Per-Arne Amundsen

Heavy metal contamination in freshwater fish from the border region between Norway and Russia

The contents of Cd, Cu, Cr, Hg, Ni and Zn in muscle, liver and gills were studied in whitefish, perch, pike, brown trout, burbot and vendace from three lake localities in a watercourse in the border region between Norway and Russia, in the vicinity of mining activity and several metallurgic smelters. The contents of Cd and Ni in fish tissue increased with increasing proximity to the smelters, whereas the other elements showed similar concentrations at the three localities. The recorded heavy metal concentrations appeared to be within the ranges reported for fish from other metal-contaminated lakes, and higher than comparable observations from unpolluted systems. The heavy metal concentrations were usually lowest in muscle and highest in the liver or the gills. Significant differences in metal concentration levels were found between different fish species, but Hg was the only metal where these species differences were possibly related to biomagnification. For the other elements, the concentrations generally appeared to be inversely related to the trophic level of the fish species.

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.

Culling prey promotes predator recovery - Alternative states in a whole-lake experiment

Many top-predator fish stocks in both freshwater and marine systems have collapsed as a result of overharvesting. Consequently, some of these communities have shifted into seemingly irreversible new states. We showed, for predators feeding on prey that exhibit food-dependent growth, that culling of fish prey may promote predator recovery. We removed old stunted individuals of a prey-fish species in a large, low-productive lake, which caused an increase in the availability of small-sized prey and allowed the predator to recover. The shift in community state has been sustained for more than 15 years after the cull ended and represents an experimental demonstration of an alternative stable state in a large-scale field system. Because most animals exhibit food-dependent growth, shifts into alternative stable states resulting from overcompensating prey growth may be common in nature and may require counterintuitive management strategies.

THE COMPETITIVE EDGE OF AN INVADING SPECIALIST

Introduced species represent major threats to native and natural biodiversity. On the other hand, biologists may increase the understanding of ecological interactions by following communities during establishment of exotic species. Accordingly, feeding ecology and habitat use were studied in native whitefish (Coregonus lavaretus) and recently invading vendace (C. albula) in two lake localities situated 50 km apart within the subarctic Pasvik River system, northern Norway and Russia. Whitefish originally dominated the native fish communities of both lakes. The recent invasion and successive downstream expansion of vendace allowed comparisons between two sites: one in which the influence of the new potential competitor on the native fish species was weak, and one in which the influence was strong. In the downstream lake vendace was recorded for the first time at the time of the study, and only in small numbers, whereas in the upstream lake vendace had established a high population density and was the dominant fish species in the pelagic zone. No vertical segregation in pelagic habitat use was found between the two fish species in either lake. In the downstream lake both whitefish and vendace fed exclusively on zooplankton and had almost identical diets. In the upstream lake, in contrast, whitefish fed predominantly on zoobenthos and surface insects, while vendace fed mainly on zooplankton. Thus, the strong presence of vendace as a specialized planktivore reduced the availability of zooplankton as prey for the more generalist whitefish. The food segregation between the two fish species in the upstream lake was apparently interactive and caused by a strong asymmetrical competition for zooplankton, vendace being the superior species. The ecological consequences (including reduced zooplankton size and species diversity, alteration of the pelagic food web, and eutrofication as a possible cascading effect on the primary production) of the vendace invasion in the Pasvik watercourse are considerable, even after a few years, and are likely to proceed and intensify in the future.

Food web topology and parasites in the pelagic zone of a subarctic lake.

1. Parasites permeate trophic webs with their often complex life cycles, but few studies have included parasitism in food web analyses. Here we provide a highly resolved food web from the pelagic zone of a subarctic lake and explore how the incorporation of parasites alters the topology of the web. 2. Parasites used hosts at all trophic levels and increased both food-chain lengths and the total number of trophic levels. Their inclusion in the network analyses more than doubled the number of links and resulted in an increase in important food-web characteristics such as linkage density and connectance. 3. More than half of the parasite taxa were trophically transmitted, exploiting hosts at multiple trophic levels and thus increasing the degree of omnivory in the trophic web. 4. For trophically transmitted parasites, the number of parasite-host links exhibited a positive correlation with the linkage density of the host species, whereas no such relationship was seen for nontrophically transmitted parasites. Our findings suggest that the linkage density of free-living species affects their exposure to trophically transmitted parasites, which may be more likely to adopt highly connected species as hosts during the evolution of complex life cycles. 5. The study supports a prominent role for parasites in ecological networks and demonstrates that their incorporation may substantially alter considerations of food-web structure and functioning.

Competitive exclusion after invasion

The ‘Competitive Exclusion Principle’ is a foundation stone in the understanding of interspecific competition and niche relationships between species. In spite of having the status of a biological law, the principle has limited empirical support. In this study, we document strong effects of competition from the invading fish species vendace Coregonus albula over a 14-year period in the sub-arctic Pasvik watercourse. The native d.r. whitefish, that shared food and habitat niche with the invader, was displaced from its original niche and showed a more than 90% decline in population density over the study period. The study thus provides a unique record of how an exotic fish species excludes a native species from its original niche. Our data support the competitive exclusion principle, but also indicate that the vulnerability of the inferior competitor depends on a lack of alternative resources and on indirect ecological interactions.

Incipient speciation through niche expansion: an example from the Arctic charr in a subarctic lake

Two reproductive isolated morphs of Arctic charr (Salvelinus alpinus), termed profundal and littoral charr according to their different spawning habitats, co-occur in the postglacial lake Fjellfrøsvatn in North Norway. All profundal charr live in deep water their entire life and have a maximum size of 14 cm, while the littoral charr grow to 40 cm. Some small and young littoral charr move to the profundal zone in an ontogenetic habitat shift in the ice-free season and the rest of the population remains in epilimnic waters. The two morphs had different diet niches in the profundal zone: the profundal charr ate typical soft-bottom prey (chironomid larvae, pea mussels and benthic copepods), while the young littoral charr mainly consumed crustacean zooplankton. In four other lakes without a profundal morph (i.e. monomorphic populations), young charr also performed ontogenetic habitat shifts to the profundal zone and fed on zooplankton. The profundal morph of Fjellfrøsvatn therefore utilize a food resource niche that neither the littoral morph nor comparable monomorphic populations exploit. This suggests that intraspecific resource competition has driven incipient ecological speciation of the profundal charr of Fjellfrøsvatn. The exploitation of the soft-bottom resources by the profundal charr supports earlier experimental findings that the profundal morph is genetically different in trophic behaviour and morphology. The sympatric ecological divergence within the profundal habitat is possible because unexploited food resources (soft-bottom profundal prey) are available. Apparently, this represents a case of incipient segregation by expansion to new resource types (niche invasion), and not by subdivision of one broad ancestral niche.

The role of gill raker number variability in adaptive radiation of coregonid fish

Gill raker divergence is a general pattern in adaptive radiations of postglacial fish, but few studies have addressed the adaptive significance of this morphological trait in foraging and eco-evolutionary interactions among predator and prey. Here, a set of subarctic lakes along a diversifying gradient of coregonids was used as the natural setting to explore correlations between gill raker numbers and planktivory as well as the impact of coregonid radiation on zooplankton communities. Results from 19 populations covering most of the total gill raker number gradient of the genus Coregonus, confirm that the number of gill rakers has a central role in determining the foraging ability towards zooplankton prey. Both at the individual and population levels, gill raker number was correlated with pelagic niche use and the size of utilized zooplankton prey. Furthermore, the average body size and the abundance and diversity of the zooplankton community decreased with the increasing diversity of coregonids. We argue that zooplankton feeding leads to an eco-evolutionary feedback loop that may further shape the gill raker morphology since natural selection intensifies under resource competition for depleted prey communities. Eco-evolutionary interactions may thus have a central role creating and maintaining the divergence of coregonid morphs in postglacial lakes.

Temporal stability of individual feeding specialization may promote speciation.

1. Inter-individual differences in trophic behaviour are considered important in the disruptive selection process for resource specialization and may represent an early phase in the evolution of polymorphism and adaptive radiation. Here, we provide evidence of high stability of individual trophic niches of a fish predator from a 15-year study. 2. Individual resource specialization was investigated by combining data from analyses of stomach contents (recent trophic niche), trophically transmitted parasites (long-term niche) and trophic morphology (niche adaptations) from single specimens of a postglacial fish (Arctic charr) population sampled from contrasting pelagic and littoral habitats. 3. Based on the relationships between morphology, parasites and diet, high inter-individual temporal consistency of narrow niches (zooplanktivorous vs. benthivorous) was evident through the ontogeny of the charr, indicating low degree of switching both in habitat utilization and feeding strategy of individual fish. Co-occurrence of differently specialized behavioural phenotypes was sustained over multiple generations. 4. The stable long-term habitat and feeding specializations may represent an important initial step in an adaptive radiation process, and our findings suggest a case of sympatric speciation into two incipient forms diverging along the littoral-pelagic resource axis.

Seasonal and ontogenetic variations in resource use by two sympatric Arctic charr morphs

The study compares the resource utilization of two sympatric Arctic charr morphs over an annual period in a subarctic lake. The two morphs are reproductively isolated in time and place of spawning, and are referred to as the littoral and profundal morphs (L-morph and P-morph) according to their spawning habitats. Fish were sampled monthly (ice-free season) or bimonthly (winter) using gillnets in the main lake habitats. The spatial range of the P-morph was restricted to the profundal zone throughout the whole annual period. The L-morph in contrast utilized all main habitats, exhibiting distinct seasonal and ontogenetic variations in habitat distribution. In the spring, the whole L-morph population was located along the bottom profile of the lake, in profundal and littoral habitats. During summer and autumn, habitat segregation occurred between different life-stages, juveniles mainly utilizing the profundal, pre-adults the pelagic and adult fishes the littoral zone. During winter the whole population was assembled in the littoral habitat. The L-morph also had large seasonal and ontogenetic variations in their feeding ecology, with littoral zoobenthos, zooplankton and surface insects being important prey. The P-morph had a narrower diet niche mainly consisting of chironomid larvae and other profundal zoobenthos. Hence, the two Arctic charr morphs exhibited a consistent resource differentiation during all annual seasons and throughout their life cycles, except for a dietary overlap between P-morph and juvenile L-morph charr in the profundal during summer. The findings are discussed in relation to resource polymorphism and incipient speciation.