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Symbiosis includes commensalism, mutualism, and parasitism, however in many symbiotic relationships, it is difficult to determine which type they are in. Since trophic-linkage is a necessary factor to determine this, data on feeding ecology of symbionts is essential to understand symbiont-host relationships. Amphipods of the genus Dactylopleustes are known as obligate urchin symbionts. Though their ecology remains largely unknown, Dactylopleustes was recently reported to aggregate on diseased hosts, suggesting that Dactylopleustes feeds on diseased urchins’ tissues and uses urchins as both a habitat and prey. We investigated by DNA metabarcoding analyses, the feeding ecology of Dactylopleustes yoshimurai in relation to growth and whether host is disease or not. Contrary to our hypothesis, sequence reads from the gut contents were dominated by planktonic copepods regardless of body size or host disease. These results suggest that they mainly feed on copepod fecal pellets deposited on sediments, and do not have a strong trophic-linkage with their host. Large individuals on diseased urchins feed more on urchins than those on healthy urchins. However, their main prey still remains copepods, implying that host disease has a limited effect on the feeding behavior. In conclusion, our study supports this species is not a strong parasite but more of a commensal of urchins.

We examined the feeding habits of Benthosema pterotum in Kagoshima Bay. Benthosema pterotum undertook diel vertical migration and preyed in shallow waters at night. The most frequently consumed prey were copepods, and B. pterotum selectively foraged on poecilostomatoids. Poecilostomatoids were more predominant in the Kagoshima Bay zooplankton community than in neighboring sites, which suggested that the B. pterotum preference for poecilostomatoids may be an adaptation by the local population to prey availability. Benthosema pterotum probably plays an important role as a biological pump contributor and as a mediator that links secondary production and higher trophic levels.

The Kuroshio Current has been thought to be biologically unproductive because of its oligotrophic conditions and low plankton standing stocks. Even though vulnerable life stages of major foraging fishes risk being entrapped by frontal eddies and meanders and encountering low food availability, they have life cycle strategies that include growing and recruiting around the Kuroshio Current. Here we report that phytoplankton growth and consumption by microzooplankton are stimulated by turbulent nitrate flux amplified by the Kuroshio Current. Oceanographic observations demonstrate that the Kuroshio Current topographically enhances significant turbulent mixing and nitrate influx to the euphotic zone. Graduated nutrient enrichment experiments show that growth rates of phytoplankton and microheterotroph communities were stimulated within the range of the turbulent nitrate flux. Results of dilution experiments imply significant microzooplankton grazing on phytoplankton. We propose that these rapid and systematic trophodynamics enhance biological productivity in the Kuroshio.

Trophic sources and pathways supporting early life stages are crucial for survival of forage fishes recruiting around the oligotrophic and unproductive Kuroshio. However, information is limited for the Kuroshio planktonic food web and its trophodynamics because of its high biodiversity. Here, we explore trophic sources and linkages in the Kuroshio plankton community using metabarcoding analysis of gut-content DNA for 22 mesozooplankton groups. The major prey was dinoflagellates and calanoids for omnivorous groups, and calanoids and gelatinous organisms for carnivorous groups. Larvaceans and hydrozoans were the most frequently appeared prey for both omnivores and carnivores, whereas they were minor constituents of the available prey in water samples. Although calanoids overlapped as major prey items for both omnivores and carnivores because they were the most available, contributions from phytoplankton and gelatinous prey differed among taxonomic groups. Further analysis of the metabarcoding data showed that in addition to omnivorous copepods like calanoids, gelatinous groups like larvaceans and hydrozoans were important hubs in the planktonic food web with their multiple trophic linkages to many components. These findings suggest that gelatinous organisms are important as supplementary prey and provide evidence of niche segregation on trophic sources among mesozooplankton groups in the Kuroshio.

The Osumi Strait and its neighboring waters are major spawning grounds for commercially important small pelagic and coastal fish in the Pacific coastal area of Japan. Mesopelagic fish larvae are also abundant here, co-occurring with the larvae of commercial species. However, it is unknown whether these fish groups compete for prey. We examined the diet of four commercial fish groups (Engraulis japonicus, Trachurus japonicus, Triglidae spp., and Sebastiscus spp.) and two mesopelagic species (Sigmops gracilis, Myctophum asperum) during their larval stages to determine whether they had the same prey source. Morphological analysis and DNA metabarcoding of gut contents showed that the main prey species of all six fish groups were calanoid copepods. However, results from metabarcoding differed greatly from morphological analysis, showing that appendicularians were abundant in the guts of S. gracilis, M. asperum, and Sebastiscus spp. This may be because the methods differ in how they determine prey composition, with morphological analysis relying on counts of identifiable prey parts and metabarcoding being only a semi-quantitative method. In addition, appendicularians might be underestimated by metabarcoding analysis due to variation in genome size, copy number of target genes, and primer mismatches. Stable isotope analysis supported the importance of appendicularians as prey for S. gracilis, M. asperum, and Sebastiscus spp. Our results indicate that the two mesopelagic species and Sebastiscus spp. may compete for prey, but E. japonicus, T. japonicus, and Triglidae spp. occupy niches different from those of the two mesopelagic species. Our results imply that the trophic pathway via appendicularians may support the feeding of dominant mesopelagic species and enable coexistence with commercial species in the study area.

The Kuroshio current greatly affects biological processes in coastal areas. The Kagoshima Bay has good spawning ground for major migratory fish, serves as an important feeding area for fish larvae, and is affected by the Kuroshio current; however, there is little information concerning the biological response to the Kuroshio intrusion in this location. During the mixing period, the Kuroshio current approaches the Kagoshima Bay and a density flow, consisting of surface inflow and bottom outflow, is generated. We hypothesize that the bottom water in the bay, having a high nutrient concentration, mixes with the surface layer when the water flows out of the bay and that this contributes to the high biological production at the bay mouth. In this study, we conducted a field survey of the water quality in March 2016 and, using a three-end member mixing model, examined whether the bottom water of the bay could be a nutrient source supporting the phytoplankton bloom in the bay mouth. We discuss the fate of the organic matter derived from the primary production promoted by a nutrient supply event. The results indicate that the N-based phytoplankton biomass represented less than 80% of the N uptake by the phytoplankton, as estimated by subtracting the measured nitrate concentration from the supplied nitrate concentration based on the mixing model, indicating that the phytoplankton bloom at the bay mouth was supported by the supply of bottom water from the bay. Based on the C/N molar ratio (6.4) calculated by C/chlorophyll a (Chl-a) and N/Chl-a in this study, the highest value of nitrogen uptake was 12 mg C m−3 d−1 in terms of the carbon assimilation. The feeding rate of the mesozooplankton was estimated to be 13.6 mg C m−3 d−1 using a physiological model. An analysis of the food web structure is required to assess the effects of feeding by mesozooplankton; however, most of the phytoplankton bloom caused by mixing could be efficiently transferred to mesozooplankton in this study area.

The Kuroshio Current has been thought to be biologically unproductive due to oligotrophic conditions and low plankton standing stocks. Nevertheless, major foraging fishes are known to grow and recruit around the Kuroshio Current. While mixing and advection supplying nutrients to the euphotic zone are happened by eddies and meanders but limited at the Kuroshio front, there is a risk that survival of vulnerable life stages is encountered under the low food availability. Here we report that phytoplankton productivity is stimulated by turbulent nitrate flux amplified with the Kuroshio Current and rapidly transferred to microzooplankton through their grazing. Oceanographic observations demonstrate that the Kuroshio Current topographically enhances significant turbulent mixing and nitrate influx to the euphotic zone. Gradual nutrient enrichment experiments show growth rates of phytoplankton and microzooplankton communities stimulated within a range of the turbulent nitrate flux. Dilution experiments imply a significant microzooplankton grazing on phytoplankton. We propose that these rapid and systematic trophodynamics enhance invisible biological productivity in the Kuroshio.

[Abstract] / Vertical nitrate fluxes associated with turbulent mixing and upwelling around a small reef in the Kuroshio are quantified by continuously deploying a turbulence microstructure profiler with an attached submersible ultraviolet nitrate analyzer while drifting from the upstream to the downstream of the reef. Flow separations and trains of Kelvin-Helmholtz billows (thickness = 60 m) are identified using a shipboard ADCP and an echo-sounder. The turbulence diffusivity associated with the vigorous turbulent mixing reaches up to O(10^−1 m^2 s^−1), resulting in strong nitrate fluxes of O(1–10^3 mmol m^−2 day^−1). In addition, large differences between the upstream and downstream density profiles suggest a strong upwelling velocity of O(10^−3 m s^−1), as well as an upwelling nitrate flux of O(10^2 mmol m^−2 day^−1) in the entire subsurface layer. / / [Plain Language Summary] / Vertical nitrate fluxes associated with flow-topography interactions around a small reef in the Kuroshio Current are quantified using state-of-the-art oceanographic instruments. When the flow passes over a shallow sill on the flank of the reef, the velocity differences between layers intensify, resulting in a substantial vertical overturning and mixing of the water column. This turbulent mixing causes the observed nitrate flux from deep water to reach a value among the highest observed worldwide. / / [Key Points] / • Flow separations and trains of Kelvin-Helmholtz billows mix the water column around a small reef in the Kuroshio / • Doming of isopycnals/nitraclines suggests strong upwelling in the lee of the reef / • Turbulent nitrate fluxes reach up to O(10^3 mmol m^−2 day^−1)

The annual flux of biologically produced organic carbon from surface waters is equivalent to annual net community production (NCP) at a steady state and equals the export of particulate and dissolved organic carbon (POC and DOC, respectively) to the ocean interior. NCP was estimated from carbon budgets of salinity-normalized dissolved inorganic carbon (nDIC) inventories at two time-series stations in the western subarctic (K2) and subtropical (S1) North Pacific Ocean. By using quasi-monthly biogeochemical observations from 2004 to 2013, monthly mean nDIC inventories were integrated from the surface to the annual maximum mixed layer depth and corrected for changes due to net air–sea CO2 exchange, net CaCO3 production, vertical diffusion from the upper thermocline, and horizontal advection. The annual organic carbon flux at K2 (1.49 ± 0.42 mol m−2 year−1) was lower than S1 (2.81 ± 0.53 mol m−2 year−1) (p

We investigated seasonal changes in carbon demand and flux by mesozooplankton communities at subtropical (S1) and subarctic sites (K2) in the western North Pacific Ocean to compare the impact of mesozooplankton communities on the carbon budget in surface and mesopelagic layers. Fecal pellet fluxes were one order higher at K2 than at S1, and seemed to be enhanced by copepod and euphausiid egestion under high chlorophyll a concentrations. The decrease in pellet volume and the lack of any substantial change in shape composition during sink suggest a decline in fecal pellet flux due to coprorhexy and coprophagy. While respiratory and excretory carbon by diel migrants at depth (i.e., active carbon flux) was similar between the two sites, the actively transported carbon exceeded sinking fecal pellets at S1. Mesozooplankton carbon demand in surface and mesopelagic layers was higher at K2 than S1, and an excess of demand to primary production and sinking POC flux was found during some seasons at K2. We propose that this demand was met by supplementary carbon sources such as feeding on protozoans and fecal pellets at the surface and carnivory of migrants at mesopelagic depths.

Seasonal changes in mesozooplankton biomass and their community structures were observed at time-series stations K2 (subarctic) and S1 (subtropical) in the western North Pacific Ocean. At K2, the maximum biomass was observed during the spring when primary productivity was still low. The annual mean biomasses in the euphotic and 200- to 1000-m layers were 1.39 (day) and 2.49 (night) g C m−2 and 4.00 (day) and 3.63 (night) g C m−2, respectively. Mesozooplankton vertical distribution was bimodal and mesopelagic peak was observed in a 200- to 300-m layer; it mainly comprised dormant copepods. Copepods predominated in most sampling layers, but euphausiids were dominant at the surface during the night. At S1, the maximum biomass was observed during the spring and the peak timing of biomass followed those of chlorophyll a and primary productivity. The annual mean biomasses in the euphotic and 200- to 1000-m layers were 0.10 (day) and 0.21 (night) g C m−2 and 0.47 (day) and 0.26 (night) g C m−2, respectively. Copepods were dominant in most sampling layers, but their mean proportion was lower than that in K2. Mesozooplankton community characteristics at both sites were compared with those at other time-series stations in the North Pacific and with each other. The annual mean primary productivities and sinking POC fluxes were equivalent at both sites; however, mesozooplankton biomasses were higher at K2 than at S1. The difference of biomasses was probably caused by differences of individual carbon losses, population turnover rates, and trophic structures of communities between the two sites.

Many foraging fishes spend their vulnerable life stages with low food availability in the oligotrophic Kuroshio. Although they are likely dependent on microbial production as their energy source, there is limited knowledge of the trophodynamics of the planktonic food web in the Kuroshio of the East China Sea (ECS-Kuroshio). Here, we demonstrate energy sources and feeding impacts of microzooplankton on the phytoplankton community based on size-fractionated dilution experiments in the ECS-Kuroshio. Naked ciliates were the most predominant among micro-sized heterotrophs biomass throughout the study area. Pico- to nano-phytoplankton such as Prochlorococcus and haptophytes contributed to total chlorophyll a concentrations. The apparent growth rates in non-diluted bottles without nutrient enrichment were significantly negative for Prochlorococcus and haptophytes. Microzooplankton grazing rates were higher for nano-phytoplankton (0.17–1.45 d−1) compared with those for pico- (0.18–1.23 d−1) and micro-phytoplankton (0.06–0.92 d−1). The community grazing impacts accounted for up to 122% of primary production. These results suggest that one major pathway at lower trophic levels in the ECS-Kuroshio is from pico- and nano-phytoplankton to microzooplankton. This pathway could support energy transfer to higher trophic levels through food resources of mesozooplankton and fish larvae.

Drifting sediment trap experiments were conducted during various seasons to elucidate the characteristics of particles sinking through the upper 200 m of the water column in the western Pacific at subarctic station K2 and subtropical station S1. The sinking particle flux increased when primary productivity was high at each station, during June–July at K2 and during February at S1. Biogenic opal (Opal) and CaCO3 were the major components of the fluxes at K2 and S1, respectively. Contrary to the expectation of a high flux at the eutrophic station K2 and low flux at the oligotrophic station S1, the annual average organic carbon fluxes at 100 m were comparable at both stations: 62.7 mg C m−2 day−1 at K2 and 56.1 mg C m−2 day−1 at S1. The similarity of the fluxes was perhaps a reflection of the unexpectedly high primary production at S1. At K2, the organic carbon export ratio (organic carbon flux/primary productivity) was significantly and negatively correlated with primary production and tended to decrease with depth. The magnitude of the rate of attenuation of the organic carbon flux with depth was larger at S1 than at K2. This rate of attenuation tended to decrease and increase with primary production at K2 and S1, respectively. The explanation for these patterns may be that the flux of labile organic carbon at relatively shallow depths decreased with increasing primary production at K2, and zooplankton grazing pressure increased with increasing primary productivity at S1.

Several new approaches for measuring zooplankton growth and production rates have been developed since the publication of the ICES (International Council for the Exploration of the Sea) Zooplankton Methodology Manual (Harris et al., 2000). In this review, we summarize the advances in biochemical methods made in recent years. Our approach explores the rationale behind each method, the design of calibration experiments, the advantages and limitations of each method and their suitability as proxies for in situ rates of zooplankton community growth and production. We also provide detailed protocols for the existing methods and information relevant to scientists wanting to apply, calibrate or develop these biochemical indices for zooplankton production. Versión del editor

Zooplankton growth rates ultimately shape the functional response of marine ecosystems to regional and global climate changes, because they determine the quantity and distribution of matter and energy within the zooplankton community available to higher trophic levels. Despite the variety of techniques available for measuring zooplankton growth, no or few approaches have been universally applied to the natural zooplankton populations or community and there are only a limited number of comparisons among the methods. Here we review and compile data for the traditional methods for estimating metazooplankton weight-specific growth rates, describe the principles and underlying assumptions of each method, and finally their advantages and disadvantages. This review encompasses the analysis of time-series (i.e., Natural Cohort method), three experimental approaches (i.e., Artificial Cohort, Molting Rate and Egg Production) and several empirical models that have been applied to specific stages, populations or community guilds of metazooplankton in the field. Whereas, some methodological problems and their resolution have been proposed in the past, no single method adequately addresses the high biodiversity of metazooplankton communities and resolves our limited capacity determining rates. We recommend a more formal comparison of methodologies be undertaken that would allow for their direct cross-calibration to facilitate future cross-site synthesis.

We compared wintertime depth distributions of the mesozooplankton community and dominant copepods between the subtropical (S1) and subarctic (K2) Pacific Oceans to evaluate the relative importance of actively transported carbon by vertical migrants to sinking particulate organic carbon flux. Primary production was higher and the ratio of sinking particulate organic carbon flux to primary production was lower at S1 compared with those at K2. The mesozooplankton community was lower in abundance and biomass at S1 compared to K2. Copepods were the dominant group among both mesozooplankton abundance and biomass throughout the water column down to 1000 m at both sites. The depth distribution showed that diel vertical migration was obvious for the mesozooplankton abundance and biomass at S1 but was not apparent for the abundance at K2, because the dominant component was diurnally migrating species at S1 and overwintering species residing at mesopelagic depths at K2. The major components of diel migrants were copepods and euphausiids at S1 and only euphausiids at K2. Respiratory flux by the diurnally migrating mesozooplankton was estimated to be 2 mgC m −2 day −1 at S1 and 7 mgC m −2 day −1 at K2. The respiratory flux was equivalent to 131% of sedimentary fecal pellet flux at S1 and 136% of that at K2. Because pathways of downward carbon flux are facilitated by the mesozooplankton community, the actively transported carbon (respiration of dissolved inorganic carbon, excretion of dissolved organic carbon and egestion of fecal pellets at depth) might be larger during winter than the flux of sinking fecal pellets.

We compared food availability of filter-feeding bivalves, Anadara spp., between two Fijian sites of different mangrove richness to evaluate impacts of environmental variables on Anadara spp. abundance and body size. Suspended particles including planktonic organisms and detritus were more abundant in the fishery grounds of the mangrove-rich site (MR) than in the mangrove-poor site (MP). Athough no substantial difference was observed in abundance of Anadara spp., dry weights of soft tissue were heavier for animals at MR than those at MP. Respiration rates (i.e., minimum metabolic requirements) of Anadara spp. decreased with increasing animal weight. Unicellular planktonic biomass was estimated to support the Anadara community metabolic requirements (i.e., minimum food requirement) for 9.2 to 85.7 days at MR and 1.4 to 67.4 days at MP, indicating that the planktonic biomass cannot support sufficient growth of the bivalve population at some locations. These results suggest that suspended particles support increased shell sizes of Anadara spp. and that resuspended detritus is a supplement or alternative food resource for these bivalves in mangrovecoral associated ecosystems.

We investigated individual DNA, RNA and protein (PRO) content of the ontogenetically migrating copepods Neocalanus cristatus, N. plumchrus and Eucalanus bungii collected from surface layers; of the northwestern Pacific Ocean during the spring phytoplankton bloom and mesopelagic depths during winter. RNA:DNA and RNA:PRO ratios for dormant and adult stages residing at mesopelagic depths were significantly lower than those for animals collected from the surface layers. Compiling the values of all species and stages together, log-transformed individual content of RNA (logRNA) was significantly correlated with log-transformed individual contents of DNA (logDNA) and protein (logPRO) in both layers. While logRNA:logDNA and logRNA:logPRO ratios were significantly lower for specimens residing at mesopelagic depths, the difference of the logRNA:logPRO ratio was not clear for some specimens appearing in the surface layer. These results indicate that in these ontogenetically migrating copepods, RNA to DNA decreases considerably during dormant and adult stages residing at mesopelagic depths. We propose the RNA:DNA ratio as a biochemical indicator for dormancy of ontogenetically migrating copepods.

Regional differences in the mechanisms of temporal variation in the lower trophic levels in the western, central, and eastern subarctic North Pacific were studied using the nitrogen stable isotope ratio (δ 15 N) of the major copepod species, Neocalanus cristatus , Neocalanus flemingeri , and Neocalanus plumchrus . We used formalin-preserved specimens collected in the Oyashio region (OY), three sections from north to south along the 180° longitudinal line (180LineSA, TN, and TS), off Vancouver Island (Off-Van), and at Sta. P, during the periods of 1960–2000, 1979–1997, 1981–2007, and 1996–2007, respectively. The regional mean δ 15 N of the three species roughly corresponded to the surface nitrate distribution and the extent of its drawdown from winter to spring; it was higher in regions of larger seasonal drawdown as observed in the coastal regions OY and Off-Van (7–10‰), but lower in regions with less seasonal drawdown, such as in the offshore regions at St. P and stations along the 180Line (3–6‰). Time series analysis revealed possible region-specific mechanisms for temporal variation in Neocalanus δ 15 N. First, δ 15 N indicated shifts in feeding strategies between herbivorous to omnivorous/carnivorous at OY and 180LineSA, where δ 15 N tended to be lower in the years with warmer winters, suggesting that Neocalanus took advantage of enhanced phytoplankton production under favorable light availability due to increased stratification. Conversely, wind-induced latitudinal advection of surface water was considered to be the initial cause of interannual variation in Neocalanus δ 15 N at 180LineTN, 180LineTS, and Off-Van, where δ 15 N was higher in the years with strong southerly or westerly winds at 180LineTN and TS, and the Off-Van site. This suggests that pole-ward transport of relatively oligotrophic, southern water might enhance the uptake of the heavier isotope by phytoplankton, which Neocalanus feed upon. Another possibility at the Off-Van site, where high δ 15 N was observed ( c.a. , 8–10‰), is a switch in the Neocalanus feeding strategy induced by decreased phytoplankton availability. This study demonstrated the usefulness of zooplankton δ 15 N as an indicator of interannual variation in lower trophic level environments and food web structures, which are caused by region-specific mechanisms.

Seasonal changes in abundance, growth and mortality of heterotrophic bacteria were investigated monthly from collections and dilution experiments in Kagoshima Bay, the southernmost of Japan. Bacteria occurred abundantly with considerable variation in the surface layers where chlorophyll a concentrations were high, whereas seasonal variations were obscure below 100 m. Especially, bacteria showed a decline of cell density toward summer when heterotrophic nano-flagellates increased their abundance. Seasonal and vertical variations in bacterial cell number during the study period were positively correlated with those of water temperature and pico-sized chlorophyll a concentration. Maximum growth and mortality rates showed positive correlations with water temperature but no positive relationships to size-fractionated chlorophyll a. Net increase rate (i.e. in situ rate if abundance changes) was negatively correlated with cell density of heterotrophic nano-flagellates. It is concluded that bacterial abundance is controlled by the resultant (i.e. net growth rate) of the balance between maximum growth and predatory mortality by heterotrophic nano-flagelllates which are both dependent on ambient temperature.

Seasonal changes in the shape and size composition of fecal pellets were investigated with sediment trap samples from 50 and 150 m in Kagoshima Bay to evaluate how the mesozooplankton community affects fecal pellet flux. Deep vertical mixing was evident in March, and thermal stratification was developed above 50 m in June, August and November. Chlorophyll a, suspended particulate organic carbon (POC) and copepod abundance were uniform throughout the water column during the seasonal mixing and concentrated above 50 m in the stratified seasons. Calanoids were the most predominant copepods in March and poecilostomatoids composed more than 45% of the copepod community in June, August and November. Fecal pellet fluxes at 50 and 150 m were the highest in March, nearly half of POC flux. The relative contribution declined considerably in the other months, especially for less than 4% of POC flux in August. The decline was corresponded to the predominance of cyclopoids and poecilostomatoids. Cylindrical pellets dominated the fecal matters at both depths throughout the study period, while larger cylindrical pellets nearly disappeared at 150 m in June, August and November. Copepod incubation revealed that cylindrical and oval pellets were egested by calanoids and the other copepods, respectively. We suggest that cylindrical fecal pellets produced by calanoid copepods contribute to feces flux but the predominance of poecilostomatoids and/or cyclopoids decreases feces flux via the increase of oval pellets and fragmentation of larger cylindrical pellets.

We have evaluated development and growth of the ontogenetically migrating copepods in the northwestern subarctic Pacific by frequent sampling and by molting rate experiments during the spring phytoplankton bloom. Since different water masses frequently moved into the surface layers at our sampling station, the copepod abundance and species composition fluctuated greatly. Early copepodite stages of Eucalanus bungii and Neocalanus plumchrus appeared abundantly when a warmer and more saline water mass was present. Despite the fluctuating temporal pattern, the population structure revealed that each copepod species had a different life history strategy during the bloom. Eucalanus bungii were in the stages of gonad maturation, spawning and naupliar development when chlorophyll a concentrations were high. The phytoplankton bloom was utilized for development and lipid accumulation by late copepodites of Neocalanus cristatus and Neocalanus flemingeri and for development of nauplii and early copepodites of N. plumchrus. Molting experiments showed that carbon weight of ‘molters’ in the incubations was greater than that of ‘non-molters’, indicating that primarily animals with heavier body weight are those that molt into the next stage. Mean stage durations estimated by the molting rate method were on the order of 9.7 (C3) to 16.6 days (C4) for N. flemingeri, 13.9 (C3) to 29.1 days (C4) for N. plumchrus, and 12.2 days (C2) for E. bungii. Large fluctuations were observed for stage duration estimates, suggesting different development histories (i.e. ages-within-stage distributions) among the replicate incubations. From these results, we discuss development and growth of the ontogenetically migrating copepods during strongly fluctuating hydrographic conditions in the Oyashio.