Depth gradients in food‐web processes linking habitats in large lakes: Lake Superior as an exemplar ecosystem

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Diurnal vertical and seasonal horizontal distribution patterns of Mysis relicta in a large Norwegian lake

Mysis relicta was sampled intensively along a depth profilein Lake Snåsavatnet using benthic trawls and verticalnet hauls. Samples were made around 12.00 and 24.00 h in May,June, August and September 1984. The light regime and temperaturewere also recorded. Mysis relicta showed a size-dependent diurnalvertical migration and seasonal benthic horizontal migrationbetween shallow and deep areas. The young individuals left themarsupium in spring while the adults were staying benthic inshallow areas. As they grew in size the young individuals changedhabitat, migrating into the pelagic zone and the deeper benthicareas. Mysis showed a bimodal vertical distribution patternon our deepwater station, with one density peak in the watercolumn and another along the bottom both day and night betweenMay and September. The bimodal distribution pattern is discussedin relation to the light regime and an asynchronic verticalmigration model. The density of Mysis was low in the pelagiczone compared to the shallow benthic area throughout the samplingperiod. Together with the difference in distribution patternin the pelagic and benthic zones, this may influence the distributionpattern of salinonids which seek Mysis as a food item in LakeSnåsavatnet.     

Diel vertical migration and feeding rhythm of Daphnia longispina and Bosmina corexoni in Lake Toya, Hokkaido, Japan

Diel vertical migration (DVM) and diel feeding rhythm of two cladocerans, Daphnia longispina and Bosmina coregoni were investigated at the pelagic area of Lake Toya (Hokkaido, Japan) in May, August and October 1992. Both species performed nocturnal DVM. The amplitude of DVM, however, became smaller from May to October. Such seasonal variations in DVM could not be explained by light penetration and/or water temperature. The two species had a clear feeding rhythm; they fed at night in May and October but also after sunrise in August. These feeding rhythms appeared to be related to the light-dark cycle, but were not necessarily associated with their DVM. We suggest that the diel feeding rhythm and DVM are regulated independently by light cues.     

Behavioral,Ecological and Genetic Differentiation in an Open Environment—A Study of a Mysid Population in the Baltic Sea

Diel vertical migration (DVM) is often assumed to encompass an entire population. However, bimodal nighttime vertical distributions have been observed in various taxa. Mysid shrimp populations also display this pattern with one group concentrated in the pelagia and the other near the bottom. This may indicate alternative migratory strategies, resembling the seasonal partial migrations seen in birds, fishes and amphibians, where only a subset of the population migrates. To assess the persistence of these alternative strategies, we analyzed the nitrogen and carbon stable isotope signatures (as proxies for diet), biochemical indices (as proxies for growth condition), and genetic population divergence in the Baltic mysid Mysis salemaai collected at night in the pelagia and close to the bottom. Stable isotope signatures were significantly different between migrants (pelagic samples) and residents (benthic samples), indicating persistent diet differences, with pelagic mysids having a more uniform and carnivorous diet. Sequencing of the mitochondrial cytochrome subunit I (COI) gene showed genetic differentiation attributable to geographic location but not between benthic and pelagic groups. Divergent migration strategies were however supported by significantly lower gene flow between benthic populations indicating that these groups have a lower predisposition for horizontal migrations compared to pelagic ones. Different migration strategies did not convey measurable growth benefits as pelagic and benthic mysids had similar growth condition indices. Thus, the combination of ecological, biochemical and genetic markers indicate that this partial migration may be a plastic behavioral trait that yields equal growth benefits.     

Behavioural responses to co-occurring threats of predation and ultraviolet radiation in Daphnia

1. Organisms in the wild are faced with multiple threats and a common response is a change in behaviour. To disentangle responses to several threats, we exposed two differently sized species of the freshwater invertebrate Daphnia to solar ultraviolet radiation (UVR) and predation from either moving pelagic or benthic ambush predators.
2. Using an advanced nanotechnology-based method, we tracked the three-dimensional movements of those mm-sized animals at the individual level. Each behavioural trial was performed both under conditions resembling night (no UVR) and day (UVR) and we examined patterns of the depth distribution and swimming speed by Daphnia across three treatments: no predator (control); bottom-dwelling damselfly (Calopteryx sp.); and fish (stickleback, Pungitius pungitius) predators. We also quantified the actual predation rate by the two predators on the two Daphnia species, Daphnia manga and Daphnia pulex.
3. We show that individual Daphnia are able to identify predators with different feeding habitats, rank multiple and simultaneously occurring risks and respond in accordance with the actual threat; complex responses that are generally associated with larger animals.
4. In a broader context, our results highlight and quantify how a cocktail of everyday threats is perceived and handled by invertebrates, which advances our understanding of species distribution in space and time, and thereby of population dynamics and ecosystem function in natural ecosystems.

     

The impact of diel vertical migration of <Emphasis Type="Italic">Daphnia</Emphasis> on phytoplankton dynamics

Diel vertical migration (DVM) of large zooplankton is a very common phenomenon in the pelagic zone of lakes and oceans. Although the underlying mechanisms of DVM are well understood, we lack experimental studies on the consequences of this behaviour for the zooplankton’s food resource—the phytoplankton. As large zooplankton species or individuals migrate downwards into lower and darker water strata by day and upwards into surface layers by night, a huge amount of herbivorous biomass moves through the water column twice a day. This migration must have profound consequences for the phytoplankton. It is generally assumed that migration supports an enhanced phytoplankton biomass and a change in the composition of the phytoplankton community towards smaller, edible algae in the epilimnion of a lake. We tested this assumption for the first time in field experiments by comparing phytoplankton biomass and community assemblage in mesocosms with and without artificially migrating natural stocks of Daphnia hyalina. We show that DVM can enhance phytoplankton biomass in the epilimnion and that it has a strong impact on the composition of a phytoplankton community leading to an advantage for small, edible algae. Our results support the idea that DVM of Daphnia can have strong effects on phytoplankton dynamics in a lake.     

Benthic carbon is inefficiently transferred in the food webs of two eutrophic shallow lakes

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Diel feeding ecology of the nymphs of Aeshna multicolor and Lestes unguiculatus (Odonata)

SUMMARY.

• 1 Diel feeding periodicity and diet composition of nymphs of Aeshna multicolor and Lestes unguiculatus were examined at 6 h intervals in a small pond. Feeding intensity of A. multicolor increased from 06.00 hours to approximately 18.00 hours whereas food intake by L. unguiculatus was uniformly high between 12.00 and 18.00 hours. Feeding of both species was reduced during the night.
• 2 Diel variation in diet composition was substantial in each species. Chironomid larvae were the major prey (48.9%) in the diet of A. multicolor. Zooplankton, mainly cladocerans (42.0%) and ostracods (21.1%), were the principal prey of L. unguiculatus. Detritus composed a substantial portion of the diets of both species.
• 3 Diet overlap (Cλ) at each 6 h interval was high (0.43–0.72) when detritus was included in the calculation and was low (0.48–0.54) when detrital material was omitted.

     

Climate and productivity shape fish and invertebrate community structure in subarctic lakes

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Hunt warm, rest cool: bioenergetic strategy underlying diel vertical migration of a benthic shark

1. Diel vertical migration (DVM) is a widespread phenomenon among marine and freshwater organisms and many studies with various taxa have sought to understand its adaptive significance. Among crustacean zooplankton and juveniles of some fish species DVM is accepted widely as an antipredator behaviour, but little is known about its adaptive value for relatively large-bodied, adult predatory fish such as sharks. Moreover, the majority of studies have focused on pelagic forms, which raises the question of whether DVM occurs in bottom-living predators. 2. To investigate DVM in benthic predatory fish in the marine environment and to determine why it might occur we tracked movements of adult male dogfish (Scyliorhinus canicula) by short- and long-term acoustic and archival telemetry. Movement studies were complemented with measurements of prey abundance and availability and thermal habitat within home ranges. A thermal choice experiment and energy budget modelling was used to investigate trade-offs between foraging and thermal habitat selection. 3. Male dogfish undertook normal DVM (nocturnal ascent) within relatively small home ranges (-100 x 100 m) comprising along-bottom movements up submarine slopes from deeper, colder waters occupied during the day into warmer, shallow prey-rich areas above the thermocline at night. Few daytime vertical movements occurred. Levels of activity were higher during the night above the thermocline compared to below it during the day indicating they foraged in warm water and rested in colder depths. 4. A thermal choice experiment using environmentally realistic temperatures supported the field observation that dogfish positively avoided warmer water even when it was associated with greater food availability. Males in laboratory aquaria moved into warm water from a cooler refuge only to obtain food, and after food consumption they preferred to rest and digest in cooler water. 5. Modelling of energy budgets under different realistic thermal-choice scenarios indicated dogfish adopting a 'hunt warm - rest cool' strategy could lower daily energy costs by just over 4%. Our results provide the first clear evidence that are consistent with the hypothesis that a benthic marine-fish predator utilizes DVM as an energy conservation strategy that increases bioenergetic efficiency.     

Hormone‐mediated foraging strategies in an uncertain environment: Insights into the at‐sea behavior of a marine predator

1. Hormones are extensively known to be physiological mediators of energy mobilization and allow animals to adjust behavioral performance in response to their environment, especially within a foraging context.
2. Few studies, however, have narrowed focus toward the consistency of hormonal patterns and their impact on individual foraging behavior. Describing these relationships can further our understanding of how individuals cope with heterogeneous environments and exploit different ecological niches.
3. To address this, we measured between‐ and within‐individual variation of basal cortisol (CORT), thyroid hormone T3, and testosterone (TEST) levels in wild adult female Galápagos sea lions (Zalophus wollebaeki) and analyzed how these hormones may be associated with foraging strategies. In this marine predator, females exhibit one of three spatially and temporally distinct foraging patterns (i.e., “benthic,” “pelagic,” and “night” divers) within diverse habitat types.
4. Night divers differentiated from other strategies by having lower T3 levels. Considering metabolic costs, night divers may represent an energetically conservative strategy with shorter dive durations, depths, and descent rates to exploit prey which migrate up the water column based on vertical diel patterns.
5. Intriguingly, CORT and TEST levels were highest in benthic divers, a strategy characterized by congregating around limited, shallow seafloors to specialize on confined yet reliable prey. This pattern may reflect hormone‐mediated behavioral responses to specific risks in these habitats, such as high competition with conspecifics, prey predictability, or greater risks of predation.
6. Overall, our study highlights the collective effects of hormonal and ecological variation on marine foraging. In doing so, we provide insights into how mechanistic constraints and environmental pressures may facilitate individual specialization in adaptive behavior in wild populations.

     

Comparing two fish sampling standards over time: largely congruent results but with caveats

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Diel vertical migration by males,females, copepodids and nauplii in a limnetic population of diaptomus (Copepoda)

Diel vertical migration (DVM) by the sexes and life history stages of the copepod Diaptomus novamexicanus (Herrick) was investigated during two summers in a subalpine lake (Castle Lake, California, USA). Migratory behavior was quantified repeatedly by 1300 and 21–2200 hr sampling in the epilimnion. Population density and vertical distribution were estimated routinely at 1300 hr and at 2200 hr on selected dates. Males had greater night to day biomass differentials and ratios than females. The sexes showed comparable DVM participation (% of total biomass migrating) and amplitudes of migration (distance of ascent). Copepodids displayed higher participation, night/day ratios and amplitudes than younger copepods, but adult participation differed only from that of nauplii. Active participation in high amplitude ascents thus increased with age and was greatest in adult males. Analysis suggests that such relative behaviors probably occur in most limnetic Diaptomus, and that the magnitude of ascent into the epilimnion is associated with daytime vertical distribution.     

Distribution patterns of fishes in a canyon-shaped reservoir

In August 2004 and 2005, an extensive study of the fish community was carried out in the largest water supply reservoir in the Czech Republic and Central Europe, the canyon‐shaped ?elivka Reservoir, using a fleet of Nordic multimesh gillnets. Fishes were sampled at eight locations along the longitudinal profile of the reservoir and at five benthic depth layers covering depths from the surface down to 18 m (benthic gillnet 1·5 m high), and at three pelagic depth layers down to the depth of 5 m above the bottom (pelagic gillnets 4·5 m high). Catches of both juvenile (age 0+ year) and adult (fishes >1 year) fishes were highest in the upper layers of the water column (i.e. in the epilimnion down to 5 m, and down to 10 m in the benthic habitats). Along the tributary–dam axis in the pelagic habitats, both juvenile and adult fishes preferred the upper part of the reservoir, where the maximum number of species and also the greatest abundance of zooplankton were found. In the benthic habitats, fishes selected location according to factors other than trophic status. More juvenile fishes were recorded in the benthic habitats than in the pelagic habitats. Depth had the largest explanatory power for predicting fish community composition, followed by the affiliation with benthic and pelagic habitats, and location on the longitudinal axis of the reservoir. The fish community was represented mainly by cyprinids and consisted of two distinct groups of species, with bleak Alburnus alburnus, rudd Scardinius erythrophthalmus and asp Aspius aspius dominating the offshore group while perch Perca fluviatilis and ruffe Gymnocephalus cernuus were affiliated with the inshore group of the adult fish community. Roach Rutilus rutilus, bream Abramis brama and pikeperch Sander lucioperca occurred in important proportions in both the inshore and the offshore zones. All species, with the exceptions of adult perch (1+ year and older), 0+ year perch and 0+ year roach, preferred the most eutrophic tributary part of the reservoir. The fish community was relatively stable between the 2 years sampled.     

Habitat use of arctic charrSalvelinus alpinus in Thingvallavatn,Iceland

Synopsis Habitat use by four morphs of arctic charr,Salvelinus alpinus, was investigated in Thingvallavatn, Iceland, by sampling with pelagic and benthic gill nets. Sampling was done in May/June and August/September. Greatest abundance of fish was recorded in the littoral and epipelagic zone in early autumn. Catches were low in early summer. The four morphs are partly segregated in habitat. Small (SB-) and large benthivorous (LB-) charr have a more restricted spatial distribution than piscivorous (PI-), and especially planktivorous (PL-) charr. Both benthivorous morphs are mainly found in the littoral zone, and occur in largest numbers in stony shallows at depths between 0 and 10 m. PL-charr usually dominates in numbers in all habitats. PI-charr is most abundant in epibenthic habitats, although numbers are always low. All morphs are caught in higher numbers at night than during the day, but the diurnal activity difference is highest among SB-charr. The habitat use by different morphs is as may be expected from their morphology and diets. Within the population of PL-charr, young and small fish are more abundant on the bottom than in the pelagic zone, and there is a surplus of females in the pelagic zone. Along the benthic profile, young, small and immature PL-charr are more abundant in deep than in shallow waters. The results are discussed in relation to food supply, competition and predation. Possible reasons for the occurrence of four arctic charr morphs are also discussed.Contribution from the Thingvallavatn project.     

Toxic benthic freshwater cyanobacterial proliferations: Challenges and solutions for enhancing knowledge and improving monitoring and mitigation

1. This review summarises knowledge on the ecology, toxin production, and impacts of toxic freshwater benthic cyanobacterial proliferations. It documents monitoring, management, and sampling strategies, and explores mitigation options.
2. Toxic proliferations of freshwater benthic cyanobacteria (taxa that grow attached to substrates) occur in streams, rivers, lakes, and thermal and meltwater ponds, and have been reported in 19 countries. Anatoxin- and microcystin-containing mats are most commonly reported (eight and 10 countries, respectively).
3. Studies exploring factors that promote toxic benthic cyanobacterial proliferations are limited to a few species and habitats. There is a hierarchy of importance in environmental and biological factors that regulate proliferations with variables such as flow (rivers), fine sediment deposition, nutrients, associated microbes, and grazing identified as key drivers. Regulating factors differ among colonisation, expansion, and dispersal phases.
4. New -omics-based approaches are providing novel insights into the physiological attributes of benthic cyanobacteria and the role of associated microorganisms in facilitating their proliferation.
5. Proliferations are commonly comprised of both toxic and non-toxic strains, and the relative proportion of these is the key factor contributing to the overall toxin content of each mat.
6. While these events are becoming more commonly reported globally, we currently lack standardised approaches to detect, monitor, and manage this emerging health issue. To solve these critical gaps, global collaborations are needed to facilitate the rapid transfer of knowledge and promote the development of standardised techniques that can be applied to diverse habitats and species, and ultimately lead to improved management.

     

Taxonomic and functional homogenisation of macroinvertebrate communities in recently intermittent Alpine watercourses

1. Mountain streams in southwestern European Alps are currently shifting from perennial to intermittent flow due to the combined effects of climate change and local anthropogenic pressures. Given that flow intermittency is a recently documented phenomenon in the Alps, only scattered studies have investigated functional and taxonomical diversity of benthic invertebrate communities in recently intermittent Alpine streams.
2. We used a hierarchical sampling design to investigate patterns in taxonomic and functional diversity of benthic invertebrate communities in 13 recently intermittent Alpine streams in north-west Italy. in April 2017, we sampled benthic communities in two reaches of each stream with different hydrological conditions: a control reach, with permanent flow; and an intermittent reach, which recently experienced non-flow periods in summer.
3. We tested for the response of taxonomic richness at multiple spatial scales by partitioning total diversity into the average richness of local communities and the richness due to variation among local communities both within and among reaches. By partitioning total diversity (γ) into its local (α) and turnover (β) components we showed a decrease in local and regional species richness both within and among reaches, whereas variation among communities was significantly lower in intermittent reaches at the reach scale only.
4. The analysis of multidimensional trait space of macroinvertebrate communities in reaches with different hydrological conditions revealed a significant reduction of functional diversity, dispersion, and evenness in intermittent reaches. There was trait overdispersion in intermittent reaches, as these hosted both typical Alpine taxa and organisms adapted to flow intermittency. In particular, we observed the replacement of taxa with aquatic respiration and those preferring medium- to fast-flowing oligotrophic waters by taxa adapted to lentic habitats, air breathing and with larval dormancy phases.
5. These results indicate that recent flow intermittency has caused drastic changes in benthic invertebrate communities in Alpine streams. Our work highlights the importance of integrating taxonomic and functional diversity to thoroughly assess the impacts of flow intermittency.

     

Life history, growth and production of Mysis relicta in the large, fiord-type Lake Mjøsa, Norway

SUMMARY.

• 1 Biomass, production and life history of Mysis relicta were studied in the large Lake Mjøsa during the years 1976–80. Biomass fluctuations were large, but biomass averaged about 1 g wet wt m^?2 or 200 ind. m^?2. Cumulative net production during summer ranged from 1.6 to 2.1 g wet wt m^?2. Mean population P/B-ratio was close to 2.
• 2 Neither production or biomass of Mysis were correlated with the biomass of phytoplankton or the main Zooplankton prey species (Daphnia and Bosmina), suggesting a predominant ‘top down’ control of Mysis in this lake.
• 3 The life cycle was well synchronized with development of food items, but predation pressure also affected life-history patterns. The proportion of juveniles apparently increased in years with strong fish predation pressure, producing a predominantly 1-year life-cycle pattern the following year.
• 4 Mysis had a predominant 2-year life history, but a traction of the population may reproduce after 1 or 3 years. Reproduction occurred exclusively during winter. Even in such a large and stable system, where Mysis has coexisted with its main food items and predators for 8–10 000 years, a flexible life history is maintained and is probably an important buffer against year-to-year fluctuations in food and predator abundance.

     

Diel differences in fish assemblages in nearshore eelgrass and kelp habitats in Prince William Sound,Alaska

The importance of a particular habitat to nearshore fishes can be best assessed by both diurnal and nocturnal sampling. To determine diel differences in fish assemblages in nearshore eelgrass and understory kelp habitats, fishes were sampled diurnally and nocturnally at six locations in western Prince William Sound, Alaska, in summer 2007. Abundance of fish between day and night were similar, but species composition and mean size of some fish changed. Species richness and species diversity were similar in eelgrass during the day and night, whereas in kelp, species richness and species diversity were greater at night than during the day. In eelgrass, saffron cod (Eleginus gracilis) was the most abundant species during the day and night. In kelp, the most abundant species were Pacific herring (Clupea pallasii) during the day and saffron cod at night. Diel differences in fish size varied by species and habitat. Mean length of saffron cod was similar between day and night in eelgrass but was greatest during the day in kelp. Pacific herring were larger at night than during the day in kelp. Diel sampling is important to identity nearshore habitats essential to fish and help manage fish stocks at risk.     

Diel vertical migration of the copepod <Emphasis Type="Italic">Thermocyclops inversus</Emphasis> (Kiefer, 1936) in a tropical reservoir: the role of oxygen and the spatial overlap with <Emphasis Type="Italic">Chaoborus</Emphasis>

The effect of water transparency, dissolved oxygen concentration and the invertebrate predator Chaoborus brasiliensis on the day–night vertical distribution of the copepod cyclopoid Thermocyclops inversus was investigated in a shallow tropical reservoir, Nado Reservoir, Belo Horizonte, Brazil. Diel cycles were carried out over a period of 12 consecutive months, between October 1999 and September 2000. The different developmental stages of T. inversus exhibited diel vertical migration (DVM) and displayed a clear ontogenetic trend, with the amplitude of DVM increasing with the age of the organism, and ranging from 0.4 m to 0.8 m for nauplii, 0.4 m to 1.2 m for copepodite, and 1.1 m to 2.1 m for adults. We observed that seasonal changes in dissolved oxygen and C. brasiliensis directly influenced the vertical distribution of the copepod population in this reservoir. Furthermore, it was showed that the diurnal vertical migration is an important predator avoidance behavior since it diminished the spatial overlap between prey and its potential predator. This finding supports the hypothesis that the vertical migration is a defense mechanism against predation. Thus, T. inversus is able to remain in the anoxic layers during day light hours, and at night they move upwards avoiding hypolimnetic waters to escape from predation by Chaoborus.