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.     

Asymmetric admixture and morphological variability at a suture zone: parapatric burbot subspecies (Pisces) in the Mackenzie River basin,Canada

The burbot (Lota lota L.) is a northern freshwater fish with a circumpolar distribution. Two subspecies diverged due to isolation during glacial maxima: Lota lota lota is the Eurasian-Beringian subspecies and Lota lota maculosa the North American subspecies. We sampled burbot from the Great Slave Lake and Mackenzie River area, Canada, the only known contact zone of these two lineages. Using molecular methods (microsatellite loci and mtDNA sequence) we found that the subspecies’ distributions abut in the Mackenzie River delta, with L. l. lota in the lower delta and L. l. maculosa in all upstream rivers and lakes. Admixture between subspecies was minimal, decreased with increasing geographic distance, and was asymmetrical: mitochondrial and nuclear genetic introgression was from L. l. lota into L. l. maculosa but not the reverse. Within subspecies, there was low inter-population genetic differentiation, no isolation-by-distance, and no evidence for sex-biased dispersal. We did not identify a difference in body length between subspecies per se, though mean lengths differed among localities. Thus, genetic data demonstrate that burbot subspecies are reproductively isolated though the extent to which morphologically variability relates to local versus subspecific variation remains unclear.     

Influence of substrate preference and complexity on co-existence of two non-native gammarideans (Crustacea: Amphipoda)

Substrate choice, swimming activity and risk to predation by burbot (Lota lota) of the well established Gammarus roeselii and the invader Dikerogammarus villosus were studied in mixed and single-species aquarium experiments. We used stones, gravel and aquatic weeds (Elodea, Chara) as substrates. We hypothesized that both species have different substrate preferences and that substrate affects the predation risk. We also assumed that presence of D. villosus influences substrate preference and predation risk of G. roeselii since the invader is known to affect the behavior of other gammarids. Adults of D. villosus in single species experiments and juveniles in mixed and single species experiments were evenly distributed over the different substrates but adults in mixed species experiments were more likely to prefer stone substrate. In contrast, adults and juveniles of G. roeselii clearly preferred aquatic weeds independent of the presence/absence of the invader. Both species preferred substrates with fissured surface over substrates with smooth surface. Gammarus roeselii was observed swimming more often than D. villosus in the open water but its swimming activity was lower when its preferred substrate was present compared with its swimming activity if non-preferred substrates were present. Predation rate of burbot on D. villosus was comparatively low and independent of the substrate. Burbot consumed many more G. roeselii than D. villosus, both in mixed and single species experiments. But when the preferred substrate of G. roeselii (weeds) was used in the experiments, predation rate of burbot on G. roeselii was somewhat lower than that when non-preferred substrates were present. The results of the experiments support our hypothesis that the gammarids studied here have different substrate preferences and that presence of the preferred substrate can affect predation risk. However, there is no evidence that presence of D. villosus affected substrate choice or predation risk in G. roeselii. We consider that differences in use of spatial niches permit co-existence of G. roeselii and D. villosus in the wild when substrates are diverse. The fact that G. roeselii than D. villosus is more often observed swimming in the open water may explain its higher risk of being captured by fish.     

Evidence for predatory control of the invasive round goby

We coupled bioenergetics modeling with bottom trawl survey results to evaluate the capacity of piscivorous fish in eastern Lake Erie to exert predatory control of the invading population of round goby Neogobius melanostomus. In the offshore (>20 m deep) waters of eastern Lake Erie, burbot Lota lota is a native top predator, feeding on a suite of prey fishes. The round goby invaded eastern Lake Erie during the late 1990s, and round goby population size increased dramatically during 1999–2004. According to annual bottom trawl survey results, round goby abundance in offshore waters peaked in 2004, but then declined during 2004–2008. Coincidentally, round goby became an important component of burbot diet beginning in 2003. Using bottom trawling and gill netting, we estimated adult burbot abundance and age structure in eastern Lake Erie during 2007. Diet composition and energy density of eastern Lake Erie burbot were also determined during 2007. This information, along with estimates of burbot growth, burbot mortality, burbot water temperature regime, and energy densities of prey fish from the literature, were incorporated into a bioenergetics model application to estimate annual consumption of round goby by the adult burbot population. Results indicated that the adult burbot population in eastern Lake Erie annually consumed 1,361 metric tons of round goby. Based on the results of bottom trawling, we estimated the biomass of yearling and older round goby in offshore waters eastern Lake Erie during 2007–2008 to be 2,232 metric tons. Thus, the adult burbot population was feeding on round goby at an annual rate equal to 61% of the estimated round goby standing stock. We concluded that the burbot population had high potential to exert predatory control on round goby in offshore waters of eastern Lake Erie.     

The genetic diversity of burbot (<Emphasis Type="Italic">Lota lota</Emphasis> L., 1758) of Western Siberia (the analysis of the mtDNA control region polymorphism)

The genetic variability of burbot (Lota lota L., 1758) inhabiting the Ob-Irtysh and Taz river basins in Western Siberia has been studied based on the polymorphism of the hypervariable fragment of mtDNA control region (407 bp). The analysis of 134 fish samples revealed 30 haplotypes, 23 of which were new. Among haplotypes, previously detected in Eurasia and North America, EB30 was the most frequently found in Western Siberia (45.5% frequency). The results of our study are in agreement with previous research pointing to the genetic differentiation of two burbot subspecies, L. l. lota and L. l. maculosa, and indicate that burbot inhabiting the Ob-Irtysh and Taz river basins belong to the Eurasian-Beringian clade (nominative subspecies L. l. lota). However, a high genetic diversity of burbot in Western Siberia, along with a relatively high differentiation of burbot groups within studied territory, points to a regional specificity of burbot population.     

Assessment of burbot Lota lota (L. 1758) population sustainability in central European reservoirs

A novel sampling scheme, using a combination of electrofishing, visual exploration by scuba divers, two types of fyke nets and longlines, was tested in four reservoirs (including their inlets and outlets) to monitor a population of burbot Lota lota. This was supplemented by fry trawling and vertical hydro‐acoustics, to detect L. lota larvae in two deep reservoirs that have had a long‐term stocking programme. The majority of the L. lota detected were juveniles, captured by electrofishing in the littoral zones of the reservoirs and in running waters. Older individuals were rarely captured with longlines or fyke nets in deeper zones or structured habitats within the reservoirs. A combination of multiple sampling methods provided an assessment of the whole population. Population establishment could not be demonstrated as the age structure of the sampled fish corresponded with that of the stocked fish. Low post‐stocking survival, migratory behaviour, interactions with other species and warmer water temperatures are considered the potential drivers for unsuccessful establishment of L. lota populations in these reservoirs.     

Food,growth, habitat,and community interactions of young-of-the-year burbot,Lota lota L., in a Precambrian Shield lake

The burbot, Lota lota, is a widely distributed gadid of the northern circumpolar regions of North America and Eurasia. Despite its near ubiquity over much of its range, relatively little is known about its biology during the first year of life.Burbot sac-fry of 3 mm total length, hatched under the ice in early May in Shebandowan Lake. Their first foods following atrophy of the yolk-sac were copepods and cladocerans which they captured pelagically. At first the fry swam in small schools, high in the water column of the near-shore littoral, and fed during the daytime. Upon reaching 30 mm in total length, the burbot fingerlings became solitary and benthic, and fed primarily at night, almost exclusively on the amphipod, Hyalella azteca. Amphipods constituted about 75 per cent by number of all the food consumed by burbot fingerlings in their first year of life. Growth was rapid from May to the end of July, tapered off during August to October, and effectively stopped by November at freeze-up. The principal habitat of burbot fingerlings during the benthic stages of their life was the near-shore littoral, in depths ranging from 0.5 m to 4.0 m. There, they were sheltered by boulder shoals, sunken trees or other debris, or beds of quillworts (Isoetes sp.). The principal ichthyofauna that cohabited with the burbot included the sculpin (Cottus cognatus), the smallmouth bass (Micropterus dolomieui), two etheostomatines (Etheostoma nigrum) and (E. exile) and the yellow perch (Perca flavescens). Burbot fingerlings were subjected to low levels of predation from the nocturnal foraging of walleyes (Stizostedion vitreum).     

Effects of temperature on sperm motility of burbot Lota lota: spontaneous activation and calcium dependency

Several factors regulating activation of spermatozoon motility in Eurasian burbot, Lota lota, including osmolality, calcium (Ca²⁺) ions, and temperature were investigated. Spermatozoon motility in Eurasian burbot, Lota lota was assessed at 4 and 30°C in seminal fluid, isotonic media (with and without Ca²⁺) and hypotonic media (with and without Ca²⁺). Spermatozoa were spontaneously activated in seminal fluid at 20°C and the maximum motility was recorded at 30°C, which is out of the spawning temperature range, indicating that no risk of activation occurs during routine semen handling in artificial insemination. Initiation of spermatozoon motility in L. lota is mediated by Ca²⁺ and sensitivity to Ca²⁺ is dependent on temperature.     

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.     

Phylogenetic relationships among Palearctic and Nearctic burbot (Lota lota): Pleistocene extinctions and recolonization

The burbot (Lota lota Linnaeus, 1758) is the only freshwater species from the cod family. Various taxonomic hypotheses were tested against molecular data by sequencing the mitochondrial cytochrome b locus of 120 burbot from 41 populations together with the related species Molva molva (ling) and Brosme brosme (tusk), which represented the other Lotinae genera. Within the genus Lota two distinct phylogroups were observed: one in North America south of the Great Slave Lakes (Lota lota maculosa) and one in Eurasia and the remainder of the Nearctic region (Lota lota lota). The burbot lineage separated 10 Myr BP from the other Lotinae, while the genetic variation within burbot appeared to be approximately 1 Myr old. However, fossil evidence suggested that burbot already existed in the Early Pliocene in Europe, from were it probably colonized North America in the Early Pleistocene. While Nearctic burbot survived climatic oscillations and diverged in several refugia, the Eurasian form became extinct or was reduced to a very small population. In the Late Pleistocene the species recolonized the Palearctic region to establish its present distribution range.     

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.

     

A synopsis of ‘hydropeaking’ studies on the response of juvenile Atlantic salmon to experimental flow alteration

In insular Newfoundland, Canada, studies were conducted from 1999 to 2003 on the effects of ‘simulated’ hydropeaking power generation on juvenile Atlantic salmon (Salmo salar). In 1999, Atlantic salmon parr were released into an experimental reach below a hydroelectric facility and flow was manipulated over a range of discharge (1.0–4.2 m³ s⁻¹) during a series of ‘experiments’ simulating hydropeaking in both summer and fall. Fish were implanted with radio transmitters, manually tracked, precisely located (±1 m), habitat selection evaluated, and movement response determined. Experiments were continued in 2002 and 2003 to contrast response of salmon between summer and winter, the magnitude of flow changes were greater (0.7–5.2 m³ s⁻¹) and changes were made more rapidly (instantly). As discharge was increased, velocity and depth use by parr increased, and fish adapted behaviourally by increased contact with the substrate. Salmon parr also exhibited two distinct movement patterns in the summer and fall of 1999 studies; high site fidelity or considerable movement during trials. Salmon were more mobile during both static and dynamic flow conditions and throughout the diel cycle in the summer of 2002 experiment, and 2 fish were stranded and died, the only time this happened in the four series of experiments. Within each experiment generally there were no differences between movements at static high and low flows for day and night movements, with one exception, and night time movements were always greatest, again with one exception. During dynamic flow changes, within each experiment, distances moved during down ramping and up ramping were not significantly different except in the summer of 1999. Overall, comparing between experiments for up and down ramping events, distances moved in the summer of 2002 were statistically higher than for all other experiments. Not surprisingly, the home ranges of fish in the summer of 2002 were also the greatest while the smallest home ranges were in the winter of 2003. Results suggest hydropeaking regimes may be energetically costly potentially affecting over-winter survival which is related to energy reserves obtained during summer. Collectively these studies provide comprehensive information on the response of juvenile Atlantic salmon parr to hydropeaking, on both diel and seasonal scales, and will assist hydro producers and regulators design and operate hydropeaking regimes to minimize ecological impact. Guest editors: R. L. Welcomme & G. Marmulla Hydropower, Flood Control and Water Abstraction: Implications for Fish and Fisheries     

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.     

Diurnal shifts in use of summer habitat by age-0 brown trout in a regulated mountain stream

Diurnal shifts in habitat use by age-0 (< 50 mm total length) brown trout (Salmo trutta) were observed from June to July 1989 and 1990 in low-gradient (< 1% channel slope) reaches of Douglas Creek, Medicine Bow National Forest, Wyoming. Fish were visually located during day and night, and water depth, water velocity, distance from the stream edge, substrate, habitat type, and cover were determined at each location. The fish were found predominantly in locations with slow water velocities in stream-margin and backwater-pool habitats. They were observed in slower water, closer to the stream edge at night than during the day.     

Similar distribution pattern of different phenotypes of <Emphasis Type="Italic">Limnocythere inopinata</Emphasis> (Baird) in a brackish-water lake in Inner Mongolia

It has been recognized long ago that Limnocythere inopinata is a widely distributed ostracod and comprises various phenotypes. The taxonomy of the different phenotypes and their ecological differences, however, are still under debate. In this study, we investigated the relationships between ostracod distribution, water depth, and the substrate based on the analyses of species composition, abundance, and age structure of ostracods in 51 surface-sediment samples from Dali Lake in Inner Mongolia in order to reveal the distribution pattern and habitat characteristics of the ostracods. The identification of a total of 32,182 valves indicates that the dominant ostracod in Dali Lake is L. inopinata, which includes unnoded, 1-, 2- and 3-node and 1-carina phenotypes. The distribution of L. inopinata is affected both by the water depth and by the sand content of the substrate, and the phenotypes show their maximum abundances in the intermediate zone of the lake. All the phenotypes of L. inopinata display a similar preference for water depth and substrate, implying that the occurrence of different phenotypes is controlled by other factors. The age structure of L. inopinata is mainly related to the sand content, with the percentage of adults increasing and that of early juveniles decreasing with increasing sand content. Our data suggest that the occurrence of L. inopinata is closely related to water depth and substrate grain size, and different phenotypes of L. inopinata respond in a similar pattern to water depth and substrate.     

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

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Burbot (Lota lota) exhibit plasticity in life-history traits in a small drainage at the southwestern-most extent of the species' native range

Little is known about the life-history traits exhibited by burbot (Lota lota) throughout their circumpolar range. Monitoring burbot movements between lentic and lotic habits and collection of demographic data (length, age, sex, and maturity) were used to answer the following questions in the Torrey Creek drainage of west-central Wyoming, USA: (a) is there plasticity in the life-history traits of the burbot population, (b) do Trail Lake origin and Torrey Creek origin burbot interchange during the spawning period, and (c) is there a difference in growth and age at sexual maturity between burbot captured in Trail Lake and Torrey Creek? Results indicated that burbot in Trail Lake and Torrey Creek exhibit plasticity in their life history traits. Directional movement of PIT-tagged burbot in Trail Lake and Torrey Creek was monitored nearly continuously by tandem stream-width antennas. Thirty-five percent of lentic-origin burbot migrated upstream into Torrey Creek, and 11% of lotic-origin burbot migrated downstream of the antennas near the Torrey Creek inlet to Trail Lake. Migratory activity of burbot was highest during the late winter and early spring at a time that coincided with spawning. Additionally, Torrey Creek-origin burbot were smaller and younger than Trail Lake-origin burbot. In addition to documenting migration, the sampling of small, sexually mature burbot in Torrey Creek suggests that stream-resident burbot reach maturity earlier than adfluvial and lacustrine burbot in Trail Lake. Furthermore, high catch rates of age-0 burbot indicate that Torrey Creek upstream from Trail Lake provides nursery habitat to the burbot population. Life history trait plasticity and spawning and nursery habitats documented in this study should be considered when selecting conservation actions for this unique burbot population.     

Relationship between current velocity,depth and the invertebrate community in a stable river system

The relationship between invertebrate densities, current velocity and water depth was studied in the Dan River, northern Israel. Maximum current preferences ranged from 5–120 cm sec^–1, and depth preferences ranged from 5–60 cm. Thirty-five taxa of invertebrates were collected by means of colonization cages. Larval and adult stages of 3 Elmidae (Coleoptera) species were treated separately: Limnius letourneuxi, Grouvellinus caucasicus and Elmis rioloides. Differences in current preference were observed between larval and adult stages of the same species of Elmidae. Taxa were also grouped according to preference for turbulence. Wide ranges of depth and current velocity preferences were observed. Most of the taxa were found at between 80–100 cm sec^–1 and at depths of less than 30 cm. A correlation between species diversity and current velocity was established. Velocities of 60–80 cm sec^–1 contained the greatest overlap of faunal preference. The sensitivity of selected species to stream flow reduction is discussed.IES Laboratory, Department of Zoology, Hebrew University; and the Nature Reserves Authority     

Oxygen consumption of burbot, Lota lota(L.), fed different rations of vendace, Coregonus albula L.

The oxygen consumption of adult burbot, Lota lota (L.), was examined to determine the effects of fasting and meal ration on oxygen consumption rates. The temperature (2.1 °C) was selected to represent ambient conditions for burbot in winter. The average pre‐feeding oxygen consumption rate was 29.5 mg kg^?1 h^?1. Feeding affected the oxygen consumption rates since the apparent heat increment correlated significantly with the meal ration. However, the meal ration (i.e. vendace, Coregonus albula L.) did not affect the proportion of the ingested meal energy used in the apparent heat increment. When the meal ration was increased, peak oxygen consumption ranged from 1.2 to 2.2 times higher than the fasting level. Observed low oxygen consumption values suggest that the metabolic processes of burbot are strongly reduced in low temperatures.     

Trends in body condition of native piscivores following invasion of Lakes Erie and Ontario by the round goby

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