Karl Øystein Gjelland

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.

Environmental influence on transmitter detection probability in biotelemetry: developing a general model of acoustic transmission

Summary 1. Environmental factors, such as wind, may have a strong influence on the detection probability and detection rate of acoustic telemetry tags. The effect of environmental factors may obscure biological effects and distort the interpretation of acoustic telemetry data. 2. This study was undertaken with fish internally tagged with acoustic transmitters containing depth sensors and monitored by an array of automatic receivers. The influence of environmental factors on the hourly detection rate was evaluated using environmental data from a nearby climate station. The signal detection probability was modelled within the framework of general theory of sound propagation in water. 3. Wind was found to have the strongest influence on the detection rate. Transmitter depth range and rain also contributed signif icantly to the variation in detection rate. 4. By modelling the attenuation coefficient as a function of wind speed, we show that the probability of detecting a free-swimming acoustically tagged animal can be successfully modelled using general sound propagation theory. 5. The approach of modelling detection probability as a function of the attenuation coefficient offers a wide applicability, as it implies a direct link between detection probability and physical characteristics of the water at the study site. Correcting for varying detection probability is in many cases extremely important to do, since rhythms in biological/behavioural factors are often confounded with environmental variables that influence detection probability (e.g. sea breeze, tide).

Predation by brown trout (Salmo trutta) along a diversifying prey community gradient

Predation has a fundamental role in aquatic ecosystems, but the relative importance of factors governing prey selection by predators remains controversial. In this study, we contrast five lakes of a subarctic watershed to explore how prey community characteristics affect prey selection and growth rate of the common top predator, brown trout (Salmo trutta). The lakes constitute a distinct gradient of different coregonid prey fish, ranging from monomorphic common whitefish (Coregonus lavaretus) to polymorphic whitefish co-occurring with vendace (Coregonus albula). The brown trout was a morph–species- and size-specific pelagic predator, selecting the small-sized, pelagic whitefish morph or vendace over the benthic whitefish morphs. In all lakes, the average prey size increased with predator size, but small-sized prey were also included in the diet of large predators. The selection of small-sized, pelagic prey fish appeared to be a favourable foraging strategy for the brown trout, yielding higher growth rates...

Speciation Reversal in European Whitefish (Coregonus lavaretus (L.)) Caused by Competitor Invasion

Invasion of exotic species has caused the loss of biodiversity and imparts evolutionary and ecological changes in the introduced systems. In northern Fennoscandia, European whitefish (Coregonus lavaretus (L.)) is a highly polymorphic species displaying adaptive radiations into partially reproductively isolated and thus genetically differentiated sympatric morphs utilizing the planktivorous and benthivorous food niche in many lakes. In 1993, Lake Skrukkebukta was invaded by vendace (Coregonus albula (L.)) which is a zooplanktivorous specialist. The vendace displaced the densely rakered whitefish from its preferred pelagic niche to the benthic habitat harbouring the large sparsely rakered whitefish. In this study, we investigate the potential influence of the vendace invasion on the breakdown of reproductive isolation between the two whitefish morphs. We inferred the genotypic and phenotypic differentiation between the two morphs collected at the arrival (1993) and 15 years after (2008) the vendace invasion using 16 microsatellite loci and gill raker numbers, the most distinctive adaptive phenotypic trait between them. The comparison of gill raker number distributions revealed two modes growing closer over 15 years following the invasion. Bayesian analyses of genotypes revealed that the two genetically distinct whitefish morphs that existed in 1993 had collapsed into a single population in 2008. The decline in association between the gill raker numbers and admixture values over 15 years corroborates the findings from the Bayesian analysis. Our study thus suggests an apparent decrease of reproductive isolation in a morph-pair of European whitefish within 15 years (≃ 3 generations) following the invasion of a superior trophic competitor (vendace) in a subarctic lake, reflecting a situation of “speciation in reverse”.

Invader population speeds up life history during colonization

We explore the long-term developments in population biology and life history during the invasion and establishment of the fish species vendace Coregonus albula in a subarctic watercourse by comparing life-history traits and molecular genetic estimates between the source and the colonist population. The two populations exhibited highly contrasting life-history strategies. Relative to the source population, the colonist population was characterized by slower somatic growth rates, earlier sexual maturation at smaller individual size, higher mortality rates and a shorter life span. The two populations could also be significantly discriminated by the genetic markers. Limited founder effects were detected from heterozygote deficit and reduced allelic richness in the colonist population, but both populations were associated with relatively high genetic diversity. The study reveals that the invasion into a new environment induced large changes in life-history strategy, with typical r-selected traits being more prominent in the colonist than in the source population. We discuss the mechanisms that may explain the observed life-history differences between the source and the colonist population, and argue that the accelerated life history of the colonist population represents an adaptive pioneer strategy aimed at fast population increase during colonization and establishment.

Invasion genetics of vendace (Coregonus albula (L.)) in the Inari-Pasvik watercourse: revealing the origin and expansion pattern of a rapid colonization event

Species invasions can have wide-ranging biological and socio-economic effects and are generally unwanted by legislation. Identification of the source population as well as the ecology and genetics of both the invader population and the receiving community is of crucial importance. The rapid invasion of a small coregonid fish vendace (Coregonus albula) in a major northern European subarctic watercourse has resulted in a labile ecological situation in the receiving community. The ecological impact of the invasion has been thoroughly documented, but the genetics of the invasion remains to be explored. We analyzed the genetic diversity and divergence patterns among the two possible source populations from southern Finnish Lapland and three colonists populations within the Inari-Pasvik watercourse using ten microsatellite loci in order to (i) identify the most likely source of the invasion, (ii) reveal the dispersal pattern and genetic structure of the secondary expansion, and (iii) to investigate whether the initial introduction and the secondary expansion were associated with founder effects. We revealed that repeated translocation of vendace from Lake Sinettäjärvi into a tributary lake of L. Inari in 1964–1966 is the most plausible source for the invasion. Both the initial introduction and the secondary expansion were found not to be associated with significant founder effects. The secondary expansion followed a stepping stone pattern and the source and colonist populations of this expansion have undergone rapid genetic divergence within a period of 15–35 years (ca. 8–17 generations). The rapid divergence may be contributed to lack of gene flow among the source and colonist populations due to the extensive hydroelectric damming in the watercourse. Multiple introductions and substantial genetic variation in combination with the boom-and-bust population development of the species thus likely counteracted the founder effects as well as fueled the rapid establishment and expansion of this species within the Inari-Pasvik watercourse.

A planktivorous specialist turns rapacious: piscivory in invading vendace Coregonus albula

Following invasion and establishment in a subarctic watercourse, small-sized vendace Coregonus albula were surprisingly found to include fish prey in their diet. The zooplankton resource was severely depleted and only made a minor dietary contribution.

A specialised cannibalistic Arctic charr morph in the piscivore guild of a subarctic lake

Arctic charr (Salvelinus alpinus) is generally considered an inferior piscivore compared to brown trout (Salmo trutta). However, we demonstrate that a recently described profundal spawning piscivore PP-morph of Arctic charr in a subarctic lake has evolved typical piscivore traits (i.e. large jaws, robust skulls), similar to the co-occurring brown trout but different from the sympatric littoral spawning omnivore LO-morph of Arctic charr. A few large-sized LO-morph, most large-sized PP-morph and trout constitute the piscivore guild, but they differed in prey fish selection. Of the fish-eating deep-water PP-morph, 52% had consumed charr, with a peak during the polar night period (68–80% in November–January). In contrast, the LO-morph and trout were mainly (~90%) inter-specific piscivores eating stickleback. Consequently, the PP-morph was the main charr predator and started to feed on fish at a smaller size and took larger prey relative to their body length than the trout. Stable isotope (δ13C, δ15N) ratios reflect the dietary specialisations in the upper-water (LO-morph, trout) and the deep-water (PP-morph) environments, besides the piscivorous behaviour of the PP-morph and trout. The existence of a shallow-water piscivore (trout) may explain the origin of this PP-morph, taking benefit of underutilised resources of small-sized Arctic charr in the deep-water habitat.

Contrasting population and life history responses of a young morph-pair of European whitefish to the invasion of a specialised coregonid competitor, vendace.

Invasions of non-native species represent a global problem of great scientific interest. Here we study in detail the response in population and life history characteristics of closely related native species, with divergent habitat preferences, that are impacted by an invading species over a sufficient time period to allow a new stable state to become established. A time series of 20 years starting at the first occurrence of the invader (vendace Coregonus albula (L.)) allows exploration of the long term population and life history response of two ecologically, morphologically, and genetically different native sympatric morphs (DR- and SR-) of congeneric whitefish C. lavaretus (L.). The whitefish morphs are taxonomically equally related to the invading vendace, but only the planktivorous DR-whitefish share its pelagic niche. We would expect that the ecological differences between the whitefish morphs may be used as a predictor of competitive effects. Vendace exhibited an initial boom-and-bust development, and has continued to fluctuate in density. The responses of the pelagic DR-whitefish were: i) an immediate habitat shift, ii) a subsequent population decline caused by increased annual mortality, and iii) a new stable state at a lower density and apparently relaxed competition. The ecologically more distant benthivorous SR-whitefish also showed significant, but a much more limited response during this process, indicating damped indirect interactions through the food-web. This long-term case-study found that in two native eco-species equally related to the invader, only one of the eco-species was highly affected. Direct competition for resources is obviously important for species interactions, whereas the taxonomic relatedness per se seems to offer little predictive power for invasion effects.

Resource partitioning between pelagic coregonids in a subarctic watercourse following a biological invasion

If a pelagic ecosystem is invaded by an efficient planktivorous fish, competition theory predicts that habitat and/or diet segregation should occur if the invader and native planktivores are to co-exist. We compared the diet and pelagic habitat use between invading vendace (Coregonus albula) and two native morphs of whitefish (Coregonus lavaretus) in three subarctic lakes located within the same watercourse. No clear vertical habitat segregation was found as vendace prevailed throughout the entire depth range. Zooplankton generally dominated the diet of all three coregonids. No obvious resource partitioning was found in two of the lakes, while in the third lake the diets of whitefish and vendace differed significantly from each other. Here, the predominant pelagic whitefish morph fed extensively on water mites whereas vendace mainly consumed surface insects and to some extent small fish, apparently as a result of highly depleted zooplankton resources. The results suggest that various levels of resource depletion following the vendace invasion caused the observed differences between the lakes, indicating that different levels of resource partitioning were associated with dissimilar competition intensities. At present, coexistence of these congeneric species seems to be possible, although the vendace invasion has apparently resulted in a challenging and unstable situation for the planktivorous whitefish.