move further offshore to open ocean waters. This typically occurs in late autumn or early winter. Patterns of movement in the ocean are complex and are influenced by many factors including temperature, currents and food distribution. During this period, sockeye are widely distributed throughout the North Pacific where they feed on large zooplankton, small fish and squid. For most sockeye salmon, sexual maturation begins during their second year of ocean residence when they begin an accelerated spring-early summer migration back to the river of origin. Habitat Requirements by Life History Stage The health of all Pacific salmon is closely linked to the availability of productive freshwater, coastal and marine environments. Lake rearing sockeye populations require quality spawning, incubation, lake rearing and ocean habitats to thrive and remain productive. The loss of quality habitat in any of these vital areas will have a negative impact on stocks. Healthy habitat is challenged by human competition for accessible land and fresh water, for ocean spaces and for the interconnecting estuarine and coastal areas. In order to maintain productive lake rearing sockeye populations throughout the Pacific coastal region, these important habitats must be protected. Spawning Adult sockeye salmon require unimpeded access to their home spawning grounds in order to successfully reproduce. Features such as dams, debris jams, waterfalls, or rock/mud slides that block upstream migration can limit access to spawning areas and impact production. Also, if conditions such as high water temperature or extreme high or low flows are encountered when spawners arrive at their river or stream of origin, fish often mill about in the vicinity of the river mouth, waiting for conditions to improve. Such delays in river entry can have a detrimental affect on survival and on spawning success as fish are exposed to predation from marine mammals and, since feeding has stopped in preparation for spawning, vital energy reserves are used up. This is of particular concern for those sockeye populations with the longest spawning migrations. In years when river conditions present a difficult migration environment due to elevated water temperature or extreme flows, a high percentage of some stocks will run out of energy and die before they reach the spawning grounds. For some large Fraser River sockeye populations, rates of such pre-spawning mortality have been very high in some years. While it may not possible to prevent these conditions in all situations, every effort must be made to ensure that river temperatures and flows are not negatively impacted by human activity. In those years when adverse migration conditions are anticipated, it may be necessary for fishery managers to reduce the catch of stocks undertaking long spawning migrations to ensure that an appropriate number of fish reach the spawning grounds. Sockeye salmon exhibit the greatest diversity in selection of spawning habitat among the Pacific salmon. They may spawn in inlet or outlet tributaries of a nursery lake, in suitable habitat between lakes, along lakeshores, or on submerged beaches. In all of these habitats, sockeye salmon require extensive, high quality spawning gravels for optimum production. Areas selected by female sockeye for redd construction tend to be located where the substrate is small enough to be moved by the fish and large enough to allow good intragravel water flow to the incubating eggs and developing alevins. This ensures that the environment in the nest is supplied with a constant flow of water to deliver dissolved oxygen and to remove waste. Those sockeye that spawn on lake shores require access to undisturbed shorelines with quality gravel and clean, upwelling groundwater to ensure high survivals. Field observations indicate that while many spawning areas are presently under-utilized, other areas appear to be filled to capacity in some years. This observation suggests that a lack of prime spawning areas can limit sockeye production in some systems as spawners may be forced to build redds in less suitable locations or on top of previously constructed redds resulting in reduced overall production. Incubation Studies indicate that at a constant temperature, sockeye salmon have the longest incubation period of the five pacific salmon species. During incubation, eggs and alevins require a stable environment with an uninterrupted supply of clean, oxygen rich water. The percentage of eggs and alevins that survive the incubation phase depends to a large extent on stream and stream bed conditions. Flooding during the winter can result in a large amount of gravel movement which reduces survival by causing eggs or alevins to be exposed and swept downstream. Also, silt loads associated with flooding may hinder water circulation through the redd reducing available oxygen to harmful or lethal levels. Studies have shown that a higher proportion of fine sediment in the spawning gravel reduces survival and results in smaller emergent fry. Activities that contribute to flooding or siltation of incubation areas can magnify the impact of extreme winter conditions and reduce survival to emergence. Juvenile Rearing After emerging from the gravel, sockeye fry migrate directly to the nursery lake. During this migration the fry are highly vulnerable to predators such as birds and large fish. Loss due to predation during this stage varies but has been estimated to range from 15 to 90%. The availability of appropriate cover can limit exposure to predators and reduce mortality on fry during migration to the nursery lake. Upon reaching the lake, sockeye juveniles spend the first few weeks in the near shore area where they feed primarily on insect larvae. Shoreline habitats must remain favourable to the production of insect populations in order to sustain juvenile sockeye during this life stage. With the growing settlement of lakeshore areas for recreational purposes, it is important that shoreline areas remain in a natural state in order to maintain a productive habitat for juvenile sockeye. The longest period of freshwater residence for lake rearing sockeye salmon is spent in the offshore waters of the nursery lake and juveniles require quality, productive offshore habitat to thrive. During this period, sockeye juveniles employ various strategies to maximize feeding while reducing exposure to predation which include vertical migrations and schooling behaviors. In some lakes, species that are predators on juvenile sockeye have been enhanced or introduced for recreational purposes. Such practices should be avoided in sockeye nursery lakes and every effort made to maintain quality offshore lake habitats for juvenile sockeye. There are many sources and types of pollutants that can affect the nursery lake habitat including waste water, pesticides, toxic chemicals, petroleum products and organic compounds. These substances can reduce the quality of the habitat for juvenile sockeye and for the aquatic organisms on which they feed. All efforts must be made to minimize the introduction of pollutants into sockeye nursery lakes. Sockeye salmon smolts transiting from nursery lakes to the ocean require open migration corridors that are free from obstacles that could delay or prevent passage. Since predation can result in high mortality rates during this phase, it is important that migration paths have cover available to smolts. Appropriate cover includes healthy, abundant streamside vegetation as well as woody debris and large substrate material. Ocean Phase The transition zone from freshwater to saltwater provided by the river estuary is an important habitat for all transiting juvenile salmon. Sockeye smolts require a natural, productive estuary that can provide the necessary food resources while offering refuge from fish and bird predators. The time spent in the estuary helps transiting sockeye smolts adjust to a saltwater environment that is dramatically different from the freshwater habitat that has been their home during the first part of their life. As they move into the ocean, another critical life history phase begins. Research suggests that up to 90% of the natural ocean mortality occurs during the first four months that sockeye are at sea. Since survival is generally size-selective, favourable near-shore ocean productivity is important as it can result in faster growth and a shorter time to reach the size required for increased survival. As a result, coastal ocean productivity is a key habitat component that impacts the survival of sockeye salmon. Sockeye travel far during the marine phase of their life reaching the northern, nutrientrich waters of Alaska and the Arctic. The distribution of sockeye in offshore waters is dependent on ocean environmental conditions and on food availability. While detailed migration patterns and other aspects of their marine ecology remain poorly understood, ocean residence is recognized as a very important component of the life cycle of all Pacific salmon. During their time at sea, sockeye migrate varying distances while increasing in size and acquiring the energy reserves required for reproduction. While distribution patterns vary between years and stocks, all sockeye utilize coastal and off shore habitats during a period of rapid growth that is critical to reproductive success. References Anon. 1985. Alaska habitat management guide, southcentral region, Volume 1: Life histories and habitat requirements of fish and wildlife. Sockeye salmon (Oncorhynchus nerka). Alaska Dept. Fish. 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