Functional feeding morphology of the euphausiid Nyctiphanes australis

The structure of the feeding basket, mandible and stomach armature of the krill Nyctiphanes australis (G. O. Sars, 1883) was examined by scanning electron microscope with the aim of predicting its diet. N. australis were collected during February 1986 and October 1986 to February 1987 in Otago Harbour, New Zealand (45° 50 S; 170° 37 E). Predictions based on the functional morphology were tested by examining stomach contents with SEM. Intersetule distances of the feeding basket (1 to 7.5 m) are finer than in other krill species, suggesting that N. australis can efficiently collect nanoplankton-sized particles (2 to 20 m). The mandibular edge index (0.74) matched the edge index of Meganyctiphanes norvegica. This indicates, in contrast to the fine feeding-basket setulation, that N. australis has a mandible resembling that of predominantly carnivorous krill species. The ratio of mandibular palp length to mandible width is 3.2±0.2, or relatively longer than the elongate palp thought to be indicative of herbivorous habit in Euphausia superba. The fine structure of the molar shows specialized surfaces which differ from those of other krill species. The internal armature of the stomach is heavily spinose, as is common in krill of herbivorous habit. Layers of various-shaped spines at differing densities were observed. The functional morphology suggests that N. australis is an opportunistic omnivore. The stomach contents tentatively support this prediction, containing fragments of phytoplankton and detrital material. However, stomach contents were generally amorphous, making identification of dietary components in the field difficult.     

Stranding and matting behaviour in Nyctiphanes australis (Euphausiidae: Crustacea)

Numerous stranding events of the euphausiid Nyctiphanes australis Sars on the coast of south-east Tasmania during the winter months of 1984/1985 are documented. The krill were at a reproductive stage of their life cycle. Strandings almost always occurred on calm, sunny days on beaches with a northerly aspect. Typically, barometric pressure was greater than and wind speed was less than the monthly average. No significant differences were found between stranded euphausiids and those from surface swarms, and the relationship between the two forms of behaviour in this area is discussed. A new form of behaviour termed matting was observed, in which the euphausiids aggregate dorsal-side down on the substrate in shallow water. Matting usually occurs synchronously with stranding. Laboratory experiments showed that N. australis responds to changes in both the plane of polarization and intensity of light with behaviour typical of that observed in stranding and matting events. Light appears to be a key factor mediating these types of behaviour. Hypotheses for the relationship between stranding and matting are offered.     

Seasonal and interannual variability in krill (Nyctiphanes australis) stocks and their relationship to the fishery for jack mackerel (Trachurus declivis) off eastern Tasmania,Australia

Nyctiphanes australis was collected from the east coast of Tasmania between January 1989 and April 1991. Density and biomass were significantly higher in autumn than in any other season. The population was dominated by juveniles, except in autumn and spring 1990 when there was a significant increase in the proportion of adults. Our data indicated that N. australis does not regularly migrate vertically and that it forms aggregations of particular size classes which vary both temporally and spatially. Stomach fullness in Trachurus declivis, a major predator of N. australis, rose to a peak in autumn when N. australis stocks and the monthly catches by the fishery for T. declivis were at their highest. The stomachs of T. declivis were also dominated by adult size classes during this period. The virtual absence of N. australis in 1989 and the subsequent failure of the T. declivis fishery in that year underline the interrelationship between these two species. We suggest that this was the result of an influx of subtropical northern waters low in nutrients onto the shelf, which corresponded with a major La Niña cold event at that time.     

Contribution of moulting and eggs to secondary production in Nyctiphanes australis (Crustacea: Euphausiacea)

This study was designed to assess the contribution of moulting and eggs to production of the euphausiid Nyctiphanes australis G. O. Sars. For this purpose, live specimens were collected from Storm Bay, south-eastern Tasmania, between August and December 1981, while preserved samples, collected between February 1980 and February 1981, were also examined. The intermoult period of N. australis increased exponentially with increase in body length and weight. Larvae moulted approximately every 2.5 to 3 d and adults every 4 to 5 d at 15°C. A decrease in temperature from 15° to 10°C resulted in the intermoult period almost doubling. The mean weight of exuviae produced represented nearly 6% of the body dry weight of the individual. Continuous maturation of ova was observed, with an individual female capable of releasing a total of 1 100 eggs in a lifetime. These are deposited as a series of batches into a pair of external ovisacs every 30 d. The size of the batch of eggs was dependent on the size of the female. Moulting of euphausiids forms a significant contribution of organic matter to detrital food webs. A value of 42.01 mg m^-3 yr^-1 was obtained for the production of exuviae. Egg production was calculated to be between 1.41 and 4.22 mg m^-3 yr^-1. Production and mean annual biomass of N. australis was previously calculated as 78.29 mg m^-3 yr^-1 and 5.39 mg m^-3, respectively. Thus, the total production integrated for the whole of Storm Bay was 125 mg m^-3 yr^-1 or 2 309 tonnes dry wt yr^-1, representing an overall P:B ratio of 23.1.     

Biochemical composition of Nyctiphanes australis and its possible use as an aquaculture feed source: Lipids,pigments and fluoride content

Nyctiphanes australis contained, on a dry weight basis, an average of 52% crude protein and 5.0 to 9.5% lipid. The fatty acid profile of N. australis was markedly unsaturated, with a mean total 3 fatty acid content of 48.6±2.4% of total fatty acids. N. australis contained high levels of the essential long-chain polyunsaturated fatty acids eicosapentaenoic (EPA, 20:53) and docosahexaencic (DHA, 22:63), ranging from 16.6 to 36.5% and 11.1 to 24.8%, respectively. The concentration of total carotenoids ranged from 137 to 302 g g^–1 dry wt, with no significant differences in concentrations found with season or life stage. The carotenoids were comprised of 79.5% astaxanthin and 20.5% canthaxanthin. The lipid and pigment compositions of N. australis suggest that the species could serve as a suitable feed source for cultured salmonids. Like other euphausiids, N. australis contained high levels of fluoride, with a seasonal range between 277 and 3507 g g^–1 dry wt. The high fluoride levels found in N. australis would not detract from its potential as a feed source for salmonids because ingested fluoride is largely absorbed by the skeleton.     

桑沟湾大型底栖动物的次级生产力

根据2009年夏季和冬季在桑沟湾9个大面站进行的大型底栖动物定量采集样品资料,利用Brey(1990)的经验公式对调查海区进行了大型底栖动物丰度、生物量、次级生产力和P/B值的研究计算.结果表明,该海域大型底栖动物年平均丰度为1 226.7 ind m-2,年平均生物量为3.62 g(AFDW)m-2,年平均生产力为4.76 g(AFDW)m-2 a-1,P/B值平均为1.45 a-1.调查期间由湾口到湾底,次级生产力呈明显的梯度升高.相关分析表明,水深和水温是影响次级生产力的重要环境因子.通过比较,发现桑沟湾大型底栖动物次级生产力与南黄海相近,高于东海、长江口和深沪湾,低于渤海、胶州湾、大亚湾和海坛海峡;P/B值低于长江口、海坛海峡和深沪湾,高于其它海区,说明桑沟湾大型底栖动物群落中,个体小、生活史短、代谢快的种类所占比例低于海坛海峡和深沪湾,高于其它海区.图2表3参25     

Distribution of euphausiid assemblages in the Mediterranean Sea

Eleven species of euphausiids from 24 Isaac-Kidd Midwater Trawl (IKMT) night collections taken at stations throughout the Mediterranean Sea were counted. The frequency of occurrence and dominance of individual species and percent similarity faunal analysis of the euphausiid community were used to describe changes in faunal composition between geographical areas and differences in vertical distribution. Although most species were widespread, three distinct patterns of abundance were apparent: Euphausia krohnii, Nematoscelis megalops, Meganyctiphanes norvegica, and Stylocheiron abbreviatum predominated in western basin areas west of the Tyrrhenian Sea; Euphausia hemigibba, Thysanopoda aequalis, and Stylocheiron longicorne predominated in the Tyrrhenian Sea and eastern Mediterranean Sea; Euphausia brevis and Stylocheiron suhmii predominated in the eastern Mediterranean Sea. Percent similarity analysis of data from the IKMT samples and data from Ruud (1936) indicates the Tyrrhenian Sea fauna at the time of the collections was more similar to eastern Mediterranean areas than to most other areas in the western basin, although the degree of similarity was dependent, to some extent, on the depth at which the samples were collected. The composition of euphausiids living above 150 m at night in this area was more similar to eastern basin areas, while the composition of deeper living forms was more similar to those of the rest of the western basin. Comparison of euphausiids collected at three points over a 60 year time-span in the Balearic Sea shows the similarity in composition to be greater within the area over time than between adjacent areas in the western Mediterranean Sea.Contribution No. 2732 from the Woods Hole Oceanographic Institution, Woods Hole, Massachusetts, USA. This research was supported by the Atomic Energy Commission Contract AT (30-1)-3862, ref. NYO-3862-49, National Science Foundation Grant GA 29303 and Office of Naval Research Contract N00014-71-C-0284.     

Molecular dating and biogeography of the neritic krill Nyctiphanes

The genus Nyctiphanes (Malacostraca, Euphausiacea) comprises four neritic species that display antitropical geographic distribution in the Pacific (N. simplex and N. australis) and Atlantic (N. couchii and N. capensis) Oceans. We studied the origin of this distribution applying methods for phylogenetic reconstruction and molecular dating of nodes using a Bayesian MCMC analysis and the DNA sequence information contained in mtDNA 16S rDNA and cytochrome oxidase (COI). We tested hypotheses of vicariance by contrasting the time estimates of cladogenesis with the onset of the major barriers to ocean circulation. It was estimated that Nyctiphanes originated in the Pacific Ocean during the Miocene, with a lower limit of 18 miilion years ago (Mya). An Atlantic–Pacific cladogenic event (95% HPD 3.2–9.6) took place after the closure of the Tethyan Sea, suggesting that dispersal occurred from the Indo-Pacific, most likely via southern Africa. Similarly, the antitropical distribution pattern observed in the eastern Atlantic Ocean likely resulted from recent Pliocene–Pleistocene (95% HPD 1.0–4.97) northward dispersal from the southern hemisphere. Our results imply that dispersal appears to have had a significant role to play in the evolution of this group. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Larval development of the euphausiid Euphausia pacifica in the Yellow Sea

The complete larval development of Euphausia pacifica in the Yellow Sea is described and the stages are compared with larvae of E. nana. Euphausiid larvae examined in the present study were collected at 30 stations in the Yellow Sea in 1989. During the stages of Calyptopis III to Furcilia II, E. pacifica larvae are smaller than E. nana larvae but they are larger from Furcilia III onward. There are geographical variations in body size of Calyptopis III among the southern California waters, the eastern Japan waters, the Japan Sea, and the Yellow Sea; body size is smallest in the Yellow Sea and Japan Sea, while it is largest in southern California waters and intermediate in eastern Japan waters. Segmentation of antennal endopods was observed in the 56 to 51 furcilia forms in the Yellow Sea population, suggesting that this characteristic is not as reliable for identification of furcilia stages as the pleopods and terminal telson spines. Thus, we propose here that there are six furcilia stages of E. pacifica, instead of the seven previously reported by Boden (1950).     

Daytime surface swarms of the euphausiid Thysanoessa inermis off the west coast of Hokkaido,northern Japan

Biological characteristics of the daytime swarming of the euphausiid Thysanoessa inermis (Krøyer) were examined off the west coast of Hokkaido in the northeast Sea of Japan between 1982 and 1987. Swarms were composed exclusively of fully mature individuals; males possessed spermatophores at the ejaculatory ducts and females had attached spermatophores at the thelycum. Females predominated in most swarms, while males or nearly balanced swarms were infrequently encountered. Fully mature females occurred in the late swarming season, usually mid-April. Examination of stomachs indicated that feeding activity was low during the daytime surface swarming behavior. The body sizes of swarming T. inermis occurring around Yagishiri Island were significantly larger than those found simultaneously off the Shakotan Peninsula. This suggests that the euphausiid populations of the two areas are different.     

Respiration and chemical composition of the bathypelagic euphausiid Bentheuphausia amblyops

Bentheuphausia amblyops is a cosmopolitan bathypelagic euphausiid with a vertical range of from 280 to 7 000 m. Determinations of proximate and elemental composition show that B. amblyops has a more robust structure (lower water content and higher protein content) than is typical of other bathypelagic Crustacea. B. amblyops is a strong swimmer and is capable of employing its thoracic legs in raptorial fashion. Discrete depth trawls taken between depths of 400 and 2 500 m on a diel basis show no evidence of vertical migration. There was no significant difference in oxygen consumption within the species environmental temperature range (1.5° to 7.5°C), which may be partially ascribed to a limited temperature effect and partially to variability in spontaneous activity at the different temperatures. Between 7.5° and 9.5°C there was a marked temperature effect on metabolism (Q₁₀=6.4). The observed vertical distribution and metabolic response to elevated temperature preclude the migrations to the surface that have been postulated for B. amblyops. The robust composition of the species, its behavioral characteristics, and published diet information strongly suggest that the species is omnivorous with a strong predatory component in its foraging strategy.     

Investigations on some tropical euphausiid species of the Indian Ocean

In spite of the great ecological importance of euphausiids in large areas of the sea, very little is known about their early developmental stages. During the 40th and 41st cruises of RV Vityaz (USSR) in the western part of the Indian Ocean, new information was obtained on the development of 2 species common in that area: Euphausia diomedeae and Slylocheiron carinatum. In E. diomedeae, the time between fertilization and hatching of the nauplius amounts to about 16 h; the metanauplius stage lasts 2 days, the calyptopis stage I also 2 days, and the calyptopis stage III up to 4 days, at water temperatures ranging from 22° to 26°C. The developmental stages of both species are described and illustrated.     

Morphology and sinking velocities of fecal pellets of copepod, molluscan, euphausiid, and salp taxa in the northeastern tropical Atlantic

Morphological characteristics and sinking velocities of naturally occurring fecal pellets of copepods, euphausiids, salps, and pelagic mollusks collected in the northeastern tropical Atlantic were investigated during the period of May-June 1992. The fecal pellets of copepods and euphausiids were cylindrical and distinguished only by their size. Those of salps were, in general, rectangular, and slight differences were noted according to the species. The fecal pellets of the molluscan pteropod Clio sp. were conical, while those of the molluscan heteropod Carinaria sp. were spiral. The sinking velocities ranged from 26.5 to 159.5 m day^-1 for copepod fecal pellets, from 16.1 to 341.1 m day^-1 for euphausiid pellets, from 43.5 to 1167.6 m day^-1 for salps' pellets (Cyclosalpa affinis, Salpa fusiformis, Iasis zonaria, and two unidentified species), from 65 to 205.7 m day^-1 for Clio sp. pellets, and from 120.3 to 646.4 m day^-1 for Carinaria sp. fecal pellets. The measured sinking velocities were compared with estimates predicted using the equations of Komar et al. (1981; Limnol Oceanogr 26:172-180), Stokes' law, and Newton's second law, using either a constant density of fecal pellets (1.22 g cm^-3) or densities estimated with the three different equations. When a constant density was used, the three equations overestimated the sinking velocities; Stokes' law resulted in the largest overestimation, and Newton's second law, the smallest. At the taxa level, the overestimation was greatest for euphausiid 1 fecal pellets and smallest for copepod fecal pellets. When the three equations were used to estimate fecal pellet density, the density estimated using the equation of Komar et al. was the greatest, and that using Stokes' law, the smallest, resulting in over- and underestimation of sinking velocities, respectively. Newton's second law resulted in an intermediate density and gave the closest estimate of sinking velocities. We propose that measurement of sinking velocities of a portion of the fecal pellets might guide in choosing an appropriate equation to be used for a reasonable interpretation of vertical mass flux.     

Turnover and vertical transport of zinc by the euphausiid Meganyctiphanes norvegica in the Ligurian Sea

Participatory turnover time is defined as the time required to cycle an element in a system through a given material in that system. The participatory turnover time of ionic zinc by the adult Meganyctiphanes norvegica population in the Ligurian Sea ranged between 498 and 1243 years, depending upon the available food supply, and considering the food chain as the only route for zinc accumulation by the population. A total-impact turnover time was calculated as the sum of the participatory turnover time for live individuals plus the time required for dead euphausiids to lose 90% of their zinc to the water. Carcasses lost zinc to the water slower than either feces or molts, and so established the maximum loss time for all particulate excretion products; nevertheless, total-impact turnover time for zinc did not differ significantly from the participatory turnover time. The net vertical transport of zinc by M. norvegica from the sea surface to any specified depth can be calculated as the sum of the dissolved zinc excreted below the depth plus the concentrations of zinc left in feces, molts, and carcasses after they have sunk to the specified depth. Carcasses sink the fastest and lose the smallest fraction of their zinc concentration during descent; fecal pellets sink the slowest and lose the greatest fraction of their zinc concentration, and molts are intermediate. Nevertheless, feces represents the major route for delivering zinc to the bottom of the Ligurian Sea (2500 m), because concentration of the element in the pellets is so much higher than in carcasses or molts. Excretion of dissolved zinc into the water at the vertical migration depth of the living population during daylight hours was also inconsequential. Feces zinc represented over 80% of the total zinc transported to the sea floor if only marginal food supplies were available to the euphausiids, and over 90% if food was in sufficient supply. M. norvegica can effect a net transport of about 98% of its body zinc concentration below 500 m daily, in conditions of sufficient food supply and assuming that no released products are eaten during descent. If the food supply in the Ligurian Sea is considered only marginal throughout the year, M. norvegica can still effect a daily net transport below 500 m of about 36% of its body concentration, and about 6% of its body concentration will reach 2500 m daily.     

Nocturnal changes in the mean length of a euphausiid population: vertical migration,net avoidance,or experimental error?

The lengths of 9 421 Antarctic krill, Euphausia superba Dana, from 144 net samples from 4 depth ranges at the same location (north east of South Georgia) were measured over a period of 6 d. While the mean lengths showed differences between each depth range in each time period of each day, the only consistent difference occurred in the period 00.00 to 04.00 hrs local time, when the mean lengths of krill were up to 2 mm greater than those found at other times. While vertical migration may have contributed to this nocturnal change in mean length, differences in measurement techniques between observers were also implicated. Controlled laboratory experiments confirmed that there were significant differences between measurements of the same krill analysed by six different observers while there were no significant differences in replicate measurements made by a single observer. These results have important implications for future studies involving length and length-frequency analysis, and suitable precautions and experimental protocols are suggested.     

An annual cycle of flash induced luminescence in the euphausiid Thysanoëssa raschii

The euphausiid Thysanoëssa raschii (M. Sars) rarely luminesces spontaneously in the laboratory. The flash-stimulated luminescent response (FSLR) was therefore used to investigate seasonal changes in luminescent ability. Some observations were also made on the responsiveness of Meganyctiphanes norvegica. In the case of T. raschii both sexes showed maximum responsiveness from March through June and in some years to September. The strength of the tendency to respond was strongest in May and June. The FSLR delay was least during April, May and June. These results are interpreted in the light of the hypothesis that the form of the FSLR is the result of opposing inhibitory and excitatory components. It is tentatively concluded that luminescence is involved in behaviour related to spermatophore transference, which takes place from April to June.     

Metal concentration changes in growing Pacific oysters,Crassostrea gigas,cultivated in Tasmania,Australia

Pacific oyster spat (Crassostrea gigas Thunberg) transferred from the Tamar Estuary to two growing areas in southern Tasmania were monitored for their metal contents over one growing season (1974–1975). Oysters at Pipeclay Lagoon were grown with stick and tray culture while those at Dart Island were cultured with the longline technique. Metal content of the oysters increased with time and the trend was similar to the weight growth curves. Mean dry weights of oysters increased from 0.07 to 1.19 g at Pipeclay Lagoon and from 0.25 to 1.47 g at Dart Island. Metal contents (g) increased at each site, respectively: Fe 57 to 326; 91 to 446; Zn 259 to 6 555; 755 to 5 335; Cd 1.5 to 13.3; 1.9 to 16.3; Cu 26 to 142; 9 to 116; Pb 1.9 to 11.9; 0.6 to 3.8. Concentration curves generally showed a downward trend with time. The relationships of metal concentrations with weight did not differ from sample to sample at a site nor did they differ at one site compared with the other. The only exception was lead, which showed no relationship of concentration with weight at Pipeclay Lagoon and a negative one at Dart Island. It is postulated that higher winter concentrations of metals in the oysters were linked with greater solubility of metal ions in lower salinity water.     

Dominance of larval forms in euphausiid (Crustacea:Eucarida) ontogenesis

Variability and dominance of Euphausia superba Dana larval forms (Furcilia-I and Furcilia-II stages) in the plankton of the Scotia and Weddell Seas have been investigated. New forms (1, 2, 3 and 14) have been recorded. The dominant forms of Furcilia-I stage are proposed to represent the initial dominants. In accordance with the initial dominance, five types of larval form combinations have been established. During the initial dominance change — from 5 to 2, the larval form number increases. This phenomenon is due to parallel development of euphauiid larvae; it results from a developmental pathway change in the main larval mass. Such deviations are reflected in an initial dominance change. In each pathway, the larvae pass through an unequal number of moults before reaching Form 5 (in accordance with the proposed moult scheme of Furcilia-I stage larvae). In the case of initial dominance of less-developed forms, the presence of different parallel pathways increases. As a result, the total number of larval forms present in the plankton changes, causing increased variability of the total number of larval forms. Each type of larval form combination is the result of summation of all larval forms, which derive from different developmental pathways of the larvae in different localities. The most usual developmental pathway for Euphausia superba is through Form 5, which may, thus, be regarded as the main developmental pathway for this species. This pathway probably corresponds to optimal environment conditions for larval growth and morphogenesis with maximal synchronisation of both processes. Consequently, it may be suggested that changes in developmental pathways of larvae are connected with suboptimal environmental conditions.     

Effect of increasing sea water pCO2 on the northern Atlantic krill species Nyctiphanes couchii

Surprisingly little is known about potential effects of ocean acidification on krill of the Northern Hemisphere as ecologically very important food web component. Sub-adult individuals of the northern Atlantic krill species Nyctiphanes couchii (caught at Austevoll near Bergen, Norway, in January 2013) were exposed in the laboratory to four different levels of pCO₂ (430, 800, 1,100, and 1,700 µatm) for 5 weeks in order to assess potential changes in a set of biological response variables. Survival decreased and the frequency of moulting-related deaths increased with increasing pCO₂. Survival was considerably reduced at relatively high pCO₂ of 1,700 µatm and tended to be negatively affected at 1,100 µatm pCO₂. However, the experimental results show no significant effects of pCO₂ on inter-moult period and growth at pCO₂ levels below 1,100 µatm. No differences in length measurements of the carapace and uropod were observed across pCO₂ levels, indicating no effect of changing carbonate chemistry on the morphology of those calciferous parts of the exoskeleton. The results suggest that sub-adult N. couchii may not suffer dramatically from predicted near-future changes in pCO₂. However, potential detrimental effects on the moulting process and associated higher mortality at 1,100 µatm pCO₂ cannot be excluded. Further experiments are needed in order to investigate whether early life stages of N. couchii show a different sensitivity to elevated sea water pCO₂ and whether those results are transferable to other krill species of the Northern Hemisphere.