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.     

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.     

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.     

The food preference of vendace (Coregonus albula) in South Swedish forest lakes including the predation effect on zooplankton populations

Vendace (Coregonus albula, L.) is in the southern part of its distribution area during the summer period restricted to hypolimnion. Food is dominated by Bosmina coregoni and Daphnia sp. The selectivity index is highest for large cladocerans and large copepods.The pelagic crustacean fauna in vendace lakes is dominated by small species like Bosmina coregoni, Daphnia cristata, Thermocyclops and Mesocyclops sp. and Eudiaptomus sp. During periods of maximum vendace abundance only Bosmina coregoni is left of the cladocerans, while the abundance of cyclopoid copepods increases.     

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.     

Zooplankton composition and distribution off the coast of Galicia, Spain

During June and September 1984, zooplankton samples were collectedwith other hydrographic and biological data along the Galiciancoast (NW of Spain). In June copepods contributed {small tilde}60%to the total zooplankton community, with larvaceans, siphonophoresand cladocerans also abundant. In September >90% of the zooplanktonsampled were copepods. The dominant species of copepods in bothJune and September were Acartia clausi, Paracalanus parvus andTemora longicornis. The meroplankton was dominated by echinoderms,bryozoans, barnacle larvae and bivalve larvae. In June the averagezooplankton biomass was 31.08 mg C m^–3; the Septemberaverage was 41.69 mg C m^–3. The relationship between theslopes of the regression equations (biomass versus abundance)suggests that the zooplankton assemblage in June was composedby larger animals than in September. The major concentrationof zooplankton was between 0 and 50 m, with both June and Septemberdaytime surface samples having 6–7 times the amount oforganisms than the lower water column (50–100 m). Therewere no distinct differences in total zooplankton abundancesat the inshore and offshore stations; however, the inshore stationsoften had a higher percentage of meroplankton than the offshorestations. In June zooplankton abundance at the northern transectsand the western transects was similar. In September there weregreater concentrations of zooplankton in the western Galicianshelf as compared with the northern shelf. These differencesin the horizontal distribution of the zooplankton were relatedto upwelling events.     

The vertical distribution of fish in the open water area of a deep temperate mesotrophic lake assessed by hydroacoustics and midwater trawling

The fish stock of a deep temperate, mesotrophic lake was sampled at different depths using a fixed‐frame fry trawl, during two nights in mid‐September 2009. Additionally, horizontal and vertical hydroacoustics were used simultaneously to evaluate fish abundance and biomass estimates obtained by the trawl. Roach Rutilus rutilus and smelt Osmerus eperlanus were the dominant species of young‐of‐the‐year (YOY) fish in the trawl catches from the surface layers (0–9 m). Bleak Alburnus alburnus dominated the catch of older fish in the upper part of the surface profile (0–6 m). Around the thermocline (9–13 m) smelt dominated the catches of both the YOY and older fish. Beneath the thermocline (13–36 m) vendace Coregonus albula dominated the catch of YOY fish, and smelt was the only species of older fish in the trawl catches. Species composition, abundance and biomass of the YOY and older fish were heterogeneous throughout the depth profiles of the lake, but only abundance differed significantly between the layers. The hydroacoustics gave relatively similar estimates of abundance and biomass to those obtained by the trawl in all the depths sampled. Our results indicate that there is a clear separation of small fish of different species along the vertical profile of a deep temperate lake during the night, and an unequal vertical distribution of fish abundance and biomass. The similarity of the trawl and hydroacoustics estimates of abundances and biomass indicated that the trawl sampling did not cause important avoidance reactions of small fish during the night in this deep temperate lake (© 2012 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Food-web manipulations influence grazer control of phytoplankton growth rates in Lake Michigan

Stocking piscivorous salmonids in Lake Michigan produced dramaticalterations in food-web structure, including higher numbersof large-bodied zooplankton (especially Daphnia pulicaria),lower summer chlorophyll concentrations and increased watertransparency. Experimental determinations of epilimnetic phytoplanktongrowth rates and of zooplankton grazing rates indicate thatherbivorous zooplankton controlled algal dynamics during thesummer of 1983 because grazers occupied the surface waters throughoutthe day. In 1985, however, both large- and small-bodied Daphniamade approximately equal contributions to total grazer biomass,and all grazers displayed pronounced diel vertical migrations,visiting epilimnetic waters only at night. This prohibited zooplanktonfrom controlling algal dynamics because grazing losses did notexceed phytoplankton growth rates. The changes in zooplanktoncommunity composition and behavior observed in summer 1985 probablyresulted from increased predation by visually orienting planktivorousfish, especially bloater chub (Coregonus hoyi). Effects of food-webmanipulations on phytoplankton dynamics were evident only duringJuly and August. During spring and early summer copepods dominateLake Michigan's zooplankton community. Owing to their smallbody size, copepods are less susceptible to fish predation andexhibit much lower filtering rates than Daphnia. Variabilityin zooplanktivorous fish abundance probably has little effecton phytoplankton dynamics during spring and early summer.     

Pelagic fish and zooplankton species assemblages in relation to water mass characteristics in the northern Bering and southeast Chukchi seas

This research explores the distributions and community composition of pelagic species in the sub-Arctic and Arctic waters of the northern Bering and central and southern Chukchi seas during September 2007 by linking pelagic zooplankton and fish assemblages to water masses. Juvenile saffron cod (Eleginus gracilis), polar cod (Boreogadus saida), and shorthorn sculpin (Myoxocephalus scorpius) were most abundant in warm, low salinity Alaska Coastal Water (ACW) of the central Chukchi Sea, characterized by low chlorophyll, low nutrients, and small zooplankton taxa. Adult Pacific herring (Clupea pallasii) were more abundant in the less stratified Bering Strait waters and in the colder, saltier Bering Shelf Water of the northern Bering and southern Chukchi seas, characterized by high chlorophyll, high nutrients, and larger zooplankton taxa. Juvenile pink (Oncorhynchus gorbuscha) and chum (O. keta) salmon were most abundant in the less stratified ACW in the central Chukchi Sea and Bering Strait. Abundances of large zooplankton were dominated by copepods (Eucalanus bungii, Calanus glacialis/marshallae, Metridia pacifica) followed by euphausiids (juvenile Thysanoessa raschii and unidentified taxa), whereas small zooplankton were dominated by bivalve larvae and copepods (Centropages abdominalis, Oithona similis, Pseudocalanus sp.). Pelagic community composition was related to environmental factors, with highest correlations between bottom salinity and large zooplankton taxa, and latitude and fish species. These data were collected in a year with strong northward retreat of summer sea ice and therefore provide a baseline for assessing the effects of future climate warming on pelagic ecosystems in sub-Arctic and Arctic regions.     

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 changes in zooplankton composition in the Barents Sea, with special attention to Calanus spp. (Copepoda)

The Barents Sea is an important area with respect to fisheriesresources (i.e. capelin and cod). In May, June and August 1981zooplankton biomass was measured along a transect at 30°E,from the ice border southwards. A maximum was recorded in Atlanticwater by the end of June (>100 g wet weight m^–2 InAugust the biomass values were relatively low south of the Polarfront and increased northwards into Arctic water (–50g m^–2 The species composition was influenced by the distributionof cold Arctic water and warmer Atlantic water. The zooplanktonwas dominated by the copepods Calanus finmarchicus and C. glacialis;the former is regarded as an Atlantic species and C. glacialisas an Arctic species.     

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

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Rapidly changing life history during invasion

The fish species vendace ( Coregonus albula ) invaded the sub-arctic Pasvik watercourse during the second half of the 1980s, and became the dominant pelagic species in the upstream part of the watercourse within a few years. Life history traits of the pioneer population of vendace in Pasvik were recorded from 1991–2000. A rapid increase in population density in the upstream part of the watercourse was accompanied by decreased growth rates, decreased fecundity and a reduced size at first maturation. The downstream part of the watercourse showed a similar, but delayed, change in life history traits compared to the upstream part. The study documents great life history variability of a non-native fish species entering a new environment. We discuss two co-acting explanations for the observed patterns: (i) a density dependent response mediated by food depletion; and (ii) a pioneer strategy that allocates resources to favour reproduction at early developmental stages with a high number of offspring, trading off growth and size of offspring.     

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.     

Interactions between invading benthivorous fish and native whitefish in subarctic lakes

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The effect of core compression on the measurement of zinc concentrations and anthropogenic burdens in lake sediments

The whitefish (Coregonus lavaretus) in Lake Tyrifjorden, southeastern Norway, is monomorphic with regard to gill raker number: (x=32), and exhibits a bimodal length distribution. The small whitefish (<27 cm) were distributed mainly in the littoral zone and the deeper layers of the pelagic zone. The larger whitefish (>27 cm) were mainly distributed in the upper layers of the pelagic zone. Both small and large whitefish fed on zooplankton in the pelagic zone. In the littoral zone the diet of the two size groups was significantly different; the small whitefish fed mainly on zooplankton whereas the large whitefish fed mainly on zoobenthos.     

Zooplankton prey selection by juvenile fish in Nova Scotian Shelf basins

The stomach contents of fish <100 mm standard length (SL)and fish >100 mm SL collected during June and October intwo deep basins on the Scotian Shelf, showed that juvenile co(and silver hake fed primarily on the two most abundant speciesof zooplankton in the basins: Calanus finmarchicus and Meganyctiphanesnorvegica. These zooplankton were abundant in the basin: duringthe 2 months, forming high concentrations at depths below 200m during the day. In June fish (<100 mm SL) concentratedin the upper 50 m during both day and night. Cod and wolffislcollected between 195 and 240 m had a large percentage of C.finmarchicusin their stomachs suggesting that these fish exploited the highconcentrations of C.finmarchicus below 200 m depth There wasno evidence that Calanus hyperboreus was eaten by any juvenilefish species. Calanus finmarchicus was the most common preyof cod feeding in the top 50 m, but the percentages o C.finmarchicusstages IV and V were lower in the stomachs than in the watercolumn. Cod showed ; strong preference for M.norvegica, withthis prey species having a modified Ivlev's index of 1 indicatingthat cod were highly selective for this species. There was noevidence that C.hyperboreus. stage IV was eaten by any of thespecies offish. Silver hake and dogfish (>100 mm SL) bothfeed oi M.norvegica and C.finmarchicus in the deep regions ofthe basins. In October, silver hake was the most common juvenilefish in the basins, feeding primarily on small stages of M.norvegica.The populations of fish (>100 mm SL) in the basins were mainlyspecies known to feed heavily or euphausiids. The two basinsare unique regions of the shelf because of the large populationsof Calanus copepods and M.norvegica they contain all the yeararound, thereby providing an atrractive feeding ground for manyspecies of fish, particularly silver hake.     

Selective predation on zooplankton by pelagic Arctic charr, Salvelinus alpinus, in six subarctic lakes

Selective predation by planktivore fish appears to be an important regulatory factor of zooplankton communities, potentially causing large changes in species composition and size distributions within populations. In this study, prey preferences and size-selective predation on zooplankton by Arctic charr were examined in six subarctic lakes with Arctic charr as the dominant pelagic fish species. Most of the lakes had a zooplankton community dominated by copepods (Cyclops scutifer and Eudiaptomus graciloides), but the pelagic charr evidently selected cladoceran species (Bythotrephes longimanus, Daphnia sp. and Bosmina sp.), likely because the copepods have a higher mobility and evasiveness than the cladocerans. Furthermore, a strong size selection was also revealed for both Bosmina sp. and Daphnia sp., as individual prey from Arctic charr stomachs were exclusively larger than individuals sampled in the environment. Additionally, visibility due to size, morphology and pigmentation (egg-carrying females) was also a major factor for the selection of zooplankton prey. In conclusion, Arctic charr was found to be highly selective on zooplankton both in respect to species composition and individual size of Bosmina sp. and Daphnia sp.