Asynchrony in population dynamics of sockeye salmon in southwest Alaska

Ecologists have examined the synchronization of population dynamics across space as a means to understand how populations respond to climate variation. However, response diversity may reflect important variation among local population dynamics driven by population‐specific responses to regional environmental change. We used long‐term data on sockeye salmon Oncorhynchus nerka from pristine watersheds of southwestern Alaska to show that populations spawning in close proximity (<40 km) to one another have a limited degree of synchrony in their dynamics, even after accounting for density‐dependent processes. In fact, the dynamics of local populations of stream‐spawning sockeye salmon were no more coherent than those of stocks at a much coarser resolution across this region of Alaska. We examined four hypotheses to explain the observed patterns of asynchrony among stream‐spawning populations, and found that populations spawning in dissimilar habitats, and using different nursery lakes were less synchronized in their productivity. Similarity in the age structure of spawning adults was less correlated with synchrony in productivity. These results emphasize the importance of maintaining diverse spawning and rearing habitat for the conservation of Pacific salmon, and should guide conservation planning for Pacific salmon populations in regions where natural dynamics have been altered by habitat loss, hatchery practices, and over‐fishing.     

Climate variation is filtered differently among lakes to influence growth of juvenile sockeye salmon in an Alaskan watershed

The physical landscape filters regional climate variation such that the environmental conditions an organism experiences are unique to the characteristics of local habitat features. While it has become apparent that populations may show watershed‐specific responses to changing climate within a geographic region, the population dynamics of Pacific salmon Oncorhynchus spp. suggest that within watershed climate filtering is also important. Growth provides an integrated measure of habitat quality capturing the overall response of individuals to climate as filtered by their habitat and their response to the biological interactions in the ecosystem. We used two different long‐term datasets, scales from returning adults and juvenile length measurements, to assess the response of sockeye salmon O. nerka growth during their juvenile life phase to single and integrated measures of climate within a watershed between 1950 and 2010. Scale growth showed evidence for differences among stocks rearing in different lake habitats within the same Alaska Peninsula watershed. These lakes have substantially different morphometry and showed opposite responses to changes in spring and fall air temperatures. Juvenile length data were also available for one of these stocks and indicated that density effects were relatively weak in contrast to the effects of temperature. While direct measures of juvenile length and measures derived from adult scales showed some similarity in their trends over time, they indicated opposite effects of air temperature. Throughout the range of Pacific salmon, climate change is altering freshwater ecosystems through changes to temperature, precipitation and associated variables. These data suggest that sockeye salmon populations are experiencing climate filtering at a sub‐watershed scale. Maintaining connected, heterogeneous landscapes will therefore likely be important for providing productive habitat for sockeye salmon across a range of climate conditions that they are going to experience under new climate regimes.     

Going with the Flow: Spatial Distributions of Juvenile Coho Salmon Track an Annually Shifting Mosaic of Water Temperature

A critical challenge for ecologists is to understand the functional significance of habitat heterogeneity and connectivity for mobile animals. Here, we explore how a thermo-regulating fish responds to annual variation in the spatial patterning of thermal and trophic resources. In a third-order stream in coastal Alaska, juvenile coho salmon forage on sockeye salmon eggs at night in cold water and then move to warmer water to increase their digestive capacity. We mapped the spatial distributions of water temperature, juvenile coho salmon, and spawning sockeye salmon across a 5-year period during which summer discharge varied by greater than fivefold. In low flow years, warm water (9–12°C) was only available in thalweg (that is, main-channel) habitat at least approximately 400 m upstream of the cooler habitat (3–7°C) where sockeye salmon spawned. In high flow years, the entire stream thalweg was isothermal at 7–8°C, but inundated off-channel areas generated warm habitats (9–12°C) laterally adjacent to the downstream regions where sockeye salmon spawned. The daytime spatial distribution of juvenile coho salmon shifted from headwater thalweg habitats in low flow years, to downstream off-channel habitats in high flow years. In all years, the majority of juvenile coho salmon sampled during the daytime were found in warm habitat units without sockeye salmon present, yet they exhibited diet contents comprised virtually entirely of sockeye salmon eggs. Thus, thermoregulatory movements by coho salmon were able to track an annually shifting mosaic of water temperature. Our results demonstrate how the spatial habitat heterogeneity and connectivity of intact floodplains can in turn buffer aquatic organisms from high levels of temporal variation in habitat conditions and resource abundance.     

Recent local adaptation of sockeye salmon to glacial spawning habitats

Salmonids spawn in highly diverse habitats, exhibit strong genetic population structuring, and can quickly colonize newly created habitats with few founders. Spawning traits often differ among populations, but it is largely unknown if these differences are adaptive or due to genetic drift. To test if sockeye salmon (Oncorhynchus nerka) populations are adapted to glacial, beach, and tributary spawning habitats, we examined variation in heritable phenotypic traits associated with spawning in 13 populations of wild sockeye salmon in Lake Clark, Alaska. These populations were commonly founded between 100 and 400 hundred sockeye salmon generations ago and exhibit low genetic divergence at 11 microsatellite loci (F _ST < 0.024) that is uncorrelated with spawning habitat type. We found that mean P _ST (phenotypic divergence among populations) exceeded neutral F _ST for most phenotypic traits measured, indicating that phenotypic differences among populations could not be explained by genetic drift alone. Phenotypic divergence among populations was associated with spawning habitat differences, but not with neutral genetic divergence. For example, female body color was lighter and egg color was darker in glacial than non-glacial habitats. This may be due to reduced sexual selection for red spawning color in glacial habitats and an apparent trade-off in carotenoid allocation to body and egg color in females. Phenotypic plasticity is an unlikely source of phenotypic differences because Lake Clark sockeye salmon spend nearly all their lives in a common environment. Our data suggest that Lake Clark sockeye salmon populations are adapted to spawning in glacial, beach and tributary habitats and provide the first evidence of a glacial spawning ecotype in salmonids. Glacial spawning habitats are often young (i.e., <200 years old) and ephemeral. Thus, local adaptation of sockeye salmon to glacial habitats appears to have occurred recently.     

Juvenile Salmon Usage of the Skeena River Estuary

Migratory salmon transit estuary habitats on their way out to the ocean but this phase of their life cycle is more poorly understood than other phases. The estuaries of large river systems in particular may support many populations and several species of salmon that originate from throughout the upstream river. The Skeena River of British Columbia, Canada, is a large river system with high salmon population- and species-level diversity. The estuary of the Skeena River is under pressure from industrial development, with two gas liquefaction terminals and a potash loading facility in various stages of environmental review processes, providing motivation for understanding the usage of the estuary by juvenile salmon. We conducted a juvenile salmonid sampling program throughout the Skeena River estuary in 2007 and 2013 to investigate the spatial and temporal distribution of different species and populations of salmon. We captured six species of juvenile anadromous salmonids throughout the estuary in both years, and found that areas proposed for development support some of the highest abundances of some species of salmon. Specifically, the highest abundances of sockeye (both years), Chinook in 2007, and coho salmon in 2013 were captured in areas proposed for development. For example, juvenile sockeye salmon were 2–8 times more abundant in the proposed development areas. Genetic stock assignment demonstrated that the Chinook salmon and most of the sockeye salmon that were captured originated from throughout the Skeena watershed, while some sockeye salmon came from the Nass, Stikine, Southeast Alaska, and coastal systems on the northern and central coasts of British Columbia. These fish support extensive commercial, recreational, and First Nations fisheries throughout the Skeena River and beyond. Our results demonstrate that estuary habitats integrate species and population diversity of salmon, and that if proposed development negatively affects the salmon populations that use the estuary, then numerous fisheries would also be negatively affected.     

Watershed‐scale climate influences productivity of Chinook salmon populations across southcentral Alaska

The ecosystems supporting Pacific salmon (Oncorhynchus spp.) are changing rapidly as a result of climate change and habitat alteration. Understanding how—and how consistently—salmon populations respond to changes at regional and watershed scales has major implications for fisheries management and habitat conservation. Chinook salmon (O. tshawytscha) populations across Alaska have declined over the past decade, resulting in fisheries closures and prolonged impacts to local communities. These declines are associated with large‐scale climate drivers, but uncertainty remains about the role of local conditions (e.g., precipitation, streamflow, and stream temperature) that vary among the watersheds where salmon spawn and rear. We estimated the effects of these and other environmental indicators on the productivity of 15 Chinook salmon populations in the Cook Inlet basin, southcentral Alaska, using a hierarchical Bayesian stock‐recruitment model. Salmon spawning during 2003–2007 produced 57% fewer recruits than the previous long‐term average, leading to declines in adult returns beginning in 2008. These declines were explained in part by density dependence, with reduced population productivity following years of high spawning abundance. Across all populations, productivity declined with increased precipitation during the fall spawning and early incubation period and increased with above‐average precipitation during juvenile rearing. Above‐average stream temperatures during spawning and rearing had variable effects, with negative relationships in many warmer streams and positive relationships in some colder streams. Productivity was also associated with regional indices of streamflow and ocean conditions, with high variability among populations. The cumulative effects of adverse conditions in freshwater, including high spawning abundance, heavy fall rains, and hot, dry summers may have contributed to the recent population declines across the region. Identifying both coherent and differential responses to environmental change underscores the importance of targeted, watershed‐specific monitoring and conservation efforts for maintaining resilient salmon runs in a warming world.     

Interhabitat migration of juvenile Atlantic salmon in a Newfoundland river system, Canada

Migrations of juvenile Atlantic salmon between habitats (both fluvial and lacustrine) were studied in a Newfoundland river system from May through September. Salmon parr showed poor site fidelity in fluvial habitats, and high rates of migration, especially in spring-early summer. Most habitat shifts were upstream from fluvial habitats to a lake at the head of a small tributary. In September, 10% of the salmon parr caught in the lake ( n =275) had been marked in the main stem of the river or in the tributary stream (total n =641). The abundance of parr in the lake increased in May-June, then declined gradually to September. Most of the lacustrine immigrants were 1+ parr. Mature males were found amongst the autumnal emigrants. Salmon parr in the lake grew larger than those in the river, and lacustrine smolts captured in the lake were larger than those caught at the mouth of the river.     

Scale and the detection of climatic influences on the productivity of salmon populations

Ecological studies relating population parameters to climate conditions are limited by a lack of experimental control systems and rely instead on correlative evidence to draw inferences about how populations respond to environmental forcing. Consequently, some correlations turn out to be spurious and not ecologically meaningful. To strengthen inferences, multiple populations may be examined simultaneously to confirm whether relationships can be generalized across multiple systems; however, this assumes that populations respond similarly to climate drivers, ignoring the potential for ecological complexity. Using data on eight sockeye salmon populations from southwestern Alaska, we constructed a series of models based on ecological hypotheses, relating salmon population productivity to climate factors experienced at different life stages. We modeled populations at a range of organizational scales, from distinct populations, to populations grouped by common nursery lake, to all populations within a watershed, and determined the relative statistical support for climate drivers at each scale. In general, warmer lake and sea surface temperatures in the summer coincided with increased productivity of these populations, but the most sensitive life‐stage for climate effects varied among populations, particularly among nursery lakes. The best model when considering all populations together, despite strong statistical support, failed to represent the complexity which became evident when populations were modeled by common nursery lake, or independently. These results emphasize that the most appropriate organizational scale to model salmon stocks will depend on specific management, scientific, or conservation goals.     

集合生态系统研究15年回顾与展望

集合生态系统(meta-ecosystem)由法国的Loreau教授等于2003年提出,是指"由跨生态系统边界的物质流、能量流和生物体流所连接起来的一系列生态系统的集合",是对只关注生物体(organism)迁移交换的集合种群(meta-population)和集合群落(meta-community)概念的外推,也是为了给生态系统空间异质性研究提供一个重要的分析路径,对研究和理解生态系统的结构、过程、功能和异质性具有重要意义。通过对相关文献的梳理分析,简述了集合生态系统研究的基本状况,分析了对集合生态系统概念的狭义和广义两种理解,指出了探讨集合生态系统结构的两个方向,构建了分析集合生态系统研究的六维整体框架,综述了研究集合生态系统的两类方法,探讨了经验化的集合生态系统(empirical meta-ecosystem)的3种空间结构和两种构建路径。将集合生态系统概念和理论引入流域复合生态系统(integrated watershed ecosystem)的分析,为流域生态学研究提供新的概念框架。     

A wide window of migration phenology captures inter‐annual variability of favourable conditions: Results of a whole‐lake experiment with juvenile Pacific salmon

相似文献   

Use of glacier river-fed estuary channels by juvenile Coho Salmon: transitional or rearing habitats?

Estuaries are among the most productive ecosystems in the world and provide important rearing environments for a variety of fish species. Though generally considered important transitional habitats for smolting salmon, little is known about the role that estuaries serve for rearing and the environmental conditions important for salmon. We illustrate how juvenile coho salmon Oncorhynchus kisutch use a glacial river-fed estuary based on examination of spatial and seasonal variability in patterns of abundance, fish size, age structure, condition, and local habitat use. Fish abundance was greater in deeper channels with cooler and less variable temperatures, and these habitats were consistently occupied throughout the season. Variability in channel depth and water temperature was negatively associated with fish abundance. Fish size was negatively related to site distance from the upper extent of the tidal influence, while fish condition did not relate to channel location within the estuary ecotone. Our work demonstrates the potential this glacially-fed estuary serves as both transitional and rearing habitat for juvenile coho salmon during smolt emigration to the ocean, and patterns of fish distribution within the estuary correspond to environmental conditions.     

Founding events influence genetic population structure of sockeye salmon (Oncorhynchus nerka) in Lake Clark, Alaska

Bottlenecks can have lasting effects on genetic population structure that obscure patterns of contemporary gene flow and drift. Sockeye salmon are vulnerable to bottleneck effects because they are a highly structured species with excellent colonizing abilities and often occupy geologically young habitats. We describe genetic divergence among and genetic variation within spawning populations of sockeye salmon throughout the Lake Clark area of Alaska. Fin tissue was collected from sockeye salmon representing 15 spawning populations of Lake Clark, Six-mile Lake, and Lake Iliamna. Allele frequencies differed significantly at 11 microsatellite loci in 96 of 105 pairwise population comparisons. Pairwise estimates of FST ranged from zero to 0.089. Six-mile Lake and Lake Clark populations have historically been grouped together for management purposes and are geographically proximate. However, Six-mile Lake populations are genetically similar to Lake Iliamna populations and are divergent from Lake Clark populations. The reduced allelic diversity and strong divergence of Lake Clark populations relative to Six-mile Lake and Lake Iliamna populations suggest a bottleneck associated with the colonization of Lake Clark by sockeye salmon. Geographic distance and spawning habitat differences apparently do not contribute to isolation and divergence among populations. However, temporal isolation based on spawning time and founder effects associated with ongoing glacial retreat and colonization of new spawning habitats contribute to the genetic population structure of Lake Clark sockeye salmon. Nonequilibrium conditions and the strong influence of genetic drift caution against using estimates of divergence to estimate gene flow among populations of Lake Clark sockeye salmon.     

Review of proposed mechanisms for sockeye salmon population cycles in the fraser river

Prominent and persistent cyclic fluctuations in the abundance of consecutive year-classes occur in some sockeye salmon populations throughout the species' range. We review and test a number of explanations for the existence of these cycles using qualitative biological arguments, including a consideration of the synchrony of cycles among populations. Most of the hypotheses involve mechanisms that would reinforce synchronous population fluctuations within watersheds. However, the 4-year cycles characteristic of many Fraser River sockeye populations are sometimes out of phase with each other, both among populations which migrate together as mixed stocks while vulnerable to commerical fisheries, and among populations whose juveniles share the same nursery lake habitat (Shuswap Lake). Such asynchrony suggests that the mechanism(s) causing population cycles can operate independently within reproductively isolated populations. Of the mechanisms reviewed here, only those involving genetic effects on age at maturation, or on resistance to disease or parasites, or those involving depensatory predation soon after fry emergence, appear to offer satisfactory explanations.     

Juvenile sockeye salmon distribution, size, condition and diet during years with warm and cool spring sea temperatures along the eastern Bering Sea shelf

Interannual variations in distribution, size, indices of feeding and condition of juvenile Bristol Bay sockeye salmon Oncorhynchus nerka collected in August to September (2000–2003) during Bering–Aleutian Salmon International Surveys were examined to test possible mechanisms influencing their early marine growth and survival. Juvenile sockeye salmon were mainly distributed within the southern region of the eastern Bering Sea, south of 57°0' N during 2000 and 2001 and farther offshore, south of 58°0' N during 2002 and 2003. In general, juvenile sockeye salmon were significantly larger ( P < 0·05) and had significantly higher indices of condition ( P < 0·05) during 2002 and 2003 than during 2000 and 2001. The feeding index was generally higher for age 1.0 year sockeye salmon than age 2.0 year during all years. Among-year comparisons suggested that Pacific sand lance Ammodytes hexapterus were important components of the juvenile sockeye salmon diet during 2000 and 2001 (20 to 50% of the mean wet mass) and age 0 year walleye pollock Theragra chalcogramma were important components during 2002 and 2003 (50 to 60% of the mean wet mass). Warmer sea temperatures during spring and summer of 2002 and 2003 probably increased productivity on the eastern Bering Sea shelf, enhancing juvenile sockeye salmon growth.     

Ephemeral floodplain habitats provide best growth conditions for juvenile Chinook salmon in a California river

We reared juvenile Chinook salmon for two consecutive flood seasons within various habitats of the Cosumnes River and its floodplain to compare fish growth in river and floodplain habitats. Fish were placed in enclosures during times when wild salmon would naturally be rearing in floodplain habitats. We found significant differences in growth rates between salmon reared in floodplain and river enclosures. Salmon reared in seasonally inundated habitats with annual terrestrial vegetation experienced higher growth rates than those reared in a perennial pond on the floodplain. Growth of fish in the non-tidal river upstream of the floodplain varied with flow in the river. When flows were high, there was little growth and high mortality, but when the flows were low and clear, the fish grew rapidly. Fish displayed very poor growth in tidally influenced river habitat below the floodplain, a habitat type to which juveniles are commonly displaced during high flow events due to a lack of channel complexity in the main-stem river. Overall, ephemeral floodplain habitats supported higher growth rates for juvenile Chinook salmon than more permanent habitats in either the floodplain or river. Variable responses in both growth and mortality, however, indicate the importance of providing habitat complexity for juvenile salmon in floodplain reaches of streams, so fish can find optimal places for rearing under different flow conditions.     

Analysis of gene origin in the first adult returns to the Cultus sockeye salmon captive breeding program

Rigorous evaluation of the utility of captive breeding for the restoration of depleted wild salmonid fish populations has not been undertaken. In particular, little is known about the reproductive success of captively-bred individuals that are released back into an extant population and their capacity to assist in long-term population persistence. For the endangered Cultus Lake sockeye salmon population, we examined the potential genetic contribution of the first juvenile fish released from a captive breeding program upon their maturity in the natural Cultus Lake environment. Genetic analysis of 792 Cultus sockeye salmon that were spawned in captivity in 2004 and their adult progeny of 2007 and 2008 revealed a genetic bottleneck originating from 20 wild sockeye salmon hatchery-spawned at Cultus Lake in the previous generation. Pedigree analysis revealed that six of the 20 sockeye salmon spawned in 2001 (grandparents) gave rise to a majority of the hatchery spawners in 2004 (parents) and provided more than 30% of the genes in the progeny that survived to maturity in the wild. Allele frequencies and genetic diversity of the age three progeny that returned to Cultus Lake from their marine migration in 2007 reflected the bottleneck, but its genetic signature was faint among the more genetically diverse age four fish that returned in 2008. Two-generation analysis of gene origin among fish resulting from 2004 hatchery production indicated that they contained the genetic diversity expected from 36 effective ancestors.     

The distribution of juvenile Pacific salmon and associated fish species in Lake Sopochnoye,Iturup Island

A single survey using a minnow seine net showed that juvenile chum and sockeye salmon are distributed extremely unevenly along the shoreline of Lake Sopochnoye, on Iturup Island. Young-of-the-year chum salmon aggregate mainly near the estuaries of the rivers that flow into the lake; young-of-the-year and 2-year-old sockeye salmon aggregate closer to the central part of the lake. The distribution of associated fish species is also very uneven, but none of them dominates within the zone of catches; the abundance of each of these species is comparable to that of the juvenile Pacific salmon.     

Individual variation in migration speed of upriver-migrating sockeye salmon in the Fraser River in relation to their physiological and energetic status at marine approach

Little research has examined individual variation in migration speeds of Pacific salmon (Oncorhynchus spp.) in natural river systems or attempted to link migratory behavior with physiological and energetic status on a large spatial scale in the wild. As a model, we used three stocks of summer-run sockeye salmon (Oncorhynchus nerka) from the Fraser River watershed, British Columbia, to test the hypothesis that individual variation in migration speed is determined by a combination of environmental factors (i.e., water temperature), intrinsic biological differences (sex and population), and physiological and energetic condition. Before the freshwater portion of the migration, sockeye salmon (Quesnel, Chilcotin, and Nechako stock complexes) were captured in Johnstone Strait ( approximately 215 km from river entry), gastrically implanted with radio transmitters, and sampled for blood, gill tissue, and energetic status before release. Analyses focused solely on individuals that successfully reached natal subwatersheds. Migration speeds were assessed by an extensive radiotelemetry array. Individuals from the stock complex that migrated the longest distance (Nechako) traveled at speeds slower than those of other stock complexes. Females traveled slower than males. An elevated energetic status of fish in the ocean was negatively correlated with migration speed in most river segments. During the transition from the ocean to the river, migration speed was negatively correlated with mean maximum water temperature; however, for the majority of river segments, it was positively correlated with migration speed. Physiological status measured in the ocean did not explain among-individual variability in river migration speeds. Collectively, these findings suggest that there could be extensive variation in migration behavior among individuals, sexes, and populations and that physiological condition in the ocean explained little of this variation relative to in-river environmental conditions and energetic status. Interestingly, individual fish generally retained their rank in swimming speed across different segments, except when transiting a challenging canyon midway during the migration.     

Gauging resource exploitation by juvenile Chinook salmon (Oncorhynchus tshawytscha) in restoring estuarine habitat

In the context of delta restoration and its impact on salmonid rearing, success is best evaluated based on whether out‐migrating juvenile salmon can access and benefit from suitable estuarine habitat. Here, we integrated 3 years of post‐restoration monitoring data including habitat availability, invertebrate prey biomass, and juvenile Chinook salmon (Oncorhynchus tshawytscha) physiological condition to determine whether individuals profited from the addition of 364 ha of delta habitat in South Puget Sound, Washington, United States. Productivity in the restored mudflat was comparable to reference sites 3 years after dike removal, surpassing a mean total of 6 million kJ energy from invertebrate prey. This resulted from the development of a complex network of tidal channels and a resurgence in dipteran biomass that was unique to the restoration area. Consequently, a notable shift in invertebrate consumption occurred between 2010 and 2011, whereby individuals switched from eating primarily amphipods to dipteran flies; however, dietary similarity to the surrounding habitat did not change from year to year, suggesting that this shift was a result of a change in the surrounding prey communities. Growth rates did not differ between restored and reference sites, but catch weight was positively correlated with prey biomass, where greater prey productivity appeared to offset potential density‐dependent effects. These results demonstrate how the realized function of restoring estuarine habitat is functionally dependent. High prey productivity in areas with greater connectivity may support healthy juvenile salmon that are more likely to reach the critical size class for offshore survival.