Marine depth use of sea trout Salmo trutta in fjord areas of central Norway.

The vertical behaviour of 44 veteran sea trout Salmo trutta (275-580 mm) in different marine fjord habitats (estuary, pelagic, near shore with and without steep cliffs) was documented during May-February by acoustic telemetry. The swimming depth of S. trutta was influenced by habitat, time of day (day v. night), season, seawater temperature and the body length at the time of tagging. Mean swimming depth during May-September was 1·7 m (individual means ranged from 0·4 to 6·4 m). Hence, S. trutta were generally surface oriented, but performed dives down to 24 m. Mean swimming depth in May-September was deeper in the near-shore habitats with or without steep cliffs (2·0 m and 2·5 m, respectively) than in the pelagic areas (1·2 m). May-September mean swimming depth in all habitats was slightly deeper during day (1·9 m) than at night (1·2 m), confirming that S. trutta conducted small-scale diel vertical movements. During summer, S. trutta residing in near-shore habitat progressively moved deeper over the period May (mean 1·1 m) to August (mean 4·0 m) and then reoccupied shallower areas (mean 2·3 m) during September. In winter (November and February), individuals residing in the innermost part of the fjords were found at similar average depths as they occupied during the summer (mean 1·3 m). The swimming depths of S. trutta coincide with the previously known surface orientation of salmon lice Lepeophtheirus salmonis. Combined with previous studies on horizontal use of S. trutta, this study illustrates how S. trutta utilize marine water bodies commonly influenced by anthropogenic factors such as aquaculture, harbours and marine constructions, marine renewable energy production or other human activity. This suggests that the marine behaviour of S. trutta and its susceptibility to coastal anthropogenic factors should be considered in marine planning processes.

Ice-dependent winter survival of juvenile Atlantic salmon.

Changes in snow and ice conditions are some of the most distinctive impacts of global warming in cold temperate and Arctic regions, altering the environment during a critical period for survival for most animals. Laboratories studies have suggested that reduced ice cover may reduce the survival of stream dwelling fishes in Northern environments. This, however, has not been empirically investigated in natural populations in large rivers. Here, we examine how the winter survival of juvenile Atlantic salmon in a large natural river, the River Alta (Norway, 70°N), is affected by the presence or absence of surface ice. Apparent survival rates for size classes corresponding to parr and presmolts were estimated using capture-mark-recapture and Cormack-Jolly-Seber models for an ice-covered and an ice-free site. Apparent survival (Φ) in the ice-covered site was greater than in the ice-free site, but did not depend on size class (0.64 for both parr and presmolt). In contrast, apparent survival in the ice-free site was lower for larger individuals (0.33) than smaller individuals (0.45). The over-winter decline in storage energy was greater for the ice-free site than the ice-covered site, suggesting that environmental conditions in the ice-free site caused a strong depletion in energy reserves likely affecting survival. Our findings highlight the importance of surface ice for the winter survival of juvenile fish, thus, underpinning that climate change, by reducing ice cover, may have a negative effect on the survival of fish adapted to ice-covered habitats during winter.

The gill maggot Salmincola salmoneus as an indicator of repeat spawning in Atlantic salmon Salmo salar.

The potential of the gill maggot Salmincola salmoneus for use as an indicator of repeat spawning in Atlantic salmon Salmo salar was studied in adult fish captured during their return migration to the River Alta (n = 659) and River Namsen (n = 540) in Norway. Eighty-eight and 49% of previous spawners identified by scale readings were infected with S. salmoneus in the two rivers, respectively. Salmincola salmoneus can be used as a reliable, rapid and objective field indicator of repeat spawning in S. salar as nearly all infected fish (99·4%) were identified as repeat spawners, although it is important to have appropriate background information on S. salmoneus prevalence on the postspawning individuals within the same population.

An assessment of the size structure, distribution and behaviour of fish populations within a temporarily closed estuary using dual frequency identification sonar (DIDSON).

The fish populations within the East Kleinemonde Estuary, South Africa, were studied using Dual frequency IDentification SONar (DIDSON) during March 2010 which allowed for direct comparisons of the abundance, distribution and behaviour of a wide size range of fishes along the length of the system. Small schooling fishes <100 mm in total length (L(T) ) were most abundant while larger size classes 100-300 and 300-500 mm were less abundant, but evenly distributed longitudinally within the estuary. Fishes <100 mm were most abundant within the mouth region which may reflect the feeding ecology of species that fall into these size classes and the effects of higher turbidities further up the estuary. Large fishes >500 mm were similarly most abundant near the mouth, and fishes of this size being mostly piscivorous, probably showed these distribution patterns due to the location of smaller prey fishes. Differential behaviour was observed among the longitudinal sections, with small fishes forming schools in areas of low turbidity and particularly in the presence of large predatory fishes. Results from this study present a snap-shot of the composition, distribution and behaviour of the ichthyofauna within a small temporarily closed estuary, and also show that abiotic factors and interactions of various size classes can influence their distribution. There is significant potential for DIDSON in future ecological research within these systems.

Pre-winter lipid stores in young-of-year Atlantic salmon along a north-south gradient.

The pre-winter lipid stores of young-of-the-year (YOY, age 0 year) Atlantic salmon Salmo salar were analysed along a north-south gradient from c. 71 to 58 degrees N, with winter conditions ranging from >200 days of ice cover to no ice. The rivers sampled in Northern Norway represent some of the most northerly S. salar rivers. There was an increase in lipid content with increasing latitude, and mean lipid content (size adjusted to common mass) for YOY in northern rivers were almost three times higher: 0.035 g compared to 0.013 g in southern rivers. The relationship was not sensitive to variation in sampling time or variation in YOY body size. The lipid stores, however, varied markedly between rivers and also between neighbouring rivers, indicating different strategies or opportunities for pre-winter lipid storage both at latitudinal and local scales.

Migratory behaviour and survival rates of wild northern Atlantic salmon Salmo salar post-smolts: effects of environmental factors.

To study smolt behaviour and survival of a northern Atlantic salmon Salmo salar population during river descent, sea entry and fjord migration, 120 wild S. salar were tagged with acoustic tags and registered at four automatic listening station arrays in the mouth of the north Norwegian River Alta and throughout the Alta Fjord. An estimated 75% of the post-smolts survived from the river mouth, through the estuary and the first 17 km of the fjord. Survival rates in the fjord varied with fork length (LF), and ranged from 97.0 to 99.5% km(-1). On average, the post-smolts spent 1.5 days (36 h, range 11-365 h) travelling from the river mouth to the last fjord array, 31 km from the river mouth. The migratory speed was slower (1.8 LF s(-1)) in the first 4 km after sea entry compared with the next 27 km (3.0 LF s(-1)). Post-smolts entered the fjord more often during the high or ebbing tide (70%). There was no clear diurnal migration pattern within the river and fjord, but most of the post-smolts entered the fjord at night (66%, 2000-0800 hours), despite the 24 h daylight at this latitude. The tidal cycle, wind-induced currents and the smolts' own movements seemed to influence migratory speeds and routes in different parts of the fjord. A large variation in migration patterns, both in the river and fjord, might indicate that individuals in stochastic estuarine and marine environments are exposed to highly variable selection regimes, resulting in different responses to environmental factors on both temporal and spatial scales. Post-smolts in the northern Alta Fjord had similar early marine survival rates to those observed previously in southern fjords; however, fjord residency in the north was shorter.

Parallel evolution of ecomorphological traits in the European whitefish Coregonus lavaretus (L.) species complex during postglacial times.

The extensive phenotypic polymorphism in the European whitefish has triggered evolutionary research in order to disentangle mechanisms underlying diversification. To illuminate the ecological distinctiveness in polymorphic whitefish, and evaluate taxonomic designations, we studied nine Norwegian lakes in three watercourses, which each harboured pairs of divergent whitefish morphs. We compared the morphology and life history of these morphs, documented the extent of genetic differentiation between them, and contrasted the niche use of sympatric morphs along both the habitat and resource axes. In all cases, sympatric morphs differed in the number of gill rakers, a highly heritable trait related to trophic utilization. Individual growth rate, age and size at maturity, diet and habitat use also differed between morphs within lakes, but were remarkably similar across lakes within the same morph. Microsatellite analyses confirmed for all but one pair that sympatric morphs were significantly genetically different, and that similar morphs from different lakes likely have a polyphyletic origin. These results are most compatible with the process of parallel evolution through recurrent postglacial divergence into pelagic and benthic niches in each of these lakes. We propose that sparsely and densely rakered whitefish sympatric pairs may be a likely case of ecological speciation, mediated in oligotrophic lakes with few trophic competitors.

Evolutionary history of the European whitefish Coregonus lavaretus (L.) species complex as inferred from mtDNA phylogeography and gill-raker numbers.

We compared mitochondrial DNA and gill-raker number variation in populations of the European whitefish Coregonus lavaretus (L.) species complex to illuminate their evolutionary history, and discuss mechanisms behind diversification. Using single-strand conformation polymorphism (SSCP) and sequencing 528 bp of combined parts of the cytochrome oxidase b (cyt b) and NADH dehydrogenase subunit 3 (ND3) mithochondrial DNA (mtDNA) regions, we documented phylogeographic relationships among populations and phylogeny of mtDNA haplotypes. Demographic events behind geographical distribution of haplotypes were inferred using nested clade analysis (NCA) and mismatch distribution. Concordance between operational taxonomical groups, based on gill-raker numbers, and mtDNA patterns was tested. Three major mtDNA clades were resolved in Europe: a North European clade from northwest Russia to Denmark, a Siberian clade from the Arctic Sea to southwest Norway, and a South European clade from Denmark to the European Alps, reflecting occupation in different glacial refugia. Demographic events inferred from NCA were isolation by distance, range expansion, and fragmentation. Mismatch analysis suggested that clades which colonized Fennoscandia and the Alps expanded in population size 24 500-5800 years before present, with minute female effective population sizes, implying small founder populations during colonization. Gill-raker counts did not commensurate with hierarchical mtDNA clades, and poorly with haplotypes, suggesting recent origin of gill-raker variation. Whitefish designations based on gill-raker numbers were not associated with ancient clades. Lack of congruence in morphology and evolutionary lineages implies that the taxonomy of this species complex should be reconsidered.

Morphological divergence and origin of sympatric populations of European whitefish (Coregonus lavaretus L.) in Lake Femund, Norway.

Combining morphological and genetic analysis, we compared patterns of diversification within and between morphs among sympatric European whitefish (Coregonus lavaretus L.) populations in Lake Femund, Norway. Seven external populations, from potential colonization routes into Lake Femund were included. We found that deep-, shallow-, river- and bay spawning populations are distinct morphs in Lake Femund. Within morphs, populations range from being similar genetically (Fst=0-0.005) among deep-spawning populations to being highly differentiated (Fst=0.153) between bay-spawning populations. Between morphs, genetic differences ranged from a low (Fst=0.008-0.022) between deep- and shallow-spawning populations to high difference (Fst=0.125-0.143) between shallow- and bay-spawning populations. A higher proportion of molecular variance was seen among (3.9%) than within morphs (2.8%). The adaptive gene combinations behind the four morphs seem to have originated within the lake, although the lake could have been colonized from more than one source population.