1. Introduction
Anadromous and potamodromous salmonids migrate from their natal river to a feeding environment before returning for reproduction
[1,2,3][1][2][3]. Migration enables fish to exploit many temporally productive and spatially discrete habitats with various fitness benefits (e.g., growth, reproduction, predator avoidance)
[4]. Migratory life history requires unrestricted migration routes between nursery, feeding, and spawning habitats
[5]. During each life stage, salmonids utilize the habitat that is advantageous for them. Migration between habitats thus clearly has an adaptive value
[6]. Nursery and feeding habitats differ in environmental characteristics, and migrations precede adaptive physiological transformations and changes in the phenotype and behavior to be better suited for the new environment.
Smolting is the major transformation anadromous salmonids undergo before migration to feeding areas. Smolting prepares fish for downstream migration and entry to seawater. Atlantic salmon
Salmo salar L., 1758 juveniles can stay in their natal river habitat to grow for 1–8 years before migrating
[7,8][7][8]. Of the Pacific salmon, pink salmon
Oncorhynchus gorbuscha (Walbaum, 1792) and chum salmon
Oncorhynchus keta (Walbaum, 1792) can move almost directly after the emergence at the fry stage into seawater, while the others (masu salmon
Oncorhynchus masou (Brevoort, 1856),
O. tshawytscha (Walbaum, 1792),
O. nerka (Walbaum, 1792), and steelhead (rainbow) trout
Oncorhynchus mykiss (Walbaum, 1792)) spend one or more years in fresh water before migrating to the sea for feeding
[7]. When smolting, the phenotype of fish changes as follows: the coloration of smolts becomes silvery, and the body shape becomes more streamlined
[2]. This, with darkened fins, a dark back, and a white abdomen, camouflage the fish in the pelagic environment
[5]. Behavioral changes include a loss of rheotaxis, and juveniles become more pelagic. Their tendency to group also increases
[9]. Several physiological changes occur, for example, increased salinity tolerance, increased metabolism, and olfactory imprinting helping the fish locate their home stream on their return migration
[10]. Environmental cues, e.g., photoperiod, temperature, and waterflow, regulate physiological changes and initiate migration
[2,5][2][5]. Lake-living Atlantic salmon appear to smolts similarly to anadromous conspecifics (
[11], but see
[12]). This is apparently an inherited trait
[13], even for salmon spending their entire life in fresh water, such as the Atlantic salmon residing in Lake Vänern, Sweden.
Both Atlantic and Pacific salmon populations have been in decline throughout their habitat ranges
[13,14,15][13][14][15]. To reverse this trend, it is important to understand the role of different environmental and anthropogenic factors in the decline
[16]. Numerous factors may impact population abundances negatively, and with the complex life history of migrating salmonids, the reasons are obviously multiple and difficult to unravel
[17[17][18],
18], although the ongoing climate warming appears particularly important, especially for low latitude populations. Anthropogenic activities have a long history of altering salmonid populations and, thus, smolt development and smolt migration. Smolts are sensitive to external impacts and behavior and survival during migrations
[2]. Several anthropogenic activities may affect smolt development, behavior, and survival during migration, such as hydropower developments, land use, pollution, fish farming, and parasites like sea lice
Lepeophtheirus salmonis (Krøyer, 1837)
[2,5][2][5]. Temperature and flow interact with the other anthropogenic pressures to affect smolting and smolt migration.
Salmonids are a cold-water species. Global warming will generally have a major impact on their success. Historically, climatic variability has affected the patterns of abundance in Atlantic salmon and Pacific salmon populations
[19,20,21,22][19][20][21][22]. Although estuarine and marine mortalities have been found to be important determinants of survival, marine mortality depends on factors acting in fresh water and during smolt migration
[23]. Thorstad et al.
[24] argue that the best strategy to mitigate the changing environmental conditions should be to ensure that the greatest number of wild smolts in the best condition migrate from rivers and coastal areas to feeding areas because mortality at sea is found to be density independent
[25]. Survival at sea depends on the size of the smolts and environmental conditions when the smolts begin their sea sojourn.
[18,23,26][18][23][26]. In research, it is important to address the links between river habitat conditions and the physiological requirements of salmonids during their juvenile life stages in freshwater habitats
[27]. Climate change will continue to affect not only smolting and migration but also instream habitats across all seasons
[27].
2. Climate Change and Salmonid Distribution
Human activities are estimated to have induced approximately 1.0 °C of global warming above pre-industrial levels (between 1880 and 2017), with a likely range of 0.8 °C to 1.2 °C. Global warming is likely to reach 1.5 °C in about 2030 if temperatures continue to increase at the current rate
[28]. For example, a higher winter discharge, earlier snowmelt, and earlier onset of summer low flow periods are predicted throughout the range of Atlantic salmon
[29,30][29][30].
Increasing global surface temperatures are very likely to lead to changes in precipitation and atmospheric moisture because of changes in atmospheric circulation, a more active hydrological cycle, and increases in the water-holding capacity throughout the atmosphere. Overall, global land precipitation has increased by about 2% since the beginning of the 20th century. There have been marked increases in precipitation in the latter part of the 20th century over northern Europe, though with a general decrease southward to the Mediterranean. Dry wintertime conditions over southern Europe and the Mediterranean and wetter-than-normal conditions over many parts of northern Europe and Scandinavia
[31] are linked to the strong positive values of the North Atlantic Oscillation (NAO), with more anticyclonic conditions over southern Europe and stronger westerly winds over northern Europe (Ref.
[32] conducted a review).
Northern Eurasia (north of approximately 40 °N) showed widespread and statistically significant increases in winter precipitation between 1921 and 2015, with values exceeding 1.2–1.6 mm mo
−1 per decade west of the Ural Mountains and along the east coast, while southern Europe exhibits coherent yet weaker amplitude drying trends that attain statistical significance over the eastern Mediterranean. These precipitation trends occur in the context of changes in the large-scale atmospheric circulation, with negative SLP (Sea Level Pressure) trends over northern Eurasia and positive SLP trends over the central North Atlantic extending into southwestern Europe
[33].
The magnitude of climate change is considered to depend on the atmospheric load of the two most important greenhouse gases: carbon dioxide (CO
2) and methane (CH
4). The terrestrial biosphere plays an important role in the global carbon balance. In boreal zones, forests and peatlands are an essential part of the global carbon cycle. Recent temperature increases have been associated with increasing forest fire activity in Canada since about 1970 and exceptionally warm summer conditions in Russia during the 2010 fire season reviewed by
[34].
Atlantic salmon is distributed from northern Portugal (42 °N) to the River Kara in northern Russia in Europe
[35], and West Atlantic salmon is distributed from the Connecticut River to the Ungava region of northern Quebec. Southern Atlantic salmon populations have declined dramatically and face the highest risk of extinction as global warming moves its thermal niche northward
[36]. The suitable thermal habitat for salmon is expected to extend northward with the invasion of new spawning, nursery, and feeding areas north of the species’ present distributional range but with the loss of the most southern populations
[37,38,39,40][37][38][39][40]. Indeed, salmon are already responding to warmer temperatures by expanding their range northward into the Arctic Ocean
[41,42][41][42] and disappearing from the southern edge of their distribution area
[7,40,43,44,45][7][40][43][44][45]. The population complex of Atlantic salmon in Europe has experienced a multidecadal decline in recruitment, resulting in the lowest population abundances observed since 1970
[46]. Atlantic salmon abundance and productivity show similar patterns of decline across six widespread regions of North America
[47]. Abundance declined in the late 1980s and early 1990s, after which it remained stable at low levels. Climate-driven environmental factors such as changes in plankton communities and prey availability at warmer ocean temperatures were linked to the low productivity of North Atlantic salmon populations
[47]. Landlocked European populations of salmonids are found in Norway, Sweden, Finland, and Russian Karelia
[48,49,50,51][48][49][50][51]. The landlocked populations of salmon have declined throughout their distribution range
[51,52][51][52]. Brown trout (
Salmo trutta L. 1758) is native to Europe and Asia, where anadromous populations are found from Portugal to the White Sea
[7]. It must be noted that the taxonomic status of the brown trout species complex is challenging, and the high morphological and ecological diversity has led to the morphological description of populations belonging to species other than
S. trutta in Europe
[53]. In the future, the living conditions for trout will probably deteriorate in the southern part of the current distribution. In the northern part of their current distribution, global warming may improve feeding opportunities, growth, and survival conditions
[7]. According to Filipe et al.
[54], future brown trout distribution will become progressively and dramatically reduced in European watercourses. Their forecasts indicate that the greatest losses in suitable habitats will take place in southern Europe.