Anders Klemetsen

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.

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.

The charr problem revisited: exceptional phenotypic plasticity promotes ecological speciation in postglacial lakes

Abstract The salmonid arctic charr Salvelinus alpinus (L.) is one of the most widespread fishes in the world and is found farther north than any other freshwater or diadromous fish, but also in cool water farther south. It shows a strong phenotypic, ecological, and life history diversity throughout its circumpolar range. One particular side of this diversity is the frequent occurrence of two or more distinct charr morphs in the same lake. This polymorphism has been termed ‘the charr problem’. Similar cases are found in other postglacial fishes, but not with the extent and diversity as with the arctic charr. This review first treats the classical case, pioneered in an advanced way by Winifred Frost, of autumn and winter spawning charr in Windermere, England, and three other cases that have received much research interest in recent years: Thingvallavatn, Iceland; Loch Rannoch, Scotland; and Fjellfrøsvatn, Norway. Then a special kind of sympatry with one morph living permanently in the profundal zone, known from a few lakes in Europe, Russia and Canada and unique for arctic charr among postglacial fishes, is reviewed. Among them is a recently discovered charr at 450 m depth in Tinnsjøen, Norway, one of the few very deep lakes in the world. With examples, the concluding discussion focuses on the variation of arctic charr polymorphisms which extends from early stages of ecological segregation to cases of reproductive isolation and speciation; and on models to explain the charr problem. The exceptional diversity of arctic charr provides a unique potential for further progress in studies on ecologically driven evolution within the frames of modern theory of developmental plasticity, adaptive radiation and adaptive speciation.

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.

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.

Takvatn through 20 years: long-term effects of an experimental mass removal of Arctic charr, Salvelinus alpinus, from a subarctic lake

Between 1984 and 1989, the experimental removal of 31 tons (666 000 fish) of stunted Arctic charr, Salvelinus alpinus, from Takvatn in northern Norway, had strong effects on the populations of Arctic charr, brown trout, Salmo trutta, and three-spined sticklebacks, Gasterosteus aculeatus. The littoral catch per unit effort (CPUE) of charr had decreased by 90% in 1990 and then increased to about 50% of the initial level by 1994 while the pelagic CPUE had decreased to zero. Growth in both charr and trout greatly improved when the charr density had decreased, and large fish of both species appeared in the catches. These large fish became predators on small charr in the littoral zone. The incidence of trout increased from below 1% to 15% from 1988 to 1999 after a brief peak at 30% in 1992 and 1993. The charr population attained a bimodal size distribution and did not return to the stunted state during the 10 years following the intensive fishing period. The mass removal experiment showed that it is possible to change the structure of a charr population by intensive fishing. Predation on small charr from cannibals and large trout was probably essential for maintaining the new population structure. An increase in the growth of young charr from 1995 to 1997 was related to a high consumption of Daphnia and Eurycercus. Rapid changes in the growth of charr followed the density fluctuations in sticklebacks, which show large annual variations in this system; the rapid changes in charr growth were probably caused by variations in the competition intensity for cladoceran prey between young charr and sticklebacks. Twenty years of data has provided important information, but even more time is needed to follow the long-term trends in northern lakes such as Takvatn.

Within-sample variabilities in stomach contents weight of fish — implications for field studies of consumption rate

Stomach contents weights in a study of Arctic charr (Salvelinus alpinus L.) were not normally distributed. This may have implications for the field estimation of consumption rates and also indicates that a more extensive handling of stomach content data is needed. Fish were sampled every 3 h throughout 24 h periods. The stomach contents showed very large variations within each 3 h sample. The distributions of stomach contents weights were skewed to the right, and the variance was dependent on the sample mean, with the variance to mean ratio equaling 1. Although mean values of stomach contents weight seemed to show diurnal variations, there were usually no significant differences between consecutive samples taken throughout each 24 h period, due to the large range in stomach contents weight within each sample. These findings question the accuracy of consumption estimates obtained using the method of Elliott & Persson (1978) and other related methods based on differences between consecutive sample mean values. Due to the large within-sample variations found in the current study, the method of Bajkov (1935) modified by Eggers (1979) is considered to give more reliable consumption estimates than the Elliott & Persson method. The generality of the findings is discussed, and it is pointed out that greater attention should be focused on within-sample variabilities in fish stomach contents studies.

New parasites and predators follow the introduction of two fish species to a subarctic lake: implications for food-web structure and functioning

Introduced species can alter the topology of food webs. For instance, an introduction can aid the arrival of free-living consumers using the new species as a resource, while new parasites may also arrive with the introduced species. Food-web responses to species additions can thus be far more complex than anticipated. In a subarctic pelagic food web with free-living and parasitic species, two fish species (arctic charr Salvelinus alpinus and three-spined stickleback Gasterosteus aculeatus) have known histories as deliberate introductions. The effects of these introductions on the food web were explored by comparing the current pelagic web with a heuristic reconstruction of the pre-introduction web. Extinctions caused by these introductions could not be evaluated by this approach. The introduced fish species have become important hubs in the trophic network, interacting with numerous parasites, predators and prey. In particular, five parasite species and four predatory bird species depend on the two introduced species as obligate trophic resources in the pelagic web and could therefore not have been present in the pre-introduction network. The presence of the two introduced fish species and the arrival of their associated parasites and predators increased biodiversity, mean trophic level, linkage density, and nestedness; altering both the network structure and functioning of the pelagic web. Parasites, in particular trophically transmitted species, had a prominent role in the network alterations that followed the introductions.