Food of 0-group ocean pout [Macrozoarces americanus (Schneider)] in eastern Newfoundland: the importance of harpacticoid copepods

Stomachs of 0-group ocean pout from Newfoundland all contained harpacticoid copepods, which comprised 82% of the diet. 0-group ocean pout were found hiding among coarsely branched algae.     

Life history differences and latitudinal variation in recruitment in two species of arctic-boreal perciform fishes,the fish doctor <Emphasis Type="Italic">Gymnelus viridis</Emphasis> and the Arctic shanny <Emphasis Type="Italic">Stichaeus punctatus</Emphasis>

Growth and recruitment were examined in two arctic-boreal, shallow water marine perciform species: the fish doctor Gymnelus viridis (Zoarcidae) and the Arctic shanny Stichaeus punctatus (Stichaeidae). G. viridis ranges in the Canadian Arctic from northern Hudson Bay to northern Ellesmere Island, whereas S. punctatus has a more southerly range from Nova Scotia coastal waters to northern Hudson Bay. At Nuvuk Islands in northeastern Hudson Bay, where the two species are sympatric in shallow water, they had comparable juvenile growth rates but temporal variation in 0+ recruitment was substantially greater in S. punctatus. This difference may stem from their contrasting early life histories. S. punctatus spawns large numbers of small demersal eggs that hatch into pelagic larvae, whereas G. viridis spawns small numbers of large demersal eggs that hatch directly into demersal juveniles with no pelagic stage, suggesting that recruitment in G. viridis should be less sensitive to yearly variation in the onset of the ice-free period in Hudson Bay and the subsequent pulse of pelagic invertebrate production. Relative to Nuvuk, recruitment variation in G. viridis was found to be greater at Resolute, Cornwallis Island, close to its northern range limit, whereas recruitment variation in S. punctatus was not evident in Newfoundland, closer to its southern range limit.     

Ecological aspects of ontogenetic shifts in prey size utilization in the bay goby (Pisces: Gobiidae)

Summary Two age classes (0+ to 3+ and 4+ to 7+) of bay gobies (Lepidogobius lepidus Girard) differed in the sizes of whole prey (except polychaetes) recovered from the digestive tract. Although older fish consumed greater amounts of larger prey they did not capture larger individuals of a given prey type for seven of nine prey classes. The remaining two, harpacticoid copepods and ostracods contribute minimally to ontogenetic differences. The switch to larger prey appears to either decrease exposure to predation through a reduction in foraging time or increase energy intake. This is facilitated through older fishes greater size and/or superiority in intraspecific competition. These differences can probably be viewed as adaptations to selective pressures imposed by fluctuating food resources on predation levels in bays and estuaries.     

Diet of marine fish larvae and juveniles that use rocky intertidal pools at the Portuguese coast

The use of tidal pools during early ontogeny is likely to enhance growth, condition and survival chances of the transient marine fish larvae and juveniles that use them. However, the diet of such individuals within tidal pools is poorly known; this knowledge is important to understand why such high numbers of individuals use these environments in spring and summer on the Portuguese coast. Transient marine fishes were sampled monthly over a two‐year period in four tidal pools of a rocky reef on the west Portuguese coast. The diet composition in the tidal pools of the most abundant marine fish larvae/juveniles, Diplodus sargus and Atherina spp., were investigated. Stomach and gut contents of 483 individuals (354 D. sargus ranging in total length from 9 to 87 mm and 134 Atherina spp. ranging in total length from 10 to 31 mm) were analyzed and dietary indices estimated for the different developmental stages. The diet of D. sargus composed mainly harpacticoid copepods, chironomid larvae, ostracods, chironomid adults and amphipods; whereas the diet of Atherina spp. was mainly harpacticoid copepods, ostracods and gastropods. Along the ontogenetic development, the proportions and diversity of food items in the D. sargus diet varied. Importance of the harpacticoid copepods decreased with increasing fish size, while that of amphipods and isopods increased. Larvae and juvenile D. sargus and Atherina spp. found in rocky reef tidal pools are opportunistic consumers of a wide range of prey and thus take advantage of the high prey availability in these habitats.     

A harpacticoid copepod Microsetella spp. from sub-Arctic coastal waters and its sensitivity towards the polyaromatic hydrocarbon pyrene

Species of the harpacticoid copepod genus Microsetella are commonly reported to occur in Arctic and sub-Arctic coastal waters, but nothing has yet been reported on their sensitivity towards toxic substances. Effects of the polyaromatic hydrocarbon pyrene on Microsetella spp. from Western Greenland were investigated by looking at survival of females, feeding status, and nucleic acid content after a 96-h exposure. Less than 10% survived at 100 nM exposure concentration, and a 50% reduced survival was also evident at 0.1 and 10 nM. The RNA:DNA ratio was significantly higher, 2.1 ng/individual, at the lowest concentration of 0.01 nM compared to 0.9 ng/individual in the control, which indicates increased metabolic activities. A reduced DNA content in higher exposure concentrations of 1–100 nM suggests inhibition of egg development. Gut content was significantly declining with increasing exposure concentrations with only 2% of exposed females displaying full guts at 100 nM pyrene. Together with reduced feeding activity, effects at the population level can be anticipated, but would have to be confirmed in future experiments. The data suggests a higher sensitivity of Microsetella spp. compared to other Arctic copepods, which implies more severe effects from oil on the pelagic food web in the areas and periods where Microsetella spp. dominates Arctic plankton food webs.     

The fourbeard rockling, Enchelyopus cimbrius (L.), in eastern Newfoundland

A population of fourbeard rocklings, Enchelyopus cimbrius, was studied in shallow water (<10 m) in eastern Newfoundland. They are nocturnal, occupying burrows in muddy substrate during the day. and often retreating into these burrows at night when disturbed. The population consisted mostly of 80–150 mm individuals, aged 2-3 years. Feeding was observed only at night, and food consisted mostly of polychaete worms, especially the scale worm Harmorhoe imbricara . The contribution of polychaetes increased with increasing size of fish.     

Physiological and ecological effects of increasing temperature on fish production in lakes of Arctic Alaska

Lake ecosystems in the Arctic are changing rapidly due to climate warming. Lakes are sensitive integrators of climate‐induced changes and prominent features across the Arctic landscape, especially in lowland permafrost regions such as the Arctic Coastal Plain of Alaska. Despite many studies on the implications of climate warming, how fish populations will respond to lake changes is uncertain for Arctic ecosystems. Least Cisco (Coregonus sardinella) is a bellwether for Arctic lakes as an important consumer and prey resource. To explore the consequences of climate warming, we used a bioenergetics model to simulate changes in Least Cisco production under future climate scenarios for lakes on the Arctic Coastal Plain. First, we used current temperatures to fit Least Cisco consumption to observed annual growth. We then estimated growth, holding food availability, and then feeding rate constant, for future projections of temperature. Projected warmer water temperatures resulted in reduced Least Cisco production, especially for larger size classes, when food availability was held constant. While holding feeding rate constant, production of Least Cisco increased under all future scenarios with progressively more growth in warmer temperatures. Higher variability occurred with longer projections of time mirroring the expanding uncertainty in climate predictions further into the future. In addition to direct temperature effects on Least Cisco growth, we also considered changes in lake ice phenology and prey resources for Least Cisco. A shorter period of ice cover resulted in increased production, similar to warming temperatures. Altering prey quality had a larger effect on fish production in summer than winter and increased relative growth of younger rather than older age classes of Least Cisco. Overall, we predicted increased production of Least Cisco due to climate warming in lakes of Arctic Alaska. Understanding the implications of increased production of Least Cisco to the entire food web will be necessary to predict ecosystem responses in lakes of the Arctic.     

Intraspecific competition and density dependence of food consumption and growth in Arctic charr

1. Intraspecific competition for restricted food resources is considered to play a fundamental part in density dependence of somatic growth and other population characteristics, but studies simultaneously addressing the interrelationships between population density, food acquisition and somatic growth have been missing. 2. We explored the food consumption and individual growth rates of Arctic charr Salvelinus alpinus in a long-term survey following a large-scale density manipulation experiment in a subarctic lake. 3. Prior to the initiation of the experiment, the population density was high and the somatic growth rates low, revealing a severely overcrowded and stunted fish population. 4. During the 6-year period of stock depletion the population density of Arctic charr was reduced with about 75%, resulting in an almost twofold increase in food consumption rates and enhanced individual growth rates of the fish. 5. Over the decade following the density manipulation experiment, the population density gradually rose to intermediate levels, accompanied by corresponding reductions in food consumption and somatic growth rates. 6. The study revealed negative relationships with population density for both food consumption and individual growth rates, reflecting a strong positive correlation between quantitative food intake and somatic growth rates. 7. Both the growth and consumption rate relationships with population density were well described by negative power curves, suggesting that large density perturbations are necessary to induce improved feeding conditions and growth rates in stunted fish populations. 8. The findings demonstrate that quantitative food consumption represents the connective link between population density and individual growth rates, apparently being highly influenced by intraspecific competition for limited resources.     

Diet composition and feeding ecology of the naked goby Gobiosoma bosc (Gobiidae) from four western Atlantic estuaries

The feeding ecology of the small‐bodied benthic naked goby Gobiosoma bosc, a western Atlantic species that occurs in estuaries and other inshore habitats from Connecticut to Texas U.S.A., was investigated in a total of four estuaries spanning South Carolina, North Carolina, Maryland and New Jersey. Gut content analysis of 391 individuals revealed that G. bosc is a benthic microcarnivore that feeds primarily on polychaetes, gammarid amphipods and harpacticoid copepods. Diet composition varied with body size, tidal creek within an estuary and geographic region. Analyses of gut fullness suggest that G. bosc is a daytime visual predator and that nest and egg guarding during the reproductive season reduce foraging activity in mature males. Additionally, G. bosc infected with adult digenean parasites of the gut foraged more intensely than uninfected individuals, a relationship that was strongest for reproductively mature males. Regionally, significant variation in dietary breadth was documented and may reflect a foraging response to a decrease in prey diversity moving from estuaries of higher salinity and lower latitude to estuaries of lower salinity and higher latitude. These results contribute to an understanding of the life history of G. bosc and the role played by this common species in estuarine food webs.     

Arctic charr in Lake Myvatn: The centennial catch record in the light of recent stock estimates

In Lake Myvatn, Arctic charr (Salvelinus alpinus) is economically the most important fish species. It is fast growing fish and its size at maturity is 35–50 cm at the age of 4 to 5 years. Catch statistics that are available for the whole 20th century show considerable long-term variation with the highest catch in the 1920s. The catches after 1970 are about 40% lower than the average from 1930–1969 and the annual catches for the last decade are the lowest. Stock size during the winter fishing season fluctuated considerably between years, with average annual fishing mortality of 83.9%. The Arctic charr population has been monitored annually since 1986, using standard series of gill nets of different mesh sizes. In 1988 one to three-year-old fish were heavily reduced in numbers during the summer months. There are indications of a similar event in 1997. In both years the charr changed its main diet from Cladocera and chironomid midges to three-spined sticklebacks Gasterosteus aculeatus and snails. At the same time its condition deteriorated. The catch in the monitoring fishery in Lake Myvatn correlates with the stock size in the beginning of the following winter fishing season deduced from Leslie's method and can give prospects for the fishery in the successive fishing season. An index of abundance of young charr also correlates with the number of chironomids and cladocerans and also with ducklings that feed on the same food as the charr. Large fluctuations in the Arctic charr population in Lake Myvatn seem to be related to changes in the main food species. The catch records available from Lake Myvatn can to a large extent be used as a measure of changes of the Arctic charr population in the lake for the past century.     

Carnivorous feeding and respiration of the Arctic under-ice amphipod Gammarus wilkitzkii

. The dominant Arctic under-ice amphipod Gammarus wilkitzkii consumes a wide range of food items. The carnivorous feeding activity and energy budget of this large species were studied using three different approaches. Maximum potential ingestion rates I_max estimated from an allometric function taken from the literature and based on body mass were 2.1ǂ.4% of body carbon day^-1. Based on respiration measurements, the specific ingestion rates required to meet metabolic demands were lower (1.4ǂ.4% of body carbon day^-1). Feeding experiments, in which co-occurring pelagic calanoid (Calanus hyperboreus) or sympagic harpacticoid (Halectinosoma sp.) copepods were offered as prey, yielded actual ingestion rates of 8.0LJ.6% of body carbon day^-1 and 0.1ǂ.1% of body carbon day^-1, respectively. These results indicate that predatory feeding on pelagic copepods may constitute an important food source for G. wilkitzkii. Abundances of G. wilkitzkii at the ice underside (median: 1.6 ind. m^-2), Calanus spp. in the upper metre below the ice (2.6 ind. m^-3), and Halectinosoma sp. in the lowermost 2-3 cm of the ice (393.5 ind. m^-2) were determined from several multi-year pack-ice floes in the northern Greenland Sea and Fram Strait. Potential predation impact of G. wilkitzkii was estimated by combining information on ingestion rates with population densities. It was very high on Calanus spp. in the under-ice water layer (61.5% of the under-ice standing stock day^-1), but comparatively low on Halectinosoma sp. in the bottom of the ice (3.8% of standing stock day^-1). The observation of G. wilkitzkii preying on pelagic copepods in the under-ice water layer represents a hitherto unknown but obviously significant process and a new direction in the cryo-pelagic coupling in the Arctic marine ecosystem.     

Is diatom size selection by harpacticoid copepods related to grazer body size?

Copepods are known as important consumers of primary production and are eaten by larger animals. They therefore form a main link to higher trophic levels. While feeding pathways and specificity of planktonic copepods have been well studied, the selectivity of the benthic harpacticoid copepods is far less documented. A better knowledge of the functional ecology of harpacticoids as important grazers on primary producers may have consequences for the re-evaluation of basic energy flow in benthic ecosystems.We tested whether size selectivity for diatoms exists in harpacticoid copepods. We hypothesized that size selectivity of harpacticoid copepod species is strongly related to body size. Because of morphological constraints, we expected smaller copepods to prefer smaller diatoms while larger copepods should be able to consume both small and large diatoms. We tested this hypothesis in four harpacticoid copepod species of varied body size: Tigriopus brevicornis, Harpacticus obscurus, Amphiascus minutus and Paramphiascella fulvofasciata. As food source we used two ¹³C labelled strains of the benthic diatom Seminavis robusta with a four-fold difference in cell biovolume.Three out of four harpacticoid species showed size selectivity: H. obscurus and A. minutus preferred the larger Seminavis cells, while P. fulvofasciata selected the smaller Seminavis cells. Based on monoclonal treatments, there was no clear preference found for T. brevicornis although there was a small preference for large cells in the mixed treatments. Except for P. fulvofasciata, all species showed a lower uptake when offered the mixed diet (both small and large cells). Although most species showed a size selectivity, our results suggest that this selectivity was not related to their body size. However, the only species that ate significantly more of small diatoms was characterised by comparatively small mouthparts in relation to its body size.     

Climate warming decreases the survival of the little auk (Alle alle), a high Arctic avian predator

Delayed maturity, low fecundity, and high adult survival are traits typical for species with a long‐life expectancy. For such species, even a small change in adult survival can strongly affect the population dynamics and viability. We examined the effects of both regional and local climatic variability on adult survival of the little auk, a long‐lived and numerous Arctic seabird species. We conducted a mark‐resighting study for a period of 8 years (2006‐2013) simultaneously at three little auk breeding sites that are influenced by the West Spitsbergen Current, which is the main carrier of warm, Atlantic water into the Arctic. We found that the survival of adult little auks was negatively correlated with both the North Atlantic Oscillation (NAO) index and local summer sea surface temperature (SST), with a time lag of 2 and 1 year, respectively. The effects of NAO and SST were likely mediated through a change in food quality and/or availability: (1) reproduction, growth, and development of Arctic Calanus copepods, the main prey of little auks, are negatively influenced by a reduction in sea ice, reduced ice algal production, and an earlier but shorter lasting spring bloom, all of which result from an increased NAO; (2) a high sea surface temperature shortens the reproductive period of Arctic Calanus, decreasing the number of eggs produced. A synchronous variation in survival rates at the different colonies indicates that climatic forcing was similar throughout the study area. Our findings suggest that a predicted warmer climate in the Arctic will negatively affect the population dynamics of the little auk, a high Arctic avian predator.     

Diet and body shape changes of pāroko Kelloggella disalvoi (Gobiidae) from intertidal pools of Easter Island

This study assesses seasonal variation in the morphology and diet of juveniles and adults of the Easter Island endemic goby Kelloggella disalvoi from intertidal pools during September–October 2015 (spring) and June–July 2016 (winter), utilizing geometric morphometric and gut‐content analyses. A set of 16 landmarks was digitized in 128 individuals. Shape changes related to size changes (i.e. allometry) were low (18·6%) and were seasonally similar. Body shape changes were mainly dorsoventral (44·2% of variance) and comprised posteroventral displacement of the premaxilla and bending of the body. The latter included vertical displacement of the anterior portion of the first and second dorsal fins and the entire base of the caudal fin. Diets mainly comprised developmental stages of harpacticoid copepods (from eggs to adults), ostracods, isopods, gastropods and bivalves. Also, trophic niche breadth remained constant throughout development and did not vary between seasons. Nonetheless, significant dietary differences were detected in specimens collected during spring (main prey items: harpacticoid copepods and copepod eggs) and winter (harpacticoid copepods and copepod nauplii). Finally, there was weak but significant covariation between diet and morphology: molluscivores were characterized by having an inferior mouth gape, whereas planktivores had an anteriorly directed premaxilla.     

Selection on Arctic charr generated by competition from brown trout

We experimentally explored population‐ and individual‐level effects on Arctic charr (Salvelinus alpinus) resulting from resource competition with its common European competitor, the brown trout (Salmo trutta). At the population level, we compared performance of the two species in their natural sympatric state with that of Arctic charr in allopatry. At the individual level, we established selection gradients for morphological traits of Arctic charr in allopatric and in sympatric conditions. We found evidence for interspecific competition likely by interference at the population level when comparing differences in average performance between treatments. The growth and feeding rates did not differ significantly between allopatric and sympatric Arctic charr despite lower charr densities (substitutive design) in sympatric enclosures indicating that inter‐ and intraspecific competition are of similar strength. The two species showed distinct niche segregation in sympatry, and brown trout grew faster than Arctic charr. Arctic charr did not expand their niche in allopatry, indicating that the two species compete to a limited degree for the same resources and that interference may suppress the growth of charr in sympatric enclosures. At the individual level, however, we found directional selection in sympatric enclosures against individual Arctic charr with large head and long fins and against individuals feeding on zoobenthos rather than zooplankton indicating competition for common resources (possibly exploitative) between trout and these charr individuals. In allopatric enclosures these relations were not significant. Diets were correlated to the morphology supporting selection against the benthic‐feeding type, i.e. individuals with morphology and feeding behaviour most similar to their competitor, the benthic feeding brown trout. Thus, this study lends support to the hypothesis that Arctic charr have evolved in competition with brown trout, and through ecological character displacement adapted to their present niche.     

The trophic importance of algal turfs for coral reef fishes: the crustacean link

On coral reefs, the epilithic algal matrix (EAM) is widely recognised as an important resource for herbivorous and detritivorous fishes. In comparison, little is known of the interaction between benthic carnivores and the EAM, despite the abundance of Crustacea within the EAM. The trophic importance of the EAM to fishes was investigated in Pioneer Bay, Orpheus Island, Great Barrier Reef. Fish densities were quantified using visual and clove oil censuses, and gut content analyses conducted on abundant fish species. Crustaceans were found to be an important dietary category, contributing between 49.5 and 100 % of the gut contents, with harpacticoid copepods being the dominant component. Of the benthic carnivores, the goby Eviota zebrina was found to consume the most harpacticoids with a mean of 249 copepods m^?2 day^?1. This represents approximately 0.1 % of the available harpacticoid population in the EAM. In a striking comparison, herbivorous parrotfishes were estimated to consume over 12,000 harpacticoids m^?2 day^?1, over 27 times more than all benthic carnivores surveyed, representing approximately 5.3 % of the available harpacticoid copepod population each day. The high consumption of harpacticoid copepods by benthic carnivores and parrotfishes indicates that harpacticoids form an important trophic link between the EAM and higher trophic levels on coral reefs.     

Intrashoot distributions of leaf dwelling harpacticoid copepods on the seagrass Zostera marina L.: implications for sampling design

The distribution of leaf dwelling harpacticoid copepods within seagrass (Zostera marina L.) shoots was investigated on four dates in 1986 and 1987. Copepods were found to be non-uniformly distributed on shoots, with higher abundances observed on older leaves. Patterns of abundance within shoots could not be explained by the surface area of individual leaves except on the sampling date with the highest copepod densities. It is suggested that harpacticoid copepod distributions on seagrass shoots are primarily determined by the pattern of epiphytic biomass. However, at high population densities, habitable surface area may be the limiting factor. Increased habitat complexity at high epiphyte loads does not seem to be the cause of the copepod distributions observed in this study. An accurate method for estimating seagrass copepod abundance per unit sediment area using intrashoot distributions is described and compared to existing methodology.     

A comparison of growth patterns between a stunted and two large predatory Arctic charr populations under identical hatchery conditions

Arctic charr are characterized by an extensive variability in growth and body size in natural waters. Although growth traits may involve a significant heritable component, most of this intraspecific variation presumably is environmentally induced and thus attributable to phenotypic plasticity. In the present study, size-at-age and length–weight relationship (body condition) were assessed for three Finnish Arctic charr populations of different geographical origins and extreme size forms (a stunted vs. two large-growing, predatory charr) held under standardized rearing conditions for 3 years (up to 37 months after hatching). In particular, our interest was to investigate whether the differences in growth between the large and the stunted charr as observed in the wild populations would diminish when the fish are offered suitable food in abundance. Population-specific mean body size and condition differed significantly in 0+, 1+, 2+ and 3+ fish. However, the identical rearing conditions resulted in the originally stunted charr reaching a comparable final mean size (317 mm/427 g) as the large charr populations (343 mm/510 g and 359 mm/497 g). Some individuals were of the same size as their parents at spawning already at the age of 0+ years. Furthermore, length–weight regression residuals of the stunted charr developed to a notably high level, indicating the largest final condition mean. The increase of size variation (CV of weight) in stunted charr lasted for over two growth seasons, whereas in large charr it remained stable since the end of the first summer. Variations in mortality and sexual maturation at age 2 seemed to be less relevant factors affecting overall growth performance. The study demonstrates an example of the high plasticity involved in the growth of fish: the stunted charr possess a tremendous capacity for growth in a benign environment, virtually corresponding to that observed in the large predatory populations.     

Cannibals and parasites: conflicting regulators of bimodality in high latitude Arctic char, Salvelinus alpinus

Unexploited populations of Arctic char (Salvelinus alpinus) sampled in autonomous lake ecosystems in northern Svalbard demonstrate extraordinary catch curves with age and size frequency distributions characterized by discrete bimodality. Analyses of size‐age relationship, summer diet and food‐related intestinal parasite intensities of modal char groups revealed a pattern of discrete ontogenetic niche shifts. Life‐history changes at age 10–15 and size 200–300 mm/50–300 g involved shifting from an initial mode of small‐sized, slow‐growing and sexually mature individuals feeding on micro‐crustaceans and aquatic insects (Chironomidae, Trichoptera), to a terminal mode of large‐sized and fast‐growing cannibals. Cannibalism, however, was found to result in accumulation of cestodan parasites, of which Diphyllobothrium ditremum increases age‐related mortality rates and may be lethal at 1500–2000 plerocercoids. Genetically allopatric populations with cannibalism demonstrated a female‐biased sex ratio, primarily in the initial mode, suggesting sexual asynchrony in their ontogeny. By contrast, a small population of large‐sized, non‐cannibalistic Arctic char feeding exclusively on the large amphipod Gammaracanthus lacustris, demonstrated unimodal size and age frequency distribution, faster growth, an excess of males and lower parasite burden. Seasonal prey shortage and slow juvenile growth in association with fitness components favoring large body size is a suggested mechanism for inducing cannibalism. Although not the basic cause of bimodality as such, it is concluded that ontogenetic niche shift by cannibalism reinforces discrete age modal divergence resulting in the numerical preponderance of large‐sized individuals in these marginal char populations. Cannibalism is thus considered an important strategy for survival of landlocked Arctic char in the High Arctic. As a conflicting cost to the more efficient use of available energy by larger individuals, the accumulation of cestodan parasites in cannibals, however, will reduce the survival rate of old individuals and accelerate their termination within this modal group.     

Lipid‐rich zooplankton subsidise the winter diet of benthivorous Arctic charr (Salvelinus alpinus) in a subarctic lake

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