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.     

Is coexistence mediated by microhabitat segregation? An in-depth exploration of a fish invasion

Vendace Coregonus albula has invaded the subarctic Pasvik hydrosystem, northern Norway and Russia, after being translocated from its native Finnish range into Lake Inari (Finland), upstream of Pasvik. The development of the invader and the effects on two native lake communities in the upstream and downstream part of the hydrosystem, respectively, have been monitored since 1991. In the upstream lake location, the invader relegated the native whitefish Coregonus lavaretus from the pelagic habitat within a few years. An expected similar development in the downstream location was, however, not observed. Data are presented on the diet and microhabitat use (vertical pelagic zone) for three types of replicated situation: (1) the invader v. the native species, (2) upstream v . downstream lakes and (3) early and late invasion stages, to explore the outcome of the competitive interactions. The use of different microhabitats available within the deeper pelagic zone in the downstream location was observed to have delayed the negative effects on native whitefish, possibly enabling a persistent coexistence between the two species in the pelagic zone of this lake.     

Interactions between invading benthivorous fish and native whitefish in subarctic lakes

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REGULAR PAPERS Habitat distribution and size structure in freshwater fish communities: effects of vendace on interactions between perch and roach

In a study of habitat selection and size structure in three fish species in 115 Swedish lakes, roach Rutilus rutilus used the pelagic habitat to a lesser extent and were lower in pelagic biomass in the presence of vendace Coregonus albula. Size distributions of roach were skewed toward larger sizes in lakes with vendace, although a similar pattern of size-dependent habitat use was found regardless of lake group. In disagreement with predictions, vendace did not affect relative perch Perca fluviatilis biomass, size structure, or the proportion of large perch. However, a higher growth for large perch indicated higher per capita energy intake in lakes with vendace.     

Thermal habitat segregation among morphotypes of whitefish (Coregonus lavaretus: Salmonidae) and invasive vendace (C. albula): a mechanism for co‐existence?

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Niche segregation of coexisting Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) constrains food web coupling in subarctic lakes

1. Generalist fish species are recognised as important couplers of benthic and pelagic food‐web compartments in lakes. However, interspecific niche segregation and individual specialisation may limit the potential for generalistic feeding behaviour. 2. We studied summer habitat use, stomach contents and stable isotopic compositions of the generalist feeder Arctic charr coexisting with its common resource competitor brown trout in five subarctic lakes in northern Norway to reveal population‐level and individual‐level niche plasticity. 3. Charr and trout showed partial niche segregation in all five lakes. Charr used all habitat types and a wide variety of invertebrate prey including zooplankton, whereas trout fed mainly on insects in the littoral zone. Hence, charr showed a higher potential to promote habitat and food‐web coupling compared to littoral‐dwelling trout. 4. The level of niche segregation between charr and trout and between pelagic‐caught and littoral‐caught charr depended on the prevailing patterns of interspecific and intraspecific resource competition. The two fish species had partially overlapping trophic niches in one lake where charr numerically dominated the fish community, whereas the most segregated niches occurred in lakes where trout were more abundant. 5. In general, pelagic‐caught charr had substantially narrower dietary and isotopic niches and relied less on littoral carbon sources compared to littoral‐caught conspecifics that included generalist as well as specialised benthivorous and planktivorous individuals. Despite the partially specialised planktivorous niche and thus reduced potential of pelagic‐dwelling charr to promote benthic–pelagic coupling, the isotopic compositions of both charr subpopulations suggested a significant reliance on both littoral and pelagic carbon sources in all five study lakes. 6. Our study demonstrates that both interspecific niche segregation between and individual trophic specialisation within generalist fish species can constrain food‐web coupling and alter energy mobilisation to top consumers in subarctic lakes. Nevertheless, pelagic and littoral habitats and food‐web compartments may still be highly integrated due to the potentially plastic foraging behaviour of top consumers.     

Discrete foraging niches promote ecological, phenotypic, and genetic divergence in sympatric whitefish (Coregonus lavaretus)

Natural populations often vary in their degree of ecological, morphological and genetic divergence. This variation can be arranged along an ecological speciation continuum of increasingly discrete variation, with high inter-individual variation at one end and well defined species in the other. In postglacial fishes, evolutionary divergence has commonly resulted in the co-occurrence of a pelagic and a benthic specialist. We studied three replicate lakes supporting sympatric pelagic and benthic European whitefish (Coregonus lavaretus (L.)) morphs in search for early signs of possible further divergence into more specialized niches. Using stomach content data (recent diet) and stable isotope analyses (time-integrated measure of trophic niche use), we observed a split in the trophic niche within the benthic whitefish morph, with individuals specializing on either littoral or profundal resources. This divergence in resource use was accompanied by small but significant differences in an adaptive morphological trait (gill raker number) and significant genetic differences between fish exploiting littoral and profundal habitats and foraging resources. The same pattern of parallel divergence was found in all three lakes, suggesting similar natural selection pressures driving and/or maintaining the divergence. The two levels of divergence (a clear and robust benthic – pelagic and a more subtle littoral – profundal divergence) observed in this study apparently represent different stages in the process of ecological speciation.     

Species assignment and population genetic studies of Gran Paraná pejerrey (<Emphasis Type="Italic">Odontesthes</Emphasis> sp., Atheriniformes,Atherinopsidae) from La Plata Basin in South America

Since 2013, the pelagic zone of Upper Lake Constance (ULC) has been subject to a massive invasion of the non-native three-spined stickleback (Gasterosteus aculeatus Linnaeus, 1758). Data from monthly monitoring of pelagic whitefish (Coregonus wartmanni Bloch, 1784) were used to compare weight-at-age and abundance of pelagic whitefish for years before (1997–2012) and after the invasion (2013–2015). Growth and abundance of pelagic whitefish is shown to be heavily influenced by stickleback presence. Mean autumn weight-at-age of whitefish decreased by 33.3% after the invasion took place and a significant decline in autumn CPUE in otherwise unfished cohorts of the population was also recorded. The results imply direct effects of stickleback presence on pelagic whitefish, including interspecific competition for food leading to reduced growth and survival, and predation of eggs and larvae, hampering recruitment. These observations coincide with a sharp decline in whitefish yield. In conclusion, this study shows that the invasion of stickleback has substantially altered the pelagic fish community of ULC, with severe consequences for commercial fisheries.     

Habitat use and diet of sympatric Arctic charr (<Emphasis Type="Italic">Salvelinus alpinus</Emphasis>) and whitefish (<Emphasis Type="Italic">Coregonus lavaretus</Emphasis>) in five lakes in southern Norway: not only interspecific population dominance?

Stable coexistence of Arctic charr and whitefish does occur in a number of native lake fish communities in Scandinavia. Even so, whitefish introductions into Arctic charr lakes have resulted in serious decline and possibly local extinction of Arctic charr. In this article, we analyze the habitat use and diet of the two species in five Norwegian lakes differing in basin shape and environmental conditions. In two of the lakes, both species are native, and appear to live in a relatively stable coexistence. Here, whitefish mainly occupy the littoral and upper pelagic zone, while Arctic charr live in the deeper habitats. Diets are generally quite different in terms of the zooplankton species eaten. In the three other lakes, either whitefish or both species have been introduced. In the shallowest lake, habitat segregation is similar to that seen in the pristine lakes, although Arctic charr appears to be on the brink of extinction. In the remaining two lakes, however, Arctic charr dominates, and occurs in higher numbers than whitefish in all the habitats. Our observations indicate that coexistence of the two species in oligotrophic and relatively pristine lakes requires an extensive profundal zone to serve as a refugium for Arctic charr. If the littoral zone is rendered inaccessible or unprofitable for whitefish due to dominance of a third competitor or predator, or as a result of lake regulation, then Arctic charr may be the dominant species.     

Food‐web structure and mercury dynamics in a large subarctic lake following multiple species introductions

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Species introduction promotes hybridization and introgression in Coregonus: is there sign of selection against hybrids?

Species introductions are considered one of the major drivers of biodiversity loss via ecological interactions and genetic admixture with local fauna. We examined two well-recognized fish species, native whitefish (Coregonus lavaretus) and introduced vendace (Coregonus albula), as well as their morphological hybrids in a single lake to test for selection against hybrids and backcrosses in the wild. A representative random subsample of 693 individuals (27.8%) was taken from the total catch of coregonids. This subsample was examined with the aim to select c. 50 individuals of pure whitefish (n = 52), pure vendace (n = 55) and putative hybrid (n = 19) for genetic analyses. The subsequent microsatellites and mitochondrial (mt) DNA analyses provided compelling evidence of hybridization and introgression. Of the 126 fish examined, four were found to be F(1) , 14 backcrosses to whitefish and seven backcrosses to vendace. The estimates of historical gene flow suggested higher rates from introduced vendace into native whitefish than vice versa, whereas estimates of contemporary gene flow were equal. Mitochondrial introgression was skewed, with 18 backcrosses having vendace mtDNA and only three with whitefish mtDNA. Hybrids and backcrosses had intermediate morphology and niche utilization compared with parental species. No evidence of selection against hybrids or backcrosses was apparent, as both hybrid and backcross growth rates and fecundities were high. Hybrids (F(1) ) were only detected in 2 year-classes, suggesting temporal variability in mating between vendace and whitefish. However, our data show that hybrids reached sexual maturity and reproduced actively, with backcrosses recorded from six consecutive year-classes, whereas no F(2) individuals were found. The results indicate widespread introgression, as 10.8% of coregonids were estimated to be backcrosses.     

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.     

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.     

Effects of invading vendace (Coregonus albula L.) on species composition and body size in two zooplankton communities of the Pasvik River System, northern Norway

Species composition and body-size distribution were studiedin the crustacean zooplankton communities of two limnologicallysimilar lake localities situated 50 km apart in the Pasvik RiverSystem, northern Norway. A recent invasion and successive downstreamexpansion of vendace (Coregonus albula), a specialized zooplanktivorousfish, allowed comparisons between sites with different predationpressures. Vendace had established a high population densityand was the dominant fish species in the pelagic of the upperlocality, but had just invaded the lower locality with a smallnumber of individuals. Whitefish (Coregonus lavaretus), a closelyrelated but less specialized zooplanktivore species, dominatedthe native fish community of both lakes. The zooplankton communityof the upper locality was dominated in June and August by Bosminalongirostris, the smallest zooplankton species represented inthe water course, and in September by Daphnia cristata. Thelower locality was dominated by the larger Holopedium gibberumand Eudiaptomus graciloides in June, by D.cristata in August,and by D.cristata and B.longirostris in September. The meanbody size of the three most abundant cladoceran species wassignificantly smaller in the upper locality, compared to thelower locality. It was concluded that the invasion and establishmentof a dense vendace population in the upper locality had increasedthe predation pressure in the pelagic, resulting in a reductionof body size and a shift towards smaller species in the zooplanktoncommunity.     

Habitat coupling in a large lake system: delivery of an energy subsidy by an offshore planktivore to the nearshore zone of Lake Superior

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Diel and seasonal habitat and food segregation of three sympatric Coregonus lavaretus forms in a subarctic lake

Habitat use of three sympatric whitefish Coregonus lavaretus forms was determined using hydroacoustics, pelagic trawling and epibenthic gillnetting in the subarctic Lake Muddusjärvi during the day and night in June, August and September. Whitefish constituted 97% of the numerical catches and whitefish with high number of gill rakers (DR) were the most abundant whitefish form. Forms with low numbers of gill rakers used only epibenthic habitats during both the day and night in all study periods: large whitefish with low numbers of gill rakers (LSR) dwelled mainly at depths 0–10 m, whereas small whitefish with low numbers of gill rakers (SSR) used deeper (>10 m) habitats. LSR and SSR whitefish consumed mainly benthic macroinvertebrates during all study occasions. The planktivorous DR whitefish used both epibenthic and pelagic habitats, but vertical habitat selection varied both over time of day and season. In June, when light intensity was continuously high, DR whitefish did not perform diel vertical migrations. In August and September, when dark nights were distinguishable, DR whitefish ascended from the bottom to the pelagic at dusk to feed on zooplankton, and descended at dawn. DR whitefish used pelagic habitats only at the lowest light intensities during the night, which was probably related to the high predation risk from brown trout Salmo trutta .     

Seasonal depletion of resources intensifies trophic interactions in subarctic freshwater fish communities

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Ontogenetic changes in habitat use by whitefish,Coregonus lavaretus

Synopsis The whitefish, Coregonus lavaretus, in the lake Mj?sa exhibited two niche changes during their life cycle. Juveniles (< 25 cm body length) were confined to the shallow (0–30 m) epibenthic zone. Medium sized whitefish (25–35 cm body length) expanded their habitat use to include the deep (30–90 m) epibenthic zone as well as the pelagic zone. From a body length of 35 cm, habitat use was restricted to the deep epibenthic zone. Small fish in the shallow epibenthic zone ate small and medium-sized prey (zooplankton, insect larvae and surface insects). Medium-sized fish in this zone were in addition feeding on the larger amphipod, Pallasea quadrispinosa. In the pelagic zone, the diet of medium-sized whitefish was dominated by zooplankton, although some larger prey like surface insects and age-0 smelt, Osmerus eperlanus, were also eaten. In the deep epibenthic zone, the diet of both medium-sized and large (< 35 cm) whitefish consisted mainly of the large prey P. quadrispinosa.     

Long-term responses of zooplankton to invasion by a planktivorous fish in a subarctic watercourse

1. Introduced or invading predators may have strong impacts on prey populations of the recipient community mediated by direct and indirect interactions. The long-term progression of predation effects, covering the invasion and establishment phase of alien predators, however, has rarely been documented.
2. This paper documents the impact of an invasive, specialized planktivorous fish on its prey in a subarctic watercourse. Potential predation effects on the crustacean plankton, at the community, population and individual levels, were explored in a long-term study following the invasion by vendace ( Coregonus albula ).
3. Over the 12-year period, the density and species richness of zooplankton decreased, smaller species became more abundant and Daphnia longispina , one of the largest cladocerans, was eliminated from the zooplankton community.
4. Within the dominant cladocerans, including Daphnia spp., Bosmina longispina and Bosmina longirostris , the body size of ovigerous females and the size at first reproduction decreased after the arrival of the new predator. The clutch sizes of Daphnia spp. and B. longirostris also increased.
5. Increased predation pressure following the vendace invasion induced many effects on the crustacean zooplankton, and we document comprehensive and strong direct and indirect long-term impacts of an introduced non-native predator on the native prey community.     

Inter- and intra-morph patterns in helminth communities of sympatric whitefish morphs

Infection patterns of trophically transmitted helminth parasites were compared with feeding ecology in two sympatric whitefish Coregonus lavaretus morphs from two lake systems in northern Norway. In both lakes, the pelagic morph was an obligate zooplanktivore, while the benthic morph utilized both the benthivore and zooplanktivore trophic niches. The differences in niche utilization between the two morphs were associated with differences in trophic morphology (gill raker numbers), suggesting that they were genetically dissimilar and reproductively isolated. The benthic morph had the highest number of helminth species, probably because they exhibited a broader niche width compared to the pelagic morph. In both lakes, the species composition and intensities of helminths reflected the trophic diversification of the whitefish ecotypes with respect to different habitat choice (benthic v . pelagic) and dietary specialization (benthivore v . zooplanktivore feeding strategies within the benthic whitefish morph). Zooplanktivorous fish from both morphs acquired parasites mainly from pelagic copepods and in almost equal quantities. The benthivore feeders within the benthic morph had the highest proportion of parasites with transmission stages from benthic organisms. Host feeding behaviour seemed to be a major determinant of the helminth community structure, and helminths appeared to be useful indicators of long-term trophic specialization of whitefish ecotypes.