Relationships between stomach contents and vertical migration in Meganyctiphanes norvegica, Thysanoessa raschii and T. inermis (Crustacea Euphausiacea)

Stomach contents of Thysanoëssa raschii, T. inermis andMeganyctiphanes norvegica collected in the Gulf of St. Lawrenceshowed the Thysanoëssa species fed primarily during thenight in the upper 75 m of water. M. norvegica fed on copepodswhile at their daytime depth of about 125 m and on phytoplanktonand copepods during the night above 75 m depth. M. norvegicafed on a wider range of organisms than the other two speciesduring the three sample periods of spring, fall and early winter.There was no difference in the minimum size of the cells ingestedbetween any of the species. No evidence was found of any ofthe species feeding on bottom deposits during the sample periods.     

Effects of temperature on metabolism, growth and growth efficiency of Thysanoessa longipes (Crustacea: Euphausiacea) in the Japan Sea

Growth (assessed from intermolt period and molt increment) andmetabolism (oxygen consumption) of juvenile and adult Thysanoessalongipes from central Japan Sea were determined at eight differenttemperatures ranging from 0 to 14°C. The intermolt periodshortened progressively as temperature increased from 0 to 14°C.The molt increment was not assessed satisfactorily in the rearingexperiments, and therefore this was estimated from the naturalgrowth curve and habitat temperature, combined with laboratory-obtainedintermolt data. Oxygen consumption rates increased exponentiallyfrom 0 to 8°C and then leveled off. From these results,the growth was expressed as a function of temperature and bodysize and metabolism as a function of temperature. Because ofthe differential effects of temperature on growth and metabolism,the net growth efficiency [NGE: 100 x growth/(growth + metabolism)]changed with temperature. The temperature at which T. longipesattained maximum NGE varied from 0 to 8°C, depending onthe body length of specimens of 5, 10, 15 and 20 mm. The presentresults are compared with our previous data on Euphausia pacificain relation to the body composition and habitat usage of thesetwo trophically important species in the food web of the JapanSea.     

Vertical distribution, population structure and life history of Thysanoessa longipes (Crustacea: Euphausiacea) around Yamato Rise, central Japan Sea

The diel vertical migration, growth and spawning season of theeuphausiid, Thysanoessa longipes, were investigated using seasonalsamples collected from waters around the Yamato Rise, centralJapan Sea, during the period 1987 to 1999. Thysanoessa longipeswas present throughout a broad bathymetric layer extending downas deep as 1000 m. There was a clear trend for larger specimensto occur at deeper depths. The peak of abundance of the totalpopulation occurred at depths of 30–300 m at night, and150–500 m during the day, and the distance of the dielvertical migrations of the total population was estimated tobe between 100 and 150 m. Population structure analysis revealedthe occurrence of three cohorts aged 0+, 1+ and 2+ years, withfemales attaining a larger body size than males. Growth as determinedby body length was found to fit well to the von Bertalanffygrowth equation. The estimated life span for males and femalesis 3 years, and females reach maturity in 2 years. Based onthe occurrence of calyptopis larvae, spawning of T. longipeswas estimated to occur over only a limited period of the yearbetween April and May.     

Die Calyptopis- und Furcilia-Stadien vonEuphausia hanseni (Crustacea: Euphausiacea)

The larval development is discussed and calyptopis stages I–III as well as furcilia stages I–VII are described. The larval development is similar to that ofE. spinifera, E. longirostris andE. triacantha (group d of genusEuphausia). Antennule, antenna, mandible, maxillule, maxilla and spines on the telson are unsuitable for identifying the larvae of different species of the genusEuphausia. The following characteristics can be used to distinguish the different larvae: form and marginal spines of the carapace, form of the rostrum, spines on the segments of the abdomen, and number and form of the pleopods. The mandible of larval Euphausiacea produces an asymmetric lacinia mobilis and a pair of processus incisivus accessorius (spine row). These two appendages are lacking in adult Eucarida.

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Mitglied der Taxonomischen Arbeitsgruppe an der Biologischen Anstalt Helgoland     

Spatial distribution of Euphausia pacifica (Euphausiacea: Crustacea) in the Yellow Sea

Euphausia pacifica were collected in the Yellow Sea in summer(August, 1997) and winter (February, 1998), and their distributionwas investigated in terms of the developmental stages in relationto environmental factors (temperature, salinity and chlorophylla). In summer, the water column was highly stratified with athermocline between 10–30 m depth, whereas the water columnwas well mixed in winter. Seasonal variation in temperaturewas large, ranging between 6.3 and 28.8°C in summer and3.9 and 12.2°C in winter. Chlorophyll a concentration changedlittle seasonally but a high concentration was observed in coastalareas. Salinity in the two seasons varied little spatio-temporally.In the studied area and sampling period, E.pacifica was themost dominant euphausiid species (99.7 and 99.8% in summer andwinter, respectively), and comprised adults and juveniles (74.6%in summer and 41.9% in winter), furcilia (17.5 and 44.9%) andcalyptopis (7.9 and 13.1%). The spatial distribution of theE.pacifica population varied seasonally depending on developmentalstage and appeared to be related primarily to the seawater temperatureand secondly to the chlorophyll a concentration, but not tosalinity. In summer, adults were abundant in areas below 10°C,and furcilia and calyptopis above 9°C. In winter, adultswere confined to areas between 7 and 10°C, and furciliaand calyptopis, to areas between 6 and 8°C. Furcilia andcalyptopis were concentrated in the vicinity, or at the centerof chlorophyll a-rich water masses, whereas adults seemed toavoid those water masses.     

Ommochrome pigments in the eyes of Euphausia superba (Crustacea,Euphausiacea)

Summary Ommin and a xanthommatin-like ommochrome pigment have been extracted from the eyes of Euphausia superba, and identified by their absorbance spectra and redox reactions invarious solvents. The absorbance spectrum of ommin is relatively stable, but that of the xanthommatin pigment is sensitive to light, pH, and hydroxylamine. Both pigments are readily soluble in 2% digitonin. They are therefore likely contaminants of visual pigment extracts, and may consequently introduce artifacts into visual pigment characterizations. Microspectrophotometric measurements show that both pigments are components of the distal and proximal screening pigment granules in E. superba ommatidia.This paper is dedicated to the memory of Professor Mary Alice McWhinnie, to whom I owe a great debt of gratitude for her patient tutelage and the opportunity to participate in her Antarctic research, for the encouragement to pursue my own interests, and for her generous friendship. It was a cherished association which ended sadly with her untimely death in March, 1980. She will always be fondly remembered     

DNA barcodes for species identification of euphausiids (Euphausiacea, Crustacea)

Many species of euphausiids (Euphausiacea, Crustacea) are distinguishedby subtle or geographically variable morphological characters,and erroneous identification of euphausiid species may be morefrequent than currently acknowledged. DNA barcodes (short DNAsequences that discriminate species and aid in recognition ofunknown species) are of use for this group. A 650 bp regionof mitochondrial cytochrome oxidase I (mtCOI) was sequencedfor 40 species of 10 euphausiid genera: Bentheuphausia, Euphausia,Meganyctiphanes, Nematobrachion, Nematoscelis, Nyctiphanes,Stylocheiron, Tessarabrachion, Thyssanoessa and Thysanopoda.mtCOI sequence variation discriminated all species; pairwisedifferences averaged 16.4% (range 7–24%); mean generalizedtime reversible (GTR) genetic distance was 26.7%. mtCOI reliablyidentified euphausiid species: variation within species wastypically < 1% and GTR distance was typically < 2%. Atlanticand Pacific Ocean populations of Euphausia brevis differed by13% (GTR genetic distance = 28%) and may deserve status as distinctspecies. mtCOI gene trees were reconstructed for five generausing maximum parsimony, maximum likelihood and Bayesian algorithms;best-fit models of nucleotide evolution were determined foreach genus. The mtCOI gene tree for 20 species of Euphausiareproduced one of three morphologically defined species groups.mtCOI resolved relationships among closely related species ofmost genera, usually in accord with morphological groupings.A comprehensive DNA barcode database for euphausiids will helpensure accurate species identification, recognition of crypticspecies and evaluation of taxonomically meaningful geographicvariation.     

Feeding behaviour in Meganyctiphanes norvegica (M. Sars) (Crustacea: Euphausiacea)

Meganyctiphanes norvegica (M. Sars) will feed upon the centric diatom Thalassiosira weissflogii (Grunow) G. Fryxell & Hasle but cannot fulfil its energy requirement for metabolism on this food. Its daily metabolic requirement can be exceeded when the euphausiid feeds upon the copepods Calanus finmarchicus Gunner or Centropages typicus Krøeyer, but not when feeding upon the smaller copepods Pseudocalanus spp. or Acartia spp. When feeding upon a natural copepod assemblage Meganyctiphanes norvegica requires high concentrations of copepods to achieve its metabolic requirements, suggesting that the euphausiid may exploit vertically patchy concentrations of prey. Short-term predation rates on Pseudocalanus spp. were also used to estimate feeding rates. Feeding in Meganyctiphanes norvegica appears to be adapted to a spatially variable food supply and rapid exploitation of food sources concentrated into patches or layers. The filter area of the feeding basket of M. norvegica is proportionally smaller than the filter area of Euphausia superba Dana, but has the same allometric length exponent. The filter area probably reflects the difference between the primarily carnivorous diet of Meganyctiphanes norvegica and herbivorous diet of Euphausia superba.     

The organization of the olfactory lobes in Euphausiacea and Mysidacea (Crustacea,Malacostraca)

Summary The structural organization of the olfactory lobes in representatives of euphausiid and mysid crustaceans was investigated and compared, also with these structures described in other crustaceans and in insects. In the investigated euphasiid and mysid species, the olfactory-globular tract and the position of cell clusters associated with the olfactory lobes show a similar arrangement. This arrangement is in agreement with that described in decapod crustaceans. The olfactory lobe neuropil in representatives of both taxa shows glomerular arrangement. These glomeruli are partly enclosed in a glial wrapping, and they represent the only site where synaptic contacts are established within the olfactory lobes. This glomerular arrangement appears similar to that described in the antennal lobe of insects, but differs from the columnar arrangement described in decapod crustaceans. Furthermore, about 15–20 FMRFamide-like immunoreactive globuli cells were labeled and they occupy a similar position in the investigated species. Neurites from these cells terminate only in the olfactory lobe glomeruli, and they are therefore regarded as intrinsic interneurons. The absence of serotonin-like immunoreactivity in the olfactory lobes is a feature only ascribed to the euphausiids and mysids. A specific neuropil area is present in male mysids, and it occupy a position forward of the olfactory lobe. The male-specific neuropil in mysids and the macro-glomerular in insects complex are interpreted as analogous structures.     

Morphology of Urospora chiridotae (Sporozoa: Gregarinomorpha: Eugregarinida) from sea cucumber Chiridota laevis (Echinodermata: Holothuroidea: Apoda)

Several morphological forms (morphotypes) of Urospora chiridotae gamontes are found in White Sea holothuroid Chiridota laevis. All these morphotypes are differed by localization in the body of host, form and cytological features. The gregarines are situated in several host biotopes, such as blood vessels, intestine and mesenteries. In the blood vessels elongate skittle-like cells supplied with long thin cytopillia are observed. On the external surface of the intestine spherical gregarines are found. These parasites commonly covered with one layer of coelomic epithelium's cells. In some holothuria intratissue spherical cells of parasites located in intestinal epithelium are presented. Both of these types of parasites lack cytopillia, and folds or ridges on its surface. On different mesenteries, connections between intestine and body wall, and also on intestine elongate ounce-shaped cells and gamontocysts are observed. These cells are situated on the apices of finger-like processes of the intestine and mesenteries surface. Ounce-shaped gregarines have cytopillia shorter than in skittle-like gregarines. The differences between morphotypes of Urospora chiridotae are probably caused by different environmental conditions. In the narrow rift of blood vessel elongate cells are developed. The cytopillia may serve for making more or less wide space around gregarines, which is necessary for food uptake. Spherical cells surrounded by host's cells and have the form typical for tissue parasites. In the wide coelomic cavity where convection of liquid proceeds better than in blood vessel, ounce-shaped gregarines with short cytopillia are developed. We found only typical for Urospora chiridotae ovoid oocysts with dissimilar ends, anterior collar and spine-like posterior end. Thus, the all above-mentioned morphotypes undoubtedly belong to the same species. The relationships between defense host cells and the different morphotypes of trophozoites are variable.     

The morphological variability of four species of the euphausiid genus Thysanoessa Brandt, 1851 from Kamchatkan waters

The morphological variability of four species of the euphausiid genus Thysanoessa from Kamchatka waters was studied. The neglecta form and 18 variants of the abdomen structure were found in Th. inermis. Th. raschii and Th. inermis exhibited four variants of eye shape; Th. longipes had three variants. In Th. longipes and Th. raschii, a tooth on the lateral edge of the carapace can be positioned closer to the middle. The most variable morphological features in Th. raschii were the location of a tooth on the carapace; in Th. longipes, the length of the second pair of thoracic legs and the position of a tooth on the carapace; in Th. inspinata, the length of the second pair of legs; in Th. inermis, the eye shape and the structure of the abdomen. A new key for the identification of euphausiids of the genus Thysanoessa that inhabit the Russian Far Eastern seas is proposed.     

Behavioural differences in relation to pycnoclines during vertical migration of the euphausiids Meganyctiphanes norvegica (M. Sars) and Thysanoessa raschii (M. Sars)

An observation on differences in the vertical migration patternbetween Meganyctiphanes norvegica (M. Sars) and Thysanoessaraschii (M. Sars) caused by the presence of a thermocline isreported. Generally, M.norvegica did not migrate through a thermoclineat 50–60 m depth while T.raschii did. This observationis discussed in the context of how the combination of differencesin vertical migration and advection over sill depth may affectthe abundance of fjord stocks of these two euphausiid species.     

Egg Size and Formation of the Geographical Range in <Emphasis Type="Italic">Thysanoessa inermis</Emphasis> (Kroyer, 1846) (Crustacea: Euphausiacea) in the Norwegian and Barents Seas

Using materials on the euphausiacean Thysanoessa inermis collected in the Norwegian and Barents seas (from 60°N to 73°N) we studied the variability of size characteristics in the eggs (diameter of embryos and egg capsules and the width of the perivitelline space) and their correlations with water temperature and salinity. The size of embryos showed almost no variability irrespective of the location of the sampling site and the water temperature and salinity; the perivitelline space performs a protective function, and its width showed a tendency to decrease with increasing water temperature and salinity. The similarities in the size of embryos in T. inermis populations thousands of kilometers away from each other might be explained, first, by the relatively small age of the populations (the age of the population from the Barents Sea did not exceed 6000–7000 years) and second, by movement of the crustaceans with currents from the Norwegian Sea to the Barents Sea.     

The mitochondrial genomes of Euphausia pacifica and Thysanoessa raschii sequenced using 454 next-generation sequencing, with a phylogenetic analysis of their position in the Malacostracan family tree

Euphausiid krill play a critical role in coastal and oceanic food webs, linking primary producers to upper trophic levels. In addition, some species support commercial fisheries worldwide. Despite their ecological importance, the genetics of these important species remain poorly described. To improve our understanding of the genetics of these ecological links, we sequenced the mitochondrial genomes of two species of North Pacific krill, Euphausia pacifica and Thysanoessa raschii, using long-range PCR and 454 GS Junior next-generation sequencing technology. The E. pacifica mitogenome (14,692 + base pairs (bp)) encodes 13 protein-coding genes (PCGs), two ribosomal RNA (rRNA) genes, and at least 22 transfer RNA (tRNA) genes. The T. raschii mitogenome (14,240 + bp) encodes 13 PCGs, two rRNA genes, and at least 19 tRNA genes. The gene order in both species is similar to that of E. superba. Comparisons between Bering Sea and Yellow Sea E. pacifica revealed a total of 644 variable sites. The most variable protein-coding gene were atp8 (7.55 %, 12 of 159 sites variable), nad4 (6.35 %, 85 variable sites) and nad6 (6.32 %, 33 variable sites). Phylogenetic analyses to assess the phylogenetic position of the Euphausiacea, using the concatenated nucleic acid sequences of E. pacifica and T. raschii along with 46 previously published malacostracan mitogenomes, support the monophyly of the order Decapoda and indicate that the Euphausiacea share a common ancestor with the Decapoda. Future research should utilize this sequence data to explore the population genetics and molecular ecology of these species.     

Phylogenetic analysis of the Malacostraca (Crustacea)

The Malacostraca comprises about 28 000 species with a broad disparity in morphology, anatomy, embryology, behaviour and ecology. The phylogenetic relationships of the major taxa are still under debate. Is the Leptostraca the sister group of the remaining Malacostraca, or is this taxon more closely related to other Crustacea? Does the Stomatopoda or the Bathynellacea represent the most basal taxon within the remaining taxa? Is the Peracarida monophyletic or are some peracarid taxa more closely related to other ‘caridoid’ taxa? Is the Thermosbaenacea part of the Peracarida or its sister group, and how much support is there for a taxon Amphipoda + Isopoda? To answer these questions a phylogenetic analysis of the Malacostraca combining different phylogenetic approaches was undertaken. In a first step, the monophyly of the Malacostraca including the Leptostraca is shown using the ‘Hennigian approach’. A computer cladistic analysis of the Malacostraca was carried out with NONA and PEE ‐WEE , based on 93 characters from morphology, anatomy and embryology. Nineteen higher malacostracan taxa are included in our analysis. Taxa whose representatives are exclusively fossils were not included. The Leptostraca was used as an operational out‐group. The present analysis supports the basal position of the Stomatopoda. Syncarida and Peracarida (including Thermosbaenacea) are supported as monophyletic, the Eucarida is not. Instead a sister‐group relationship is suggested between Euphausiacea and Peracarida (including Thermosbaenacea), with the Syncarida as the sister group to both taxa. Certain embryonic characters are interpreted as support for the monophyly of the Peracarida (without Thermosbaenacea) because convergences or reversals of these characters seem implausible. Within the Peracarida, the Mysidacea (Lophogastrida + Mysida) represents the sister group to the remaining taxa. A sister‐group relationship between Amphipoda and Isopoda is not supported.     

The effect of lunar eclipse on the vertical migration behaviour of Meganyctiphanes norvegica (Crustacea: Euphausiacea) in the Ligurian Sea

The vertical migration of a zooplankton community dominated by the euphausiid Meganyctiphanes norvegica was monitored between 16 and 23 September 1997 with a 153 kHz Acoustic Doppler Current Profiler (ADCP) and a MOCNESS net. The sampling period covered a phase in the lunar cycle when the rise of the moon (full moon) coincided initially with sunset and then became progressively later. On 16 September 1997, a lunar eclipse occurred 45 min after sunset, lasting for 2 h. At dusk, the ADCP observed the upward vertical migration of two principal backscattering bands 10 min apart with vertical velocities of up to 7 cm s^-1. After a period at the surface, a more diffuse band subsequently sank at a slower rate (1-2 cm s^-1) to a depth of 75-100 m. Net samples showed that the earlier band consisted mainly of the pteropod Cavolinia inflexa, whilst the later band was mostly euphausiids, predominantly M.norvegica. This species was also the major constituent of the band that sank. The timing of upward migration was relatively constant over the sampling period, but there was an increasing delay of the secondary sinking until 21 September. This showed as a strong correlation between the onset of sinking and the time of moonrise. The lunar eclipse on 16 September perturbed this pattern, such that animals did not sink soon after their arrival at the surface, as occurred on 17 September but remained at the surface until the end of the umbra. This suggests that M.norvegica can perceive moonlight and that this influences vertical migration. Evidence that the behaviour is not solely mediated by this erogenous factor, however, is seen in the pattern that emerged after 21 September, when midnight sinking occurred at a relatively constant time after sunset and before moonrise. These observations support the hypothesis that moonlight is a Zeitgeber for an endogenous rhythm that synchronizes secondary sinking behaviour with the lunar cycle.      

Vertical distribution and feeding pattern of Euphausiacea (Crustacea) in the eastern Banda Sea (Indonesia) during the SE and NW monsoons

The vertical distribution of Euphausiacea (Crustacea) in theeastern Banda Sea (Indonesia) during the SE and NW monsoon seasonswas studied. In August 1984 and February/ March 1985, stratifiedday and night sampling was carried out between 0 and 500 m atfour stations. A total of 31 species was found, of which mostwere present in both monsoon seasons. Species with a high presencein samples and performing diurnal vertical migration in bothmonsoon seasons were Euphausia diomedeae, Euphausia pseudogibba,Thysanopoda monacantha, Thysanopoda iricus-pidaia and Nematoscelismicrops. Species with a high presence in samples showing nodiurnal vertical migration in both monsoon seasons were Euphausiasimilis, Nematobrachion boopis. Nematoscelis tenella, Stylocheironmaximum and Thysanopoda orientalis. The density of Euphausiaceaat the sampled stations was higher, but more heterogeneous,in the nutrient-enriched SE monsoon period than in the relativelynutrient-poor NW monsoon season, when densities were lower andmore similar. The vertical distributions and diurnal verticalmigration of species did not show a unanimous and strong responseto seasonal hydrographic differences. The day–night depthdistribution pattern of the Euphausiacea population in the upper500 m is largely the same in both seasons, with comparable migrationranges. Some species showed a somewhat upward shifted depthdistribution in the nutrient-enriched period. For mesopelagicand bathypelagic species especially, food resources were thenfavourable in the upper layers, as indicated by rare recordsof Thysanopoda crisiata. Analyses of stomach contents snowedmainly a nocturnal feeding pattern for the diurnal verticalmigrating species and a continuous feeding pattern for the non-migratingspecies.