New host species and new distribution areas of <Emphasis Type="Italic">Salmincola lavaretus</Emphasis> (Copepoda: Lernaeopodidae): A parasite of coregonid fishes of Siberia

Data on the infestation of certain fish species by the parasitic copepod Salmincola lavaretus are presented for the first time. The infested fish species included Bauntovsky whitefish (Coregonus baunti), an endemic species from the Maloye and Bolshoye Kapylyushi lakes; Siberian whitefish (C. pidschian) and Siberian cisco (C. sardinella) from Bolshoye Kapylyushi Lake and Baunt Lake, which belong to the Tsypo-Tsypikan lake system (the Lena River basin, Transbaikalia); and Teletsky whitefish (C. lavaretus natio smitti) from Teletskoe Lake (the Ob River basin). Previously, S. lavaretus had been described as a parasite of Baikal omul (C. migratorius) and Baikal whitefish (C. baicalensis) from Baikal. The taxons of S. longimanus complex (S. longimanus, S. l. sibirica, S. svetlanovi, and S. lavaretus)—parasites of nasal fossae of grayling and coregonid fishes—were registered in the lakes of the Lake Baikal basin (Lake Baikal and Khovsgol Lake), the Lena River, the Yenisei and the Ob rivers (the largest rivers of the Arctic zoogeographic province), and the Kobdo River (the Western Mongolian province). It was assumed that S. longimanus complex is more widely distributed in the water bodies of the Arctic zoogeographic province of the Palearctic.     

A laboratory study of host use by the cuckoo catfish <Emphasis Type="Italic">Synodontis multipunctatus</Emphasis>

The only known non-avian vertebrate obligate brood parasite is the cuckoo catfish (Synodontis multipunctatus), a Lake Tanganyikan endemic. The cuckoo catfish parasitizes Tanganyikan mouthbrooding cichlids, and under captive conditions, will also parasitize cichlids from other Rift Valley lakes. Here we examine the frequency of parasitism by the cuckoo catfish of Ctenochromis horei from Lake Tanganyika and three species from Lake Malawi and the greater Lake Victorian system in a laboratory setting. C. horei was parasitized significantly less (17%) than the allopatric species Haplochromis latifasciatus, Haplochromis nubilus, and Metriaclima estherae (combined parasitism rate of 28%). The lower rates of parasitism in C. horei may be due to differences in the mating ritual, oviposition (e.g., long periods of pseudo-spawning before actual oviposition), and behavioral adaptations (e.g., increased aggression towards the cuckoo catfish). The number of catfish eggs per parasitized brood was similar between C. horei, H. latifasciatus, H. nubilus, and M. estherae. Our results are comparable to findings from the field for C. horei parasitism frequency and number of cuckoo catfish per brood. We also analyzed the parasitism rate of the albino morph of Metriaclima zebra, a domestic strain. Parasitism rates and number of catfish per brood were the highest in the albino morphotype suggesting that the higher levels of parasitism may be related to lower aggressive behavior, lower visual acuity, or captive influence. Cuckoo catfish and mouthbrooding cichlids provide a model system for testing brood parasitism in a laboratory setting.     

Concurrent evolution of ancient sister lakes and sister species: the freshwater gastropod genus <Emphasis Type="Italic">Radix</Emphasis> in lakes Ohrid and Prespa

Ancient sister lakes are considered to be ancient lakes lying in close geographic proximity, sharing a related origin and significant time of co-existence, usually having hydrological connection as well as a balanced degree of faunal overlap and distinctness. A paradigm for studying sister lake relationships are the ancient lakes Ohrid and Prespa in the Balkans, which are characterized by high degrees of endemicity. Three general patterns of endemic species can be distinguished for these lakes: (1) taxa that are endemic to either lake, with no close relatives in the respective sister lake, (2) closely related but distinct endemic taxa in both lakes (sister species) and (3) shared endemic taxa occurring in both lakes. In the present paper, two endemic freshwater pulmonate gastropod species, Radix relicta (Lake Ohrid) and R. pinteri (Lake Prespa), are used to study the evolution of presumed sister species based on biogeographical and comparative DNA data from world-wide Radix taxa. Phylogenetic, phylogeographical and parametric bootstrap analyses all suggest a sister group relationship of R. relicta and R. pinteri (pattern 2 of endemic diversity). Sister to these two taxa is the widespread R. ampla, which does not occur in the vicinity of lakes Ohrid and Prespa. The southern feeder spring complexes of Lake Ohrid are inhabited by another lineage (Radix sp. 1), which resembles Radix relicta in morphology/anatomy. For Lake Prespa, the widespread R. auricularia was reported in addition to the endemic R. pinteri. Comparative phylogenetic data favour a western Adriatic zoogeographical affinity of lakes Ohrid and Prespa over an Aegean-Anatolian faunal connection. The status of lakes Ohrid and Prespa as sister lakes is evaluated in the light of current knowledge on gastropod speciation and endemism in these hotspots of biodiversity.     

Comparative analyses of physiological assays and chlorophyll <Emphasis Type="Italic">a</Emphasis> variable fluorescence parameters: investigating the importance of phosphorus availability in oligotrophic and eutrophic freshwater systems

Diverse measurements of nutrient status indicators were used to test the severity of physiological phosphorus (P) limitation of phytoplankton among lake systems ranging from oligotrophic to eutrophic, based on P and chlorophyll a (Chl a) concentrations. Metabolic assays and particulate nutrient ratios were used to estimate nutrient status at sites located in Lake Erie, Lake Ontario and Lake Huron. Variable fluorescence ratios (F _v/F _m), relative electron transport rates and their response to irradiance were measured by the pulse-amplitude-modulated fluorometer. Under summer stratified conditions, P deficiency was strongest in the oligotrophic sites and nitrogen (N) status indicators and Chl a variable parameters revealed no severe N deficiency. Nutrient amendment assays showed positive associations with P additions and Chl a fluorescence parameters at P-deficient sites. In the most oligotrophic sites, N additions revealed a modest increase only detected by the Chl a fluorescence parameters. Phytoplankton communities were also associated with nutrient status, where chrysophytes and cryptophytes were important in P-deficient sites and cyanobacteria, phyrrophyta, and diatoms were prevalent in nutrient-rich sites. The results confirmed that Chl a fluorescence parameters can reveal P deficiency and indicate its severity among the range of trophic status in aquatic systems.     

Development of subfossil <Emphasis Type="Italic">Daphnia</Emphasis> and <Emphasis Type="Italic">Chaoborus</Emphasis> assemblages in relation to progressive acidification and fish community alterations in SW Sweden

Dated sediment cores from acidified and fishless Lake Gaffeln and Lake Härsvatten, SW Sweden, were analyzed for Daphnia ephippia and Chaoborus mandibles to test whether acidification history and fish extirpations could be reconstructed in a paleo-study using these easily identifiable animal remains. According to monitoring data fish were lost in both lakes from the 1950s to the 1970s. Progressive acidification prior to monitoring was confirmed by a gradual decrease and eventual loss of Daphnia ephippia in both study lakes during the first half of the twentieth century. In Lake Gaffeln mandibles of C. obscuripes appeared immediately after fish loss in 1973, and the regular presence of this species confirmed the succeeding fishless state of this lake. In Lake Härsvatten sediments C. obscuripes appeared only recently, i.e. three decades after fish extirpation, showing that the absence of C. obscuripes mandibles is not a trustworthy indicator of fish presence. Hence, the appearance of C. obscuripes was not temporally related to fish loss but confirmed the present fishless condition. Known historical presence of cyprinid fish in Lake Gaffeln was confirmed by a significantly higher proportion of fragmented mandibles of C. flavicans compared to the historically cyprinid-free Lake Härsvatten. In addition, both lake profiles displayed zero-proportions of fragmented mandibles during fishless periods. We conclude that acidification history and fish extirpations can be inferred by integrated studies on subfossil Daphnia ephippia and Chaoborus mandibles. However, during extreme ultra-oligotrophic conditions in acidified clear-water lakes subfossil Chaoborus mandibles may be too scarce to infer fish absence/presence.     

Distribution of sediment bacterial and archaeal communities in plateau freshwater lakes

Both Bacteria and Archaea might be involved in various biogeochemical processes in lacustrine sediment ecosystems. However, the factors governing the intra-lake distribution of sediment bacterial and archaeal communities in various freshwater lakes remain unclear. The present study investigated the sediment bacterial and archaeal communities in 13 freshwater lakes on the Yunnan Plateau. Quantitative PCR assay showed a large variation in bacterial and archaeal abundances. Illumina MiSeq sequencing illustrated high bacterial and archaeal diversities. Bacterial abundance was regulated by sediment total organic carbon and total nitrogen, and water depth, while nitrate nitrogen was an important determinant of bacterial diversity. Proteobacteria, Acidobacteria, Actinobacteria, Bacteroidetes, Chlorobi, Chloroflexi, Cyanobacteria, Firmicutes, Gemmatimonadetes, Nitrospirae, Planctomycetes, and Verrucomicrobia were the major components of sediment bacterial communities. Proteobacteria was the largest phylum, but its major classes and their proportions varied greatly among different lakes, affected by sediment nitrate nitrogen. In addition, both Euryarchaeota and Crenarchaeota were important members in sediment archaeal communities, while unclassified Archaea usually showed the dominance.     

Phytoplankton successions under ice cover in four lakes located in North-Eastern Sweden: Effects of liming

Phytoplankton successions under ice cover (January–March) were determined in four oligotrophic lakes (Burtjärn, Aspen, Vialamptjärn and Storkorstjärn) located in North-Eastern Sweden. The total phosphorus concentration in the lakes was less than 10 µg/L. Lake Burtjärn (reference lake) had a similar hydrology as Lake Aspen. Storkorstjärn and Vialamptjärn were of similar hydrology and had heavily colored water (>100 mg_Pt/L). Aspen as well as Vialamptjärn became continuously limed with calcium carbonate annually during the last decades. Biodiversity was considerably higher in the limed lakes (Aspen and Vialamptjärn) than in the untreated lakes (Burtjärn and Storkorstjärn). In Lake Burtjärn the most frequent species wereRhodomonas lacustris, Tabellaria flocculosa andBotryococcus braunii. Cryptophyceae (R. lacustris andCryptomonas marssonii) andDinophyceae (especiallyGymnodinium lantzschii) were common phytoplankton groups in Lake Aspen.Tabellaria flocculosa was also the most common organism in both humic lakes Storkorstjärn and Vialamptjärn, other phytoplankton groups were in the humic lakes scarce. Liming was found to have profound effects on phytoplankton communities studied.     

Decrease in young-of-the-year yellow perch growth rates following <Emphasis Type="Italic">Bythotrephes longimanus</Emphasis> invasion

Bythotrephes longimanus, an invasive zooplankter from Eurasia, has caused severe declines in native zooplankton communities in Rainy and Kabetogama lakes in northern Minnesota. Both lakes have experienced a 40–60% decrease in peak summer zooplankton biomass following B. longimanus establishment around 2006–2007. In these lakes, yellow perch (Perca flavescens) are a key fishery species, and young-of-the-year (YOY) yellow perch are mainly planktivorous during their first summer. This led to concern that their growth could be detrimentally affected by the depletion of zooplankton forage. We used seining data to compare growth rates of YOY yellow perch before (2001–2005) and after (2008–2012) B. longimanus establishment in Rainy and Kabetogama lakes. Nearby Lake Vermilion, assumed to have been unaffected by B. longimanus during this time period, was used as a reference for natural variation in YOY growth in the region. YOY yellow perch length was modeled as a linear function of cumulative growing degree days (GDD) throughout the summer, and the slope of the relationship was compared between pre- and post-B. longimanus time periods for the three study lakes. The two lakes with B. longimanus showed similar decreases in YOY yellow perch growth rate relative to GDD, whereas Lake Vermilion showed no evidence of a decline in growth rates during this period. The reduction in growth rates resulted in an approximate 10% decrease in mean length of YOY yellow perch at the end of the summer after B. longimanus establishment, which could lead to further effects of this invasive zooplankter at higher trophic levels.     

Identification of extremely halophilic archaea associated with adult <Emphasis Type="Italic">Artemia urmiana</Emphasis>

The brine shrimp, Artemia is the dominant macrozooplankton present in many hypersaline environments. Artemia urmiana is the only macroscopic organism in Urmia Salt Lake (Iran), and the high salinity of the lake makes it a suitable environment for halophilic archaea too. Because of common environment for Artemia and extreme halophiles; this investigation is concentrated on studying the relationship between Artemia and halophilic archaea in Urmia Lake. In this study first the procedure of arhaea isolation was done. Then, isolated strains were sub-cultured and DNA was extracted and amplified by PCR using specific primers for amplifying archaeal 16S rRNA. The amplified archeal DNA fragments were purified, and sequenced. 16S rRNA sequences were compared to known sequences using the NCBI BLAST program. Sequences relating to Halorubrum, Haloarcula and Halobacterium species were identified in Urmia Salt Lake water and Artemia adults and the phylogenetic tree of different species was constructed. Only Halorubrum species were present in association with Artemia. They belong to Halobacteriaceae family of archeae which are isolated from different salt lakes in different parts of world and we could show their existence in adult Artemia, another organism living in hypersaline enviroments.     

Winter ecology of shallow lakes: strongest effect of fish on water clarity at high nutrient levels

While the structuring role of fish in lakes is well studied for the summer season in North temperate lakes, little is known about their role in winter when fish activity and light irradiance potentially are lower. This is unfortunate as the progressing climate change may have strong effects on lake winter temperature and possibly on trophic dynamics too. We conducted an enclosure experiment with and without the presence of fish throughout winter in two shallow lakes with contrasting phosphorus concentrations. In hypertrophic Lake Søbygård, absence of fish led to higher biomass of zooplankton, higher grazing potential (zooplankton:phytoplankton ratio) and, accordingly, lower biomass of phytoplankton and chlorophyll a (Chl a), while the concentrations of total nitrogen (TN), total phosphorus (TP), oxygen and pH decreased. The average size of egg-bearing Daphnia and Bosmina and the minimum size of egg-bearing specimens of the two genera rose. In the less eutrophic Lake Stigsholm, zooplankton and their grazing potential were also markedly affected by fish. However, the decrease in Chl a was slight, and phytoplankton biovolume, pH and the oxygen concentration were not affected. TN was higher when fish were absent. Our results indicate that: (i) there is a notable effect of fish on zooplankton community structure and size during winter in both eutrophic and hypertrophic North temperate lakes, (ii) Chl a can be high in winter in such lakes, despite low light irradiance, if fish are abundant, and (iii) the cascading effects on phytoplankton and nutrients in winter may be more pronounced in hypertrophic lakes. Climate warming supposedly leading to reduced winter mortality and dominance of small fish may enhance the risk of turbid state conditions in nutrient-enriched shallow lakes, not only during the summer season, but also during winter.     

Secondary contact between two divergent lineages of charrs of the genus <Emphasis Type="Italic">Salvelinus</Emphasis> in the Northwest Pacific

The present-day contact zone between the Beringian and Arctic phylogenetic lineages of charrs of the genus Salvelinus in the Northwest Pacific is documented. A comparative analysis of the genetic differentiation and divergence indices for allopatric and sympatric populations of charrs and phylogenetic and genealogical analyses of the mtDNA haplotypes indicate that Lake Achchen and the Lake Pekulineiskoe are the zones of secondary contact between S. m. malma and S. taranetzi; Lake Nachikinskoe, between S. m. malma and Salvelinus sp. 4; and Lake Dal’nee, between S. m. malma and S. krogiusae. The level of divergence between phylogenetic groups of haplotypes considerably exceeds the range of intraspecific variability of S. m. malma and could not have been achieved after colonization of the lakes in conditions of sympathy. The obtained data suggest that the territory of Kamchatka was colonized by the common ancestor of the Arctic phylogroup of Taranetz charr.     

Parasitism (Trematoda,Digenea) in medusae from the southwestern Atlantic Ocean: medusa hosts,parasite prevalences,and ecological implications

Digenean are important endoparasites of fish with complex life cycles; some genera include medusae as secondary hosts. Their transmission to fish occurs when fish prey on these jelly hosts. Fish predation on jellyfish is a widespread phenomenon, even though predation by fish on jellyfish has not been determined through parasitism yet. We hypothesized that medusae with high prevalences of digeneans could be important for their transmission to fish. A total of 48,900 specimens of 50 medusa species were analyzed; 2,181 harbored digeneans. Opechona sp. and Monascus filiformis were the most frequent and abundant parasites with the widest range of hosts. Hemiuridae gen. sp. and Bacciger sp. were found in few specimens of some medusa species. Prevalences were unevenly distributed in the region. Three groups with high prevalence values were identified mainly related to frontal areas: Río de la Plata, Bahía Blanca, and North Patagonian tidal front. Eucheilota ventricularis, Clytia hemisphaerica, Proboscidactyla mutabilis, Liriope tetraphylla, and Aequoerea spp. were the medusae that contributed the most as secondary hosts to M. filiformis and Opechona sp. The high prevalences found in these medusae suggest that may be a fundamental part of the life cycles of both parasites in these areas.     

Vertical distribution of the relic species <Emphasis Type="Italic">Eurytemora lacustris</Emphasis> (Copepoda,Calanoida) in stratified mesotrophic lakes

The aim of this study was to determine factors affecting the vertical distribution of Eurytemora lacustris in mesotrophic lakes (Wigry, Hańcza, Szurpi?y; north-eastern Poland) during the summer stagnation. Eurytemora lacustris was found in all of the studied lakes, with the highest abundance (8 ind. L^?1) in Lake Wigry. In Lake Szurpi?y, E. lacustris has never been recorded before. The results of this study revealed that E. lacustris was most numerous in thermocline zones, suggesting that this species could temporarily tolerate warmer water and lower oxygen concentrations due to better food resources. During the study, it was found that a large part of the E. lacustris population had epibiont ciliates, in contrast to other species of zooplankton that did not have any epibionts. The improvement in the water quality of many deep lakes could lead to an increase in the abundance of E. lacustris. However, epibiont ciliates may be a threat for this species and may play a substantial role in determining the production, distribution, and community dynamics of E. lacustris.     

Diversity of Prokaryotes in Planktonic Communities of Saline Sol-Iletsk lakes (Orenburg Oblast,Russia)

Prokaryotic diversity was studied in the planktonic communities of six Sol-Iletsk lakes (Orenburg oblast, Russia) varying in salinity level using the Illumina technology of high-throughput sequencing. The extremely halophilic archaea of the phyla Euryarchaeota and Nanohaloarchaeota, as well as the bacterial phylum Bacteroidetes predominated in the communities of lakes with salinity of 285–300‰. Representatives of the phyla Bacteroidetes and Actinobacteria, as well as of the class Gammaproteobacteria were predominant in the lakes with salinity 110?180‰. A bloom of Cyanobacteria was observed in Bol’shoe Gorodskoe Lake (10‰ salinity). The dominant OTUs in the lakes with high salinity were represented by archaea Halonotius sp., uncultured Nanohaloarchaea, and bacteria Salinibacter sp. In the lakes with medium salinity level the dominants included gammaproteobacteria Spiribacter sp., alphaproteobacteria Roseovarius sp., flavobacteria Psychroflexus sp., unidentified archaea of the family Haloferacaceae, actinobacteria Pontimonas sp. and Rhodoluna sp. In the lake with low salinity level cyanobacteria of the genus Planktothrix were predominant. Effect of salinity on prokaryotic taxonomic richness, composition, and diversity in planktonic communities of the studied lakes was demonstrated.     

On the diversity of fishes in mountain lakes of the Amur basin

Results of the study of 11 lakes situated in the mountainous part of the Amur River basin are reported. Six fish species, namely, blunt-snouted lenok Brachymystax tumensis, Baikal-Lena grayling Thymallus baicalolenensis, Siberian stone loach Barbatula toni, Lagowski’s minnow Rhynchocypris lagowskii, Czekanowski’s minnow Rh. czekanowskii, and Amur sculpin Cottus szanaga, were found in five lakes. Four species were found in one lake (Lake Pereval’noe, basin of the Amgun’ River), and two species were found in each of the remaining lakes. Lenok and Amur sculpin were the most common species. The fauna of Lake Okonon (basin of the river Zeya) was the most specific and did not have analogs among the lake faunas; Czekanowski’s minnow and Baikal black grayling were found in this lake. The lenok ecotype that formed in the mountain lakes differed from the river lenok in some regards. The potential pathways of formation and protection of mountain lake ichthyocenoses of the Amur basin are discussed.     

Structure of Microbial Communities of the Sediments of Alkaline Transbaikalia Lakes with Different Salinity

The goal of the work was to reveal the differences in the structure of microbial communities of Transbaikalia alkaline lakes stemming from the differences in their salinity and hydrochemical parameters. The lakes studied were Verkhnee Beloe (Buryat Republic, Russia), as well as Khilganta, Gorbunka, and Borzinskoe (Transbaikal krai, Russia) with salinity from 12.3 to 430 g/L, which differed in the mineral composition of the sediments and hydrochemical parameters. Lake sediments were found to contain 47 prokaryotic phyla (42 bacterial and 5 archaeal ones). The phyla Proteobacteria, Euryarchaeota, Bacteroides, Chloroflexi, Actinobacteria, and Firmicutes were predominant, comprising over 95% of the classified sequences. Comparative abundance of archaea increased with salinity from below 1% in Lake Verkhnee Beloe to 35% in Lake Borzinskoe. The most numerous bacterial OTUs belonged to gammaproteobacteria of the genus Halomonas (up to 15% of the number of classified sequences). The most numerous archaeal OTUs were identified at the genus level as members of the genera Halorubrum and Halohasta belonging to the family Halorubraceae, which comprises extremely halophilic Euryarchaeota.     

Abundance,distribution, and territory areas of rock-dwelling Lake Tanganyika cichlid fish species

Lake Tanganyika, the second-oldest and second-deepest lake in the world, harbors an impressive cichlid fish fauna counting about 250 endemic species that are characterized by a great level of ecological, morphological, and behavioral specialization. This study describes and compares cichlid fish communities at two rocky shores with differential human impact in the south of Lake Tanganyika. Species inventories and depth-dependent abundances were elaborated. About 41 and 46 sympatric cichlid species were recorded in the two study sites, respectively. Variabilichromis moorii was the most abundant species (29–60% of total number of fishes), followed by Aulonocranus dewindti (3–19%), Tropheus moorii (12%), Ophthalmotilapia ventralis (4–10%), Eretmodus cyanostictus (6–11%), and Cyathopharynx furcifer (0.01–9%). All other species had abundances below 5%. It further emerged that large cichlids such as Petrochromis species, Cyathopharynx furcifer, and Lobochilotes labiatus were very rare at one location, with frequencies of 0.55% or less. Territorial sizes of three particularly abundant species, Variabilichromis moorii, Aulonocranus dewindti, and Tropheus moorii, were assessed by behavioral observations. We distinguished between territorial core areas and total defended area, yielding average core areas between 0.4 (V. moorii) and 1.6 m² (T. moorii), and total defended areas averaging for each species between 1.6 (V. moorii) and 5.0 m² (A. dewindti) with no significant differences between the two study sites. The data on individual densities are also relevant for evolutionary studies, in that they allow more accurate calculations of effective population sizes.     

Restoration of zooplankton in a small acidified mountain lake (Plešné Lake,Bohemian Forest) by reintroduction of key species

Atmospheric acidification brought about the extinction of several species of planktonic Crustacea in lakes of the Bohemian Forest (?umava, Böhmerwald). The recent reversal in chemical parameters of lake water towards pre-acidification values raises the question whether the return of missing species is possible. We have designed a bioassay to test water from different lakes for the survival of Daphnia longispina, which inhabited the lakes under study but disappeared from those most affected by acidification. The bioassay showed the best survival in water from Ple?né Lake, one of those where Daphnia and Cyclops became extinct. A large-scale experiment was conducted by transplanting Daphnia longispina (Crustacea, Cladocera) and Cyclops abyssorum (Crustacea, Copepoda) from Prá?ilské Lake, where they have survived until the present, to Ple?né Lake. One year later, a successful reintroduction was confirmed for Cyclops abyssorum.     

<Emphasis Type="Italic">Diaphanosoma mongolianum</Emphasis> Ueno, 1938 (Cladocera: Sididae) in Lakes of Yaroslavl Oblast (Russia)

Findings of the cladoceran Diaphanosoma mongolianum have been recorded in lakes Pleshcheevo (since 2004) and Nero (since 2007) in Yaroslavl oblast at the northern boundary of its range. Similar species D. brachyurum, previously described in both lakes, is found in insignificant numbers. In 2004–2015, the seasonal abundance, spatial distribution, and diel vertical migration of D. mongolianum were studied in deep mesotrophic Lake Pleshcheevo. The species was found in the plankton from early June until October, with the maximum abundance in July–August. The late appearance of males and gamogenetic females (September) is characteristic of its population. D. mongolianum was the most abundant in the epilimnion of the pelagic zone, where it performed diel vertical migrations, concentrating in the surface layer (0–2 m) at night and at a depth of 2–6 m during the day.