Roar Kristoffersen

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.

AGGREGATION OF HELMINTHS: THE ROLE OF FEEDING BEHAVIOR OF FISH HOSTS

Individual Arctic charr (Salvelinus alpinus) from Fjellfrøsvatn, northern Norway, could be categorized by their stomach contents as zooplanktivores or benthivores. Feeding specialization among these fish was evident from negative correlations between helminths transmitted by pelagic copepods (Diphyllobothrium dendriticum and D. ditremum) and those transmitted by the benthic amphipod Gammarus lacustris (Cystidicola farionis and Cyathocephalus truncatus). Occurrences of parasite species acquired from the same types of invertebrate were positively correlated in the fish. Strong relationships among habitat use, diet, and helminth infections among the Arctic charr indicated persistent foraging patterns involving long-term habitat use and feeding specialization. The distribution of all parasite species was highly aggregated in the fish samples, measured by the exponent k of the fitted negative binomial distributions (range: 0.5–7.5) and the variance-to-mean ratios (s2/x̄, range: 5–85). Charr specializing on either copepods or Gammarus predominantly contributed to high-intensity class intervals within the overall frequency distributions of the corresponding parasite species. Such fish had low infection intensities of helminths transmitted by other prey organisms. The detailed analyses of the parasite frequency distributions for fish with different habitat or feeding preferences evidently show how heterogeneity in trophic behavior contributes strongly to the commonly observed aggregation of helminths among hosts under natural conditions.

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.

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.

Seasonal dynamics and persistence of Gyrodactylus salaris in two riverine anadromous Arctic charr populations

The seasonal occurrence of the monogenean ectoparasite Gyrodactylus salaris Malmberg infecting Arctic charr, Salvelinus alpinus (L.) in the two rivers Skibotnelva and Signaldalselva in northern Norway was studied in the period from autumn 2003 to autumn 2005. Skibotnelva has been infected with the parasite since 1979, and treated with rotenone twice. Most likely resident Arctic charr avoided the rotenone treatment in small tributary streams, and thus was the source of the repeated re-infection of this river. G. salaris was first recorded in Signaldalselva in the year 2000 and it is still untreated. Unlike Atlantic salmon, which is highly susceptible to G. salaris, Arctic charr can display a wide range of host-responses to G. salaris infections. Arctic charr were sampled by electro fishing with a total sample of 681 Arctic charr. The results from this study demonstrate an evident seasonal dynamic in G. salaris infection in charr in both rivers. Parasite intensities fluctuated with the rise and fall in temperature through the year, with an autumn high and spring low. There was a significantly lower prevalence and mean intensity of G. salaris in Skibotnelva than in Signaldalselva. There were also a lower prevalence and intensity of G. salaris in the older than in the youngest charr. The different history of infection and treatment in the two rivers might be the underlying cause of these observed dissimilarities. The current study indicates that Arctic charr is a good natural host for G. salaris.

Food borne parasites as indicators of trophic segregation between Arctic charr and brown trout

The habitat and diet choice and the infection (prevalence and abundance) of trophically transmitted parasites were compared in Arctic charr and brown trout living sympatrically in two lakes in northern Norway. Arctic charr were found in all main lake habitats, whereas the brown trout were almost exclusively found in the littoral zone. In both lakes the parasite fauna reflected the niche segregation between trout and charr. Surface insects were most common in the diet of trout, but transmit few parasites, and accordingly the brown trout had a relatively low diversity and abundance of parasites. Parasites transmitted by benthic prey such as Gammarus and insect larva, were common in both salmonid host species. Copepod transmitted parasites were much more common in Arctic charr, as brown trout did not include zooplankton in their diets. Parasite species that may use small fish as transport hosts, were far more abundant in piscivorous fish, especially brown trout. The seasonal dynamics in parasite infection were also consistent with the developments in the diet throughout the year. The study demonstrates that the structure of parasite communities of charr and the trout is highly dependent on shifts in habitat and diet of their hosts both on an annual base and through the ontogeny, in addition to the observed niche segregation between the two salmonid species.

Ontogenetic dynamics of infection with Diphyllobothrium spp. cestodes in sympatric Arctic charr Salvelinus alpinus (L.) and brown trout Salmo trutta L.

The trophic niches of Arctic charr and brown trout differ when the species occur in sympatry. Their trophically transmitted parasites are expected to reflect these differences. Here, we investigate how the infections of Diphyllobothrium dendriticum and D. ditremum differ between charr and trout. These tapeworms use copepods as their first intermediate hosts and fish can become infected as second intermediate hosts by consuming either infected copepods or infected fish. We examined 767 charr and 368 trout for Diphyllobothrium plerocercoids in a subarctic lake. The prevalence of D. ditremum was higher in charr (61.5%) than in trout, (39.5%), but the prevalence of D. dendriticum was higher in trout (31.2%) than in charr (19.3%). Diphyllobothrium spp. intensities were elevated in trout compared to charr, particularly for D. dendriticum. Large fish with massive parasite burdens were responsible for the high Diphyllobothrium spp. loads in trout. We hypothesize that fish prey may be the most important source for the Diphyllobothrium spp. infections in trout, whereas charr predominantly acquire Diphyllobothrium spp. by feeding on copepods. Our findings support previous suggestions that the ability to establish in a second piscine host is greater for D. dendriticum than for D. ditremum.

Experiments to test if allopatric Salvelinus alpinus are suitable year-round hosts of Gyrodactylus salaris (Monogenea)

There has been an increased focus on Salvelinus alpinus as a potential long-term host to Gyrodactylus salaris and, here, both susceptibility to G. salaris and ability to sustain a parasite population seasonally, was tested using fry and parr of S. alpinus from the River Skibotnelva, northern Norway. Fry were highly susceptible. Gyrodactylus salaris survived on allopatric S. alpinus parr during the 5 month-long winter when water temperatures were c. 1 degrees C. Salvelinus alpinus fry also maintained a pulse of G. salaris infection for over 155 days from early May until autumn. Gyrodactylus salaris are thus able to reproduce and survive on S. alpinus for long periods and at low water temperatures. In spring, newly hatched fry of S. alpinus may serve as an important host to maintain a G. salaris metapopulation within a river system. The results suggest that S. alpinus are adequate long-term hosts of G. salaris independent of the presence of the co-occurring highly susceptible S. salar.

Wild and hatchery-reared landlocked Arctic charr, Salvelinus alpinus (L.), from Lake Takvatn, reared in fresh and sea water. Content of free amino acids and ninhydrin-positive substances

Abstract Wild and hatchery-reared landlocked Arctic charr were reared in fresh water, then acclimatized to full sea water strength over 2 weeks and reared for 60 days in sea water under identical conditions. Wild charr increased in weight from 100 to 140 g, hatchery-reared fish to 180 g. The mortality of wild fish was 30% and that of hatchery-reared none, after 60 days on-growing in sea water. Content of free amino acids and ninhydrin-positive substances in charr reared in fresh and sea water were determined in muscle, liver and serum. Muscle and liver accumulate different amino acids, glycine in muscle, and taurine in the liver. Fresh water charr had high and steady concentrations of glycine and taurine in serum. The concentrations of valine and leucine increased after gradual acclimatization to sea water, but thereafter remained unchanged in sea water. Blood serum osmolality increased during acclimatization to sea water.

Temporal changes and between-host variation in the intestinal parasite community of Arctic charr in a subarctic lake

The occurrence of trophically transmitted intestinal parasites in Arctic charr was analyzed from data material collected over two decades from Lake Takvatn, northern Norway. The main objectives were to investigate (i) between-year variation in parasite infracommunity composition, (ii) between-host variation in infracommunity composition, and (iii) possible interspecific interaction between parasites. The trematode Crepidostomum spp. and the cestode Eubothrium salvelini were most prevalent and had the highest mean intensities whereas the cestodes Proteocephalus sp. and Cyathocephalus truncatus were uncommon taxa. No clear indication of interspecific parasite interactions was evident. By far the most frequent infracommunity composition was E. salvelini co-existing with only Crepidostomum spp. Despite some indications of individual dietary specialization in Arctic charr, this strong species co-occurrence suggests that the majority of fish had been preying on the very different intermediate hosts of both parasites. Overall, even though some sporadic between-year variation was present, no long-term or cyclical variation in infracommunity composition and mean intensity was revealed, suggesting that the host–parasite system is in a steady state. Minor between-year variation was likely the result of changes in habitat and dietary utilization by Arctic charr, possibly linked to documented changes in the fish community structure of the lake.