Arctic charr in sympatry with burbot: ecological and evolutionary consequences

The trophic niche and parasite infection of Arctic charr (Salvelinus alpinus) were explored in two lakes with sympatric burbot (Lota lota) and two lakes without burbot in subarctic Norway. The CPUE of burbot and charr were similar in one lake, but burbot had a low population density in the other. Burbot were benthivorous in both lakes. Other co-occurring species like brown trout (Salmo trutta), Atlantic salmon parr (Salmo salar), grayling (Thymallus thymallus) and minnow (Phoxinus phoxinus) were also benthivores. At high densities, benthivorous burbot forced the whole Arctic charr population to utilise mainly the limnetic trophic niche. In contrast, at low burbot density or without burbot present, Arctic charr were primarily benthivorous in the littoral zone. Thus, a clear interactive segregation in diet was observed between Arctic charr and burbot at high burbot densities. There was also a high predation pressure from burbot on young Arctic charr along the benthic zones. The extensive use of zooplankton as prey caused a high parasite infection pressure of copepod transmitted Diphyllobothrium spp. larvae, with the potential for high negative impact on the Arctic charr population. As the benthivore trophic niche was occupied by burbot, the ecological opportunities for polymorphism with benthivorous ecotypes or morphs of Arctic charr were probably prevented. Therefore, the sympatry with burbot seems to have large ecological and evolutionary consequences for this Arctic charr population compared with neighbouring lakes where burbot is absent.     

Lake Whitefish and Lake Herring Population Structure and Niche in Ten South-Central Ontario Lakes

Synopsis This study compares simple fish communities of ten oligotrophic lakes in south-central Ontario. Species densities and population size structure vary significantly among these lake communities depending on fish species present beyond the littoral zone. Lake whitefish are fewer and larger in the presence of lake herring than in their absence. Diet analysis indicates that lake whitefish shift from feeding on both plankton and benthic prey when lake herring are absent to a primarily benthic feeding niche in the presence of lake herring. When benthic round whitefish are present, lake whitefish size and density decline and they move lower in the lake compared to round whitefish. Burbot are also fewer and larger in lakes with lake herring than in lakes without herring. Burbot, in turn, appear to influence the population structure of benthic coregonine species. Lower densities of benthic lake whitefish and round whitefish are found in lakes containing large benthic burbot than in lakes with either small burbot or where burbot are absent. Predation on the pelagic larvae of burbot and lake whitefish by planktivorous lake herring alters the size and age structure of these populations. As life history theory predicts, those species with poor larval survival appear to adopt a bet-hedging life history strategy of long-lived individuals as a reproductive reserve.     

Diel vertical migration of larval and early‐juvenile burbot optimises survival and growth in a deep,pre‐alpine lake

1. Burbot larvae (Lota lota) perform a substantial diel vertical migration (DVM) of increasing amplitude in the pelagic zone during a 3‐month period before migrating to the littoral zone as early‐juveniles. We hypothesised that feeding in the warm surface layers at night and then spending the day in cold water below the thermocline reduces metabolic costs and earns burbot larvae an energetic advantage. 2. To test our hypothesis, we mimicked the temperature conditions experienced by vertically migrating burbot in the pelagic zone. We also simulated three further scenarios, in which temperature remained constant. 3. Burbot showed the best performance (defined as specific growth rate multiplied by the probability of survival) in the treatments simulating DVM. The high temperature treatment, simulating permanent residence in the warm epilimnion, resulted in high growth combined with high mortality. At a permanently low temperature, simulating life in the hypolimnion, growth was poor and activity reduced. 4. In a deep, temperature‐stratified lake, where the apparently beneficial overall medium temperature is found in a restricted layer within the thermocline, DVM optimises performance in young burbot. Various ultimate factors might act synergistically in selecting for DVM in larval and early‐juvenile burbot.     

Looking for love under the ice: Using passive acoustics to detect burbot (Lota lota: Gadidae) spawning activity

1. Burbot (Lota lota: Gadidae) is a difficult species to manage effectively due to its preference for deep-water habitats and under-ice spawning behaviour, resulting in a poor understanding of its reproductive activity. However, the use of acoustic signalling by burbot as part of their mating system has recently been described and this behaviour may provide a means of investigating questions regarding the spatial and temporal distribution of spawning aggregations using passive acoustic monitoring.
2. We used audio and video recording to confirm that burbot vocalise and that these vocalisations can be detected under field conditions as well as to characterise the relationship between burbot acoustic signalling and spawning behaviour. We also evaluated the feasibility of locating and monitoring burbot spawning aggregations in real time using passive acoustics.
3. Burbot vocalisations were difficult to identify with only about 6% of the recordings containing calls being successfully identified as such in the field. Burbot vocalised more often between sundown and sunrise than during daylight hours. Calls recorded at night tended to be lower frequency, longer duration, and have lower bandwidth than those made during the day.
4. Burbot vocalisations could not be recorded in conjunction with video recordings of spawning activity, indicating that burbot may not call during active spawning, but may use acoustic communication to signal the onset of reproductive readiness and to form pre-spawning aggregations.
5. While burbot calls were readily identifiable, observers had a difficult time identifying burbot calls in real time under field conditions. Passive acoustic monitoring demonstrates considerable potential as a management tool to locate burbot spawning grounds and identify periods of activity, but may not be an appropriate technique for monitoring spawning activity in real time.

     

The impact of predation by burbot (<Emphasis Type="Italic">Lota lota</Emphasis> L.) on the macroinvertebrate community in the littoral zone of a large lake

We studied the effects of predation by juvenile burbot (Lota lota) on the macroinvertebrate community in mesocosm experiments in the stony littoral zone of Lake Constance, a large prealpine lake in Central Europe. Although the growth data of the burbot suggest that the benthivorous fish exerts strong predation pressure on the invertebrate community, the predicted level of consumption is only poorly reflected by changes in biomass and abundance of most invertebrate prey taxa. High exchange rates of the prey between the mesocosms and the ambient littoral environment apparently masked the true predation effects of fish. Also, life-history events such as hatching or synchronised emergence of larvae led to temporal effects that obscured the impact of predation. However, for the dominant prey organism, the freshwater amphipod Gammarus roeseli, direct lethal effects appeared when its migration was limited. When exchange with the ambient littoral zone was possible, abundance and biomass of G. roeseli were unexpectedly high in the mesocosms stocked with burbot, indicating behavioural responses to fish presence. During the experiment, the burbot gained ca. 15% in length and about 60% in body mass. According to stomach content analyses at the end of the experiment, the ingested prey consisted mainly of relatively large and abundant invertebrates. Our study indicates that predation by juvenile burbot should be an important factor in structuring the benthic invertebrate community in terms of qualitative and quantitative effects.     

Growth and habitat separation in eight cohorts of three species of cyprinids in a subalpine lake

Synopsis Distribution and growth of the embryos, larvae and juveniles of Rutilus rutilus (roach), Scardinius erythrophthalmus (rudd) and Leuciscus cephalus (chub) from an oligotrophic subalpine lake in Tyrol, Austria, were studied during the first three to four months after hatching. R. rutilus was the first to spawn, a single cohort hatching around May 23rd. Four cohorts of S. erythrophthalmus hatched between June 19 and August 1. Three cohorts of L. cephalus hatched between July 3 and 25. The length/weight relationship of all species changed at a length of approximately 15–16 mm. R. rutilus, hatching at the lowest temperature, also showed the lowest growth rate during early life (maximum 10.4 per cent fresh body weight day^–1). In the other two species relative growth rates up to 20% day^–1) were measured. Rudd and chub remained in the shallow littoral during the whole period of observation, whereas roach left the littoral a few weeks after hatching and migrated into deeper water. A subtle shift in vertical distribution was observed for the first cohort of rudd which moved into slightly deeper water when the second cohort made its appearance.To whom correspondence should be addressed     

The predation on Asellus aquaticus (L.) by perch,Perca fluviatilis (L.), in a small forest lake

Asellus aquaticus (L.) was the most important benthic food item for perch, Perca fluviatilis (L.), in a small, extremely humic forest lake in southern Finland. The proportion of A. aquaticus in the diet of perch varied according to the former's availability, which, in turn, depended on its life cycle. Perch 11.0–12.9 cm (total length) most frequently fed on A. aquaticus; smaller and larger perch fed more frequently on crustacean zooplankton and aquatic insect larvae, respectively. The high density of the perch population, the importance of A. aquaticus in the benthos of the study lake and the general high activity level of the prey resulted in a high predation (0.1–1.8% per day). The population of A. aquaticus was also limited by the scarcity of macrophytic vegetation and the small area of oxygenated littoral in the lake.     

Life history,population dynamics and production of Pontoporeia hoyi (Crustacea,Amphipoda) in relation to the trophic gradient of Lake George,New York

The life history characteristics, population dynamics and production of Pontoporeia hoyi in Lake George, New York, were studied from May 1981 through October 1982. P. hoyi, in terms of both density and standing crop, is the most prevalent member of the deep water macrobenthos of Lake George. It reproduces in the winter, with young being released in the late winter-early spring. At the southernmost study site, young released in the spring grew to 6–7 mm in length and bred during their first winter. At the remaining sites, P. hoyi required two years to complete its life cycle. This difference in life history characteristics can be related to food availability and temperature differences. The open waters of the south end of Lake George are not only more productive but are also more closely associated with the littoral zone, providing a wealth of bacteria-rich detritus for benthic deposit feeders. The greater food availability in the south basin of Lake George is reflected in significantly larger brood sizes and smaller size at maturity for P. hoyi populations from the south end of the lake.The southernmost study site has significantly greater P. hoyi density and standing crop than all other sites. The cohort of the year dominated density and standing crop at the southern site while the cohort of the previous year dominated standing crop at the other sites. Peak abundance ranged from 600 · m^–2 at the north site to 2 900 · m^–2 at the south site. Cohort production ranged from 2g · m^–2 at the north site to 15g · m^–2 at the south site.     

The benthic macroinvertebrate community in a coastal sand dune lake relative to habitat and changing lake levels

The benthic macroinvertebrate community (BMI) in a freshwater coastal dune lake without a surface outlet was investigated in May and October, 1986. Fifty-three invertebrate taxa were identified from Carter Lake, including three euryhaline crustacean species (Corophium spinicorne, Gnorimosphaeroma oregonensis lutea, and Acanthomysis awatchensis). Corophium spinicorne dominated the BMI communities of the littoral zones and sphaeriid clams dominated the deepwater community.The lake level dropped about 2.5 m between April and October. Based upon this decline, the lake bottom was divided into four major habitats: a sandy temporarily submerged littoral zone (A); a sandy submerged littoral zone (B); and mid-depth zone of mixed mud and sand and the macrophyte, Nitella (C); and a deep zone (D) with soft mud. The average density of BMI was highest in the littoral zones (A and B) in May and in zone B in October (zone A was dry). The lowest density occurred in zone D. In May, BMI biomass was highest in the littoral zones, but the biomass was highest in the mid-depth zone in October. The mid-depth zone in October. The mid-depth zone had the most diverse community.The two most abundant species in the temporarily submerged area, Corophium spinicorne and Juga plicifera, were found in greater numbers deeper in the lake after the water level dropped, suggesting migration by these species in response to changing water levels.     

Test of Competitive Interactions for Space Between Two Benthic Fish Species, Burbot Lota lota, and Stone Loach Barbatula barbatula

The substrate preferences, growth rates and foraging efficiency of two small benthic fish species, juvenile burbot, Lota lota, and stone loach, Barbatula barbatula, were compared in combined outdoor mesocosm and indoor laboratory experiments. Both species preferred the same stony substrate when alone, but significant differences in habitat selection were found between the two species under food deprivation and competition conditions. In burbot, preference for the stony habitat was reinforced under food-deprivation conditions and became even stronger when a potential competitor, the stone loach, was present. In contrast, stone loach switched to the gravel substrate when either starving or in the presence of a heterospecific competitor. Growth rates and foraging efficiency of burbot were significantly highest in the stony substrate and decreased with finer substrates. In stone loach, neither growth rates nor foraging efficiency were significantly different among the different substrates. The results provide an example of habitat partitioning by means of different competition styles, with a stenoecious, dominant style of the burbot and an euryoecious, evasive style of stone loach allowing coexistence of two sympatric fish species by graded interactions at an individual level in the littoral zone of a large lake.     

First Year Ontogenetic Diet Patterns in Two Coexisting <Emphasis Type="Italic">Sander</Emphasis> Species, <Emphasis Type="Italic">S. lucioperca</Emphasis> and <Emphasis Type="Italic">S. volgensis</Emphasis> in Lake Balaton

First year ontogenetic diet patterns and size-dependent diet partitioning of two coexisting piscivorous percids, pikeperch and Volga pikeperch were analysed in Lake Balaton. Diet patterns of the two pikeperches followed a common trend, catching larger preys at larger sizes, but in details they were different. Until 20–30 mm size age-0 pikeperch fed on planktonic microcrustaceans and were found only in the pelagic zone. After this size, from the end of May, a part of the stock shifted habitat and moved to the littoral zone to feed on Limnomysis benedeni. Majority of the stock stayed in the pelagic zone and shifted to Leptodora kindtii. In the pelagic group early piscivory, mainly cannibalism, occurred only sporadically. However, the few individuals that reached 100 mm size during the first year were typically piscivorous. Volga pikeperch stayed in the pelagic zone during the first year and shifted from planktonic microcrustaceans to benthic macroinvertebrates. General patterns of the first year diet ontogeny were similar in the different areas of Lake Balaton in both pikeperches with only some differences in the secondary food resources, especially in the share of chironomids. Chironomids were eaten at highest ratio by the two fish species in the Keszthely basin in accordance with their abundance pattern. Maximum prey sizes increased significantly with predator size in both pikeperch and Volga pikeperch. However, pikeperch ate much larger prey than did Volga pikeperch at the same size although they have a similar mouth gape to length ratio. These two, closely related pikeperches, although occur in the same habitat and have similar patterns of diet ontogeny, effectively partition food resources in Lake Balaton during the first growing season, except in the very beginning of their ontogeny at L = 5–10 mm size. It appears that age-0 Volga pikeperch is not a significant competitor for the economically highly rated pikeperch in Lake Balaton. On the other hand, it is age-0 pikeperch that might affect Volga pikeperch during its early ontogeny, in the planktivorous stage.     

Comparison of the life cycles of two sympatric estuarine polychaetes, Hediste diadroma and H. atoka (Polychaeta: Nereididae), in the Nanakita River estuary, northeastern Japan

The life cycles of two species of estuarine polychaetes, Hediste diadroma and H. atoka, were investigated using field surveys and colonization experiments in the Nanakita River estuary, Japan. Lactate dehydrogenase (LDH) allozyme banding patterns in starch gel electrophoresis were used to distinguish between the species. We found differences in the life cycle, reproductive ecology, and distribution of the two polychaetes. H. diadroma exhibited conspicuous reproductive swarming behavior during the spring tide in April to early May. After hatching, the trochophore larvae may be transported out to sea and return to the river estuary at the =3-setiger stage. By late July, juveniles settled into brackish water, where they completed the 1-year life span. In contrast, H. atoka never exhibited this conspicuous reproductive swarming behavior. The reproductive period of this species spanned most of the year, except winter. H. atoka juveniles hatched mainly as 3-setiger nectochaetes, which immediately became benthic. The juveniles actively dispersed during the 4-to 7-setiger stage, although they could also do so at any later stage. The life span of H. atoka was about 6 months. In small river estuaries, river flooding may flush sediment and benthic animals out to sea and create vacant habitats for colonization by polychaetes. In the Nanakita River estuary, H. atoka maintained a large and stable population in Gamo Lagoon, located 250m upstream from the river mouth, whereas H. diadroma was rarely found there. Swimming excursions of juvenile H. atoka at any stage in the life cycle may ensure its distribution to new vacant habitats, and its life-cycle characteristics may allow large populations to build rapidly. As a result, H. atoka was able to maintain dominance in both the river and the lagoon in the Nanakita River estuary.     

Trophic ecology of two congeneric pipefishes (Syngnathidae) of the lower York River,Virginia

Synopsis The northern pipefish, Syngnathus fuscus, and the dusky pipefish, S. floridae, co-exist in eelgrass beds in the lower York River. To compare the trophic ecology of both pipefishes, gut contents of 3488 northern pipefish, and 1422 dusky pipefish were examined and quantified by dry weight. Samples were taken with a dip net from May through November 1992. Syngnathus fuscus fed mainly on amphipods (Gammarus mucronatus, Amphithoe longimana, and Caprella penantis), and to a lesser degree on copepods, and isopods (Edotea spp. and Erichsonella attenuata). Syngnathus floridae fed mainly on grass shrimps (Palaemonetes pugio, P. vulgaris, and Palaemonetes eggs), and to a lesser degree on copepods, isopods, and mysids (Mysidopsis bigelowi). No seasonal trends were observed in the feeding habits of these pipefishes. Specimens of both species smaller than 100 mm TL fed mainly on copepods. Larger S. fuscus fed on amphipods, while larger S. floridae fed on grass shrimps. Schoener's diet overlap index revealed high consumption of the same prey type by both pipefish species only in July, but only for those specimens smaller than 110 mm TL. Food partitioning between both pipefish was associated with different snout sizes and shapes.     

Effects of water-level regulation on the nearshore fish community in boreal lakes

The fish community in the littoral areas of eight regulated lakes and five reference lakes in Finland was sampled by electrofishing. No significant effect of winter drawdown on species richness was recorded across lakes. Total fish density for stony bottoms of the regulated and reference lakes averaged 19.3 and 32.7 individuals per 100 m², respectively, but this difference was not statistically significant. The combined proportion of littoral fish species, including minnow (Phoxinus phoxinus), bullhead (Cottus gobio), alpine bullhead (Cottus poecilopus), nine-spined stickleback (Pungitius pungitius), and stone loach (Barbatula barbatula), supplemented with zoobenthos feeders ruffe (Gymnocephalus cernuus) and young burbot (Lota lota), was much lower in the regulated lakes than in the reference lakes. Besides winter drawdown, other variables, such as nutrient level and lake size, affected the fish community. Guest editors: K. M. Wantzen, K.-O. Rothhaupt, M. M?rtl, M. Cantonati, L. G.-Tóth & P. Fischer Ecological Effects of Water-Level Fluctuations in Lakes     

Effects of temperature and a piscivorous fish on diel winter behaviour of juvenile brown trout (Salmo trutta)

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Patches and the responses of lake benthos to sunfish nest-building

Summary Males of three sunfish species (Centrarchidae) construct nests for spawning and often share them sequentially in the littoral zone of a 4-hectare lake in New York State. To determine spatial and temporal effects of this reproductive behavior on zoobenthos in patches, I sampled bottom assemblages from inside nests and from adjacent (<1 m from nest perimeter), relatively undisturbed sites during the reproductive season of 1986 and immediately prior to nest-building in 1987. The reproductive behaviors of largemouth bass (Micropterus salmoides), redbreast sunfish (Lepomis auritus), and pumpkinseeds (L. gibbosus) altered relative abundances and significantly decreased benthic invertebrate diversity and density. These effects were extremely pronounced during the reproductive season and were partially detectable the following year. Community changes were probably the result of both bioturbation (modification of sediment size and organic content) and predation. The ecosystem-wide effects of nest-building are evaluated in terms of the number, distribution, and longevity of patches.     

Distribution and onshore migration behaviour of burbot larvae in Lake Constance, Germany

Larval abundance of burbot Lota lota in the pelagic zone of Lake Constance, Germany, peaked during April and remained at a constant but lower level until mid‐June. From the end of May onwards a significant diel vertical migration pattern was observed: while burbot larvae were distributed above as well as below the thermocline in the daytime with the highest percentages of abundance at water depths of 30 and 50 m (below the thermocline) on 5 and 25 June, they were concentrated from 2 to 15 m (above the thermocline) during the night. This migration pattern coincided with a significant decline in the food items preferred by burbot at this life stage, cyclopoid copepods. This phase of diel vertical migration lasted for c . 4 weeks until the burbot larvae suddenly disappeared from the pelagic zone. This indicated a short phase, lasting only a few days, of larval settlement into profundal benthic habitats. This was supported by a significant, abrupt decrease in the absolute growth rates of the larvae at day 56 after hatching. The day–night pattern of vertical migration observed could be assumed to be a transition phase from a pelagic to a benthic lifestyle, as reported for juveniles of several other, marine gadoid species.     

Influence of temperature and body weight on mosquito predation by the dragonfly nymph Mesogomphus lineatus

Population densities, migrations and food composition of Echinogammarus veneris in Lake Kinneret were studied during march–September 1976. Migrations are affected by changes of water level and littoral currents. Peridinium was abundant in the gut content during its blooming period in the lake while cladocerans, benthic copepods, rotifers and other algae were abundantly found in other months. Metabolic parameters that were calculated for the gammarid population indicated that its contribution to the carbon-flow system in Lake Kinneret is negligible in comparison with nematodes, molluscans or planktonic copepods and cladocerans.     

Feeding habit of larval and juvenile ayu,Plecoglossus altiyelis in the surf zone of Tosa Bay,Japan

A total of 7,000 larval and juvenilePlecoglossus altivelis was collected at semimonthly intervals with a small seine in a surf zone of Tei beach facing Tosa Bay during the period of June 1982 to May 1983. They occurred in the surf zone from middle October to middle May. About 500 larvae and juveniles (10.9–59.9 mm TL) were used to examine their feeding habit. The feeding incidences by collection dates fluctuated from 0 to 100%, with 90.6% in total incidence. They fed mainly on copepods (e.g.Paracalanus parvus andOithona spp.) throughout postlarval and juvenile stages, while they first took small benthic animals at 53.0 mm TL. Their food compositions were influenced fundamentally by the planktonic fauna of the surf zone, but larvae under 20 mm TL tended to take relatively larger copepods.     

Ontogenetic diet shift, feeding rhythm, and daily ration of juvenile yellowfin goby Acanthogobius flavimanus on a tidal mudflat in the Tama River estuary, central Japan

To ascertain the feeding habits of benthic juvenile yellowfin goby Acanthogobius flavimanus, the gut contents of 599 specimens (15–41 mm in standard length, SL), collected on a tidal mudflat in the Tama River estuary throughout the diel cycle, were examined. The major prey items changed from harpacticoid copepods to errant and sedentary polychaetes at ca. 20 mm SL. Prey width increased with fish size. Fish of 26–28 mm SL fed mainly from sunset to morning, with highest feeding intensity during twilight hours and/or high tide. Based on the gut evacuation rate estimated from a forced feeding experiment in the laboratory and data for the diel change of mean gut-content volume in the field, the daily ration of juvenile yellowfin goby (26–28 mm SL) was calculated to be 13.8 mm³ fish⁻¹ day⁻¹. This volume is approximately equivalent to 3.9 individuals of the errant polychaete Ceratonereis erythraeensis (9.7 mm in body length, BL) or 8.1 individuals of the sedentary polychaete Prionospio japonica (14.8 mm BL), both species occurring abundantly on the mudflat during the study.