Oceanic Copepods Research Articles

Comparative analysis of the molecular starvation response of Southern Ocean copepods.

Large lipid-storing copepods dominate mesozooplankton biomass in the polar oceans and form a critical link between primary production and higher trophic levels. The ecological success of these species depends on their ability to survive periods of food deprivation in a highly seasonal environment, but the molecular changes that mediate starvation tolerance in these taxa are unknown. We conducted starvation experiments for two dominant Southern Ocean copepods, Calanoides acutus and Calanus propinquus, allowing us to compare the molecular starvation response between species. These species differ in life history, diet and metabolic traits, and expressed overlapping but distinct transcriptomic responses to starvation. Most starvation-response genes were species-specific, but we identified a conserved core set of starvation-response genes related to RNA and protein metabolism. We used phylotranscriptomics to place these results in the context of copepod evolution and found that starvation-response genes are under strong purifying selection at the sequence level and stabilizing selection at the expression level, consistent with their role in mediating essential biological functions. Selection on starvation-response genes was especially strong in our focal lipid-storing lineage relative to other copepod taxa, underscoring the significance of starvation tolerance for these species. We also found that certain key lipid enzymes (elongases and desaturases) have experienced diversification and positive selection in lipid-storing lineages, reflecting the unique lipid storage needs of these animals. Our results shed light on the molecular adaptations of high-latitude zooplankton to variable food conditions and suggest that starvation-response genes are under particularly strong sequence and expression constraints.

Quantifying the feeding behavior and trophic impact of a widespread oceanic ctenophore

Oceanic ctenophores are widespread predators on pelagic zooplankton. While data on coastal ctenophores often show strong top-down predatory impacts in their ecosystems, differing morphologies, prey capture mechanisms and behaviors of oceanic species preclude the use of coastal data to draw conclusion on oceanic species. We used high-resolution imaging methods both in situ and in the laboratory to quantify interactions of Ocyropsis spp. with natural copepod prey. We confirmed that Ocyropsis spp. uses muscular lobe contraction and a prehensile mouth to capture prey, which is unique amongst ctenophores. This feeding mechanism results in high overall capture success whether encountering single or multiple prey between the lobes (71 and 81% respectively). However, multiple prey require several attempts for successful capture whereas single prey are often captured on the first attempt. Digestion of adult copepods takes 44 min at 25 °C and does not vary with ctenophore size. At high natural densities, we estimate that Ocyropsis spp. consume up to 40% of the daily copepod standing stock. This suggests that, when numerous, Ocyropsis spp. can exert strong top-down control on oceanic copepod populations. At more common densities, these animals consume only a small proportion of the daily copepod standing stock. However, compared to data from pelagic fishes and oceanic medusae, Ocyropsis spp. appears to be the dominant copepod predator in this habitat.

Ideal free flows of optimal foragers: Vertical migrations in the ocean

We consider the collective motion of animals in time-varying environments, using as a case the diel vertical migration of marine copepods. The animals are distributed in space such that each animal moves optimally, seeking regions which offer high growth rates and low mortalities, subject to costs on excessive movements as well as being in regions with high densities of conspecifics. The model applies to repeated scenarios such as diel or seasonal patterns, where the animals are aware of both current and future environmental conditions. We show that this problem can be viewed as a differential game of mean field type, and that the evolutionary stable solution, i.e., the Nash equilibrium, is characterized by partial differential equations, which govern the distributions and migration velocities of animals. These equations have similarities to equations that appear in the fluid dynamics, specifically the Euler equations for compressible inviscid fluids. If the environment is constant, the ideal free distribution emerges as an equilibrium. We illustrate the theory with a numerical example of vertical migrations of oceanic copepods, where animals are attracted to nutrient-rich surface waters while repulsed from light during daytime due to the presence of visual predators, aiming to reduce both proximity to conspecifics and swimming efforts. For this case, we show that optimal movements are diel vertical migrations in qualitative agreement with observations of copepods.

Diel metabolic patterns in a migratory oceanic copepod

Diel vertical migration of zooplankton profoundly impacts the transport of nutrients and carbon through the water column. Despite the acknowledged importance of this active flux to ocean biogeochemistry, these contributions remain poorly constrained, in part because daily variations in metabolic rates are not considered or are modeled as simple functions of temperature. To address this uncertainty, we sampled the subtropical copepod Pleuromamma xiphias at 4- to 7-h intervals throughout the daily migration and measured rates of oxygen consumption, ammonium excretion, fecal pellet production and metabolic enzyme activity. No significant patterns were detected in rates of oxygen consumption or ammonium excretion for freshly caught animals over the diel cycle. Fecal pellet production was highest during mid-night, consistent with several hours of feeding near the surface. Surface feeding resulted in fecal pellet production at depth in the morning, providing direct evidence that active flux of particulate organic carbon occurs in this region. Electron transport system activity was highest during the afternoon, contrary to our prediction of reduced daytime metabolism. Activity of both glutamate dehydrogenase and citrate synthase increased during early night, reflecting higher capacity for excretion and aerobic respiration, respectively. Overall, these results show that activities of metabolic enzymes vary during diel vertical migration. The surprising observation of elevated afternoon enzyme activity coupled with daytime fecal pellet and ammonium production suggests that additional characterization of the daytime activity of migratory zooplankton is warranted.

Copepods community structure and function under oceanographic influences and anthropic impacts from the narrowest continental shelf of Southwestern Atlantic

Studies on the Copepod community structure and function under oceanographic influences and anthropic impacts was carried out in the narrowest continental shelf of Brazil. A total of 65 taxa were identified, of which 39 were Calanoida. The most abundant species was Temora turbinata (65.26%) followed by Centropages velificatus, Paracalanuspavus, Undinula vulgaris and Clausocalanus furcatus. A mixture of coastal and oceanic species of Copepods was registered being difficult to separate both populations, however predominated the oceanic species due the narrow shelf and low influence of rivers flux. The frequency of occurrence showed that 24.62% of the species were constants, 13.85% accessories and 61.53% were accidentals. A diversity and abundance gradient were registered from neritic to oceanic waters, however diversity was very low to a shelf area under oceanic influence. The narrow shelf from the studied area permitted the movement of oceanic copepods to coastal areas. The reduced density was caused by the oligotrophic characteristic of the Tropical Water mass present in the area; however, the very low diversity was possibly caused by anthropic impacts (sewage outfalls), mainly in stations closer to the continent. Calocalanus pavo, Undinula vulgaris and Temora stylifera were the best indicators of the area and they were associated to phytoplankton abundance. The Copepod functional traits analyzed presented two sets separated by the reproductive behavior, the biggest set grouped small broadcasters, filter feeders mostly with an omnivore–herbivore strategy and a smaller group, sac-spawner that carry their eggs until hatching. The present study observed that the anthropogenic inputs had a great influence on copepods species and distribution, causing a different dominance pattern between nearshore and offshore.

De novo transcriptome assembly of the Southern Ocean copepod Rhincalanus gigas sheds light on developmental changes in gene expression

Copepods are small crustaceans that dominate most zooplankton communities in terms of both abundance and biomass. In the polar oceans, a subset of large lipid-storing copepods occupy central positions in the food web because of their important role in linking phytoplankton and microzooplankton with higher trophic levels. In this paper, we generated a high-quality de novo transcriptome for Rhincalanus gigas, the largest-and among the most abundant-of the Southern Ocean copepods. We then conducted transcriptional profiling to characterize the developmental transition between late-stage juveniles and adult females. We found that juvenile R. gigas substantially upregulate lipid synthesis and glycolysis pathways relative to females, as part of a developmental gene expression program that also implicates processes such as muscle growth, chitin formation, and ion transport. This study provides the first transcriptional profile of a developmental transition within Rhincalanus gigas or any endemic Southern Ocean copepod, thereby extending our understanding of copepod molecular physiology.

Predation risk alters life history strategies in an oceanic copepod.

The ubiquitous oceanic copepod Calanus finmarchicus is the major link between primary producers and important fish stocks in the North Atlantic Ocean and adjacent seas. Despite over a century of research on growth and development of this key species, the effect of predation risk on these processes remains elusive. We tested how food level and chemical cues from a fish predator influence growth and development of C. finmarchicus, using a predator naïve laboratory population. Copepods reached adult stage earlier both in response to high food and to predator cues in our experiment. High food also increased growth and lipid accumulation. In contrast, perceived predation risk triggered reduced size and lipid fullness, indicating a decoupling of growth and development rates. Our results demonstrate that chemical predator cues can influence life history strategies in C. finmarchicus, and suggest that present and future patterns in oceanic zooplankton size and population dynamics may also reflect differences in predation risk.

The spring mesozooplankton variability and its relationship with hydrobiological structure over year-to-year changes (2003–2013) in the southern Bay of Biscay (Northeast Atlantic)

Mesozooplankton can be considered the most important secondary producers in marine food webs because they hold an intermediate position between the phytoplankton assemblage and the upper trophic levels. They also are a robust indicator of climatic and hydrological conditions. We conducted an analysis of the interannual variability of the spring mesozooplankton assemblage, as sampled by the PELGAS fisheries survey in the southern part of the Bay of Biscay (Northeast Atlantic Ocean) between 2003 and 2013. We examined hydrology and trophic drivers to explain the variability. Our results revealed that the subsurface temperature, the subsurface salinity, the biomasses of subsurface pico-, nano-, and microphytoplankton, and the copepod assemblage exhibited a recurrent spatial pattern that was driven mainly by freshwater and nutrient inputs from the main rivers. The mesozooplankton assemblage was dominated by copepods (82%), composed of coastal, neritic, and oceanic copepod genera that paralleled the various hydrological fronts converging in the southern Bay of Biscay. The copepod community displayed high temporal-variability; there were three periods of abundant adult copepods throughout the southern Bay of Biscay. The copepod community was structured primarily around the drive for resource control, especially by the microphytoplankton biomass (24.3% of the total variability), and to a lesser extent by hydrological features (13.7% of the total variability).

Molecular evidence on evolutionary switching from particle-feeding to sophisticated carnivory in the calanoid copepod family Heterorhabdidae: drastic and rapid changes in functions of homologues

ABSTRACTThe oceanic calanoid copepod family Heterorhabdidae is unique in that it comprises both particle-feeding and carnivorous genera with some intermediate taxa. Both morphological and molecular (nuclear 18S and 28S rRNA) phylogenetic analyses of the family suggest that an evolutionary switch in feeding strategy, from transitions of typical particle-feeding through some intermediate modes to sophisticated carnivory, might have occurred with the development of a specially designed ‘poison’ injection system. In view of the small amount of genetic differentiation among genera in this family, the switching of feeding modes and re-colonization from deep to shallow waters might have occurred over a short geological period since the Middle to Late Miocene.

Non-random feeding enhances the contribution of oceanic zooplankton to the diet of the planktivorous coral reef fish Dascyllus flavicaudus

Combining gut content analysis and sampling of ambient zooplankton, we examine departures from random feeding in a planktivorous coral reef fish and explore the effects of apparent non-random feeding behavior on the relative contribution of oceanic versus reef-associated zooplankton to fish diet. The planktivorous damselfish Dascyllus flavicaudus appears to exhibit strong positive electivity for oceanic copepods including Candacia spp. and copepods from the families Oncaeidae and Corycaeidae and consistent negative electivity for cyclopoid copepods (Oithonidae). In total, prey taxa categorized as oceanic in origin contributed 10–76 % of total zooplankton biomass in fish guts. The summed contribution of oceanic prey taxa to fish diet was significantly higher than expected under a model of random feeding based on the availability of oceanic versus reef-associated prey as sampled by zooplankton net tows. The feeding behavior of D. flavicaudus appears to be visibility-selective rather than or in addition to size-selective, as electivity across prey taxa could not be explained by differences in prey size alone.

Ecological shift from piscivorous to planktivorous seabirds in the Chukchi Sea, 1975–2012

Sea ice now forms later and melts earlier than it did 4 decades years ago, and it now melts completely in all parts of the Chukchi Sea. This decline in sea ice is expected to have repercussions on the trophic structure in this environment, and there are indications that changes already have taken place in the seabird community. We compared boat-based densities of seabirds in the eastern Chukchi Sea between July and October during 1975–1981 (historical data) with densities during 2007–2012 (recent data). We related the composition of the seabird community to sea-ice cover to explore how the community may be responding to changes in oceanography. The seabird community historically was composed predominantly of piscivorous Black-legged Kittiwakes (Rissa tridactyla) and murres (Uria spp.). In contrast, the seabird community now is composed predominantly of planktivorous seabirds such as Crested Auklets (Aethia cristatella) and Short-tailed Shearwaters (Puffinus tenuirostris). Total abundance of seabirds declined in three of four strata in the eastern Chukchi Sea, largely due to declines in densities of piscivorous and omnivorous species. These changes in the abundance and community composition of seabirds were associated with changes in ice cover. Earlier ice retreat appears to contribute to an environment that is more favorable to the sustained production of large oceanic copepods and euphausiids. We propose that long-term changes (4 decades) in the abundance and composition of the seabird community reflect an increase in the availability of large zooplankton prey in the region.

Extensive genetic diversity and endemism across the global range of the oceanic copepod Pleuromamma abdominalis

Many oceanic zooplankton species have been described as cosmopolitan in distribution; however, recent molecular work has detected species complexity with highly divergent genetic lineages within several of these taxa. To further resolve the species complexity within these ecologically-important and widespread species, we performed both molecular and morphological analyses of the oceanic copepod Pleuromamma abdominalis using a comprehensive collection of material from 944 individuals collected at 46 sites across the global ocean. Phylogenetic analyses of mitochondrial cytochrome oxidase subunit I (mtCOI) sequences detected eighteen divergent evolutionary lineages within P. abdominalis, with an additional four singleton specimens that were also genetically divergent. Two phylogenetically distinct groups, PLAB1 and PLAB2, were supported by concordant sequence variation in the nuclear large subunit ribosomal RNA gene (nLSU). Within PLAB1, two mtCOI clades, 1a-1 and 1b-1 were observed, and each clade contained geographically distinct sub-clades 1a-2 and 1b-2. PLAB2 was composed of sixteen well-supported mtCOI clades (2a–2p) as well as four singletons. High genetic divergence among the mtCOI lineages within both PLAB1 and PLAB2, ranging between 9.2–11.2% and 4.3–18.9% K2P distances respectively, suggests the presence of additional species within these groups. Significant differences were observed in the presence and shape of antennule spines of adult females between sympatric clades with genetic distances greater than 5.7–7.0% (K2P). The biogeographic distributions of mtCOI clades indicated greater specialization to particular oceanographic provinces than observed in the nominal species P. abdominalis, with mtCOI clades ranging from antitropical in subtropical waters of all three ocean basins (Atlantic, Pacific and Indian; clade 1b-1 and 2a) to taxa that are endemic to a particular ocean region, for example restricted to equatorial waters of the Atlantic Ocean (clade 1b-2 and 2b). We hypothesize that many of these mtCOI clades are likely distinct, and currently undescribed species. The well-known high dominance of P. abdominalis across a range of pelagic habitats may occur as a spatial composite of genetically-distinct species with more restricted distributions and greater ecological specialization to particular marine habitats than was previously recognized.

Large-scale forcing of environmental conditions on subarctic copepods in the northern California Current system

In the ocean, dominant physical processes often change at various spatial and temporal scales. Here, we examined associations between large-scale physical forcing indexed by the Pacific Decadal Oscillation (PDO), regional ocean conditions including alongshore currents in relation to the abundance of two subarctic oceanic copepods, Neocalanus plumchrus, and N. cristatus in the offshore portions of the northern California Current (NCC) system in spring of 1998–2008. We found significant relationships between the abundance of copepods, water temperature, and alongshore currents with a lag of two or four months in response to the PDO in the NCC system. During the growth season in March/April both subarctic copepod species displayed consistent cross-shelf patterns with shoreward decreasing gradient in abundance, and were negatively correlated with the PDO, sea water temperature, and alongshore currents. Our studies highlight the responses of regional ocean conditions to large-scale physical forcing and illustrate the potential for Neocalanus copepods as unique vectors for a new understanding of the ecological response in the offshore oceanic waters of the NCC system to climate variability.

Seasonal and inter-species comparison of asymmetry in the genital system of some species of the oceanic copepod genus Metridia (Copepoda, Calanoida)

The seasonal and inter-annual changes in the asymmetry of female insemination and the male leg 5 of the planktonic calanoid copepods Metridia okhotensis and M. pacifica were investigated in the Okhotsk Sea. An inter-species comparison of both parameters was also carried out on seven Metridia species collected from oceans throughout the world. For M. okhotensis from the Okhotsk Sea, most of the females showed left-side insemination (annual average: 95.7%) and most of the males showed left-side asymmetry (99.7%) of the long inner process of the second exopodal segment in the fifth leg throughout the year. However, sympatric M. pacifica showed different ratios of asymmetry for female insemination and male morphotypes with a left : right = 1 : 2 ratio throughout the year. For the seven Metridia species from the global oceans treated in this study, ratios of asymmetry for female insemination and male morphotypes were correlated with each other. One-sided insemination (i.e., only left or only right insemination) was a common pattern for various Metridia species from global oceans, but their ratios varied by species. Previously, low hatching rates (29-68%) of eggs were reported for various Metridia spp. in laboratory experiments, but the reasons for these low rates were unclear. Because each spermatheca of Metridia spp. is connected to the oviduct on the same side, either left or right, this suggests that half of the eggs produced by unilaterally inseminated females remain unfertilized. The morphology of the genital structures and literature data of the egg hatching rates of Metridia spp. indicate that almost half of the eggs produced by females are not viable and are, thus, wasted.

Seasonal variation of the neustonic zooplankton community in southern waters of Korea

We investigated the seasonal variation of the neustonic zooplankton community in the surface layer (above 30 cm in depth) of 16 stations in southern waters of Korea from April 2002 to April 2003. In the neustonic community, copepods accounted for 59% of total organisms, as the most predominant group, while other dominant groups were invertebrate larvae, cladocerans, tunicates and fish eggs. Pontellid copepods, a major group of neustonic zooplankton, showed a marked seasonal variation and displayed a significant positive correlation with temperature (P < 0.05). Fish eggs appeared from spring to early summer in the inshore waters while tunicates and oceanic copepods (Eucalanus spp. and Euchaeta spp.) were dominant in the offshore waters during high temperature periods. This suggests that the distribution pattern of the neustonic zooplankton is seasonally affected by the hydrological conditions of in- and offshore waters. The abundance of zooplankton was higher in the surface layer (< 15 cm) than in the subsurface layer (15–30 cm), and was also higher at night than during the daytime. Of the dominant copepod families, the density of acartiid and pontellid copepods was higher in the surface layer than the subsurface layer. The density of acartiid and corycaeid copepods was significantly higher during daytime than nighttime. This indicates that the surface layer in temperate regions may provide a favorable habitat for neustonic copepods despite the dynamic environmental fluctuations.

Community structure of mesozooplankton in the western part of the Sea of Okhotsk in summer

We investigated the community structure of mesozooplankton in the western part of the Sea of Okhotsk in late summer, 2006. We recognized four communities belonging to two assemblages. A coastal assemblage dominated by the arctic planktonic snail Limacina helicina consisted of a gulf community characterized by brackish copepods and a continental shelf community characterized by the hydrozoan medusa Aglantha digitale and the arctic copepod Calanus glacialis. The other assemblage, characterized by the oceanic copepod Neocalanus plumchrus, consisted of a continental slope community characterized by a diverse species composition and a basin community characterized by the oceanic copepod N. cristatus. The continental slope community contained species from the coastal waters and was distributed along the course of the East Sakhalin current. This community may have been assembled by the incorporation of coastal water into the oceanic waters by the strong current. Small coastal copepods such as Oithona similis and Pseudocalanus spp. were the main components in all communities in terms of numbers, but larger copepods such as Neocalanus spp. and Metridia okhotensis were important in terms of weight, especially in the continental slope and basin communities. The population structures of the dominant species suggest that overall biological production is maintained by continuous reproduction or growth (or both) of L. helicina and small coastal copepods after the onset of seasonal dormancy of the large oceanic copepods in late summer.

Variation in assimilation efficiencies of dominant Neocalanus and Eucalanus copepods in the subarctic Pacific: Consequences for population structure models

The assimilation efficiency of zooplankton is an essential parameter required to estimate energy transfer to higher trophic levels in marine ecosystems. However, little information is available for large oceanic copepods, especially the Neocalanus and Eucalanus species dominant in the subarctic Pacific. In this study, the assimilation efficiencies of the C5 stages of Neocalanus cristatus, Neocalanus flemingeri and Eucalanus bungii were evaluated using eight phytoplankton species as food. The average assimilation efficiencies of N. cristatus, N. flemingeri and E. bungii ranged between 45 and 66%, 44 and 66% and 34 and 65%, respectively. The assimilation efficiency was highly variable depending on the food phytoplankton species. In all species, the assimilation efficiency showed a significant negative relationship with the ash content of the phytoplankton (r2=0.79–0.87, p<0.001). The assimilation efficiency of large-body sized N. cristatus for large-sized diatoms was higher than for the other copepod species. In population models of N. cristatus, changes in assimilation efficiency affect the growth and survival rates of the population. The Lagrangian ensemble model (LEM) for N. cristatus showed that, for assimilation efficiencies less than 57%, the population could not be maintained. Because variations in assimilation efficiency may have significant effects on the copepod population, their variability should be incorporated into marine ecosystem models in the future.

Multi‐decadal range changes vs. thermal adaptation for north east Atlantic oceanic copepods in the face of climate change

Populations may potentially respond to climate change in various ways including moving to new areas or alternatively staying where they are and adapting as conditions shift. Traditional laboratory and mesocosm experiments last days to weeks and thus only give a limited picture of thermal adaptation, whereas ocean warming occurring over decades allows the potential for selection of new strains better adapted to warmer conditions. Evidence for adaptation in natural systems is equivocal. We used a 50-year time series comprising of 117056 samples in the NE Atlantic, to quantify the abundance and distribution of two particularly important and abundant members of the ocean plankton (copepods of the genus Calanus) that play a key trophic role for fisheries. Abundance of C. finmarchicus, a cold-water species, and C. helgolandicus, a warm-water species, were negatively and positively related to sea surface temperature (SST) respectively. However, the abundance vs. SST relationships for neither species changed over time in a manner consistent with thermal adaptation. Accompanying the lack of evidence for thermal adaptation there has been an unabated range contraction for C. finmarchicus and range expansion for C. helgolandicus. Our evidence suggests that thermal adaptation has not mitigated the impacts of ocean warming for dramatic range changes of these key species and points to continued dramatic climate induced changes in the biology of the oceans.

Influences of mesoscale anticyclonic eddies on the zooplankton community south of the western Aleutian Islands during the summer of 2010

Mesoscale anticyclonic eddies have been observed south of the Aleutian Islands. Eddies farther east, in the Gulf of Alaska, are known to transport coastal water and coastal zooplankton to the offshore open ocean. The impacts of mesoscale anticyclonic eddies formed south of the western Aleutian Islands (Aleutian eddies) on the zooplankton community are not fully understood. In the present study, we describe zooplankton population structures within an Aleutian eddy and outside the eddy during July 2010. Based on the sea-level anomaly, the Aleutian eddy was formed south of Attu Island (172°54′E) in February 2010, and it moved southeastward in the next 5 months. Large oceanic copepods, Neocalanus cristatus, Eucalanus bungii and Metridia pacifica were more abundant inside the eddy than the outside. Inside the eddy, the life stage distribution of N. cristatus was more advanced than that outside, and Neocalanus spp. had accumulated more lipids. These conditions probably reflect the greater primary production in the eddy, production enhanced by nutrients advected into the eddy. The Aleutian eddy contained mostly oceanic copepods because it was formed in the offshore water and/or eddy–eddy interaction occurred after its formation. The sufficient food condition in the eddy presumably resulted in higher growth and survival rates of these oceanic copepods, resulting in the greater abundance, advanced development stages and greater lipid accumulation.

A modeling study to explore on-shelf transport of oceanic zooplankton in the Eastern Bering Sea

The Eastern Bering Sea shelf is divided into distinct hydrographic domains by structural fronts. Despite frontal obstructions to cross-shelf transport, each year large oceanic copepods—primarily Neocalanus spp.—are known to dominate the biomass of the outer-shelf zooplankton communities, and in some years are advected into the middle-shelf domain. Using ROMS (the Regional Ocean Modeling System), coupled with a float tracking model designed to represent ontogenetic vertical migration behavior of Neocalanus, we explored the mechanisms, timing, and location of the transport of oceanic zooplankton onto the eastern Bering Sea shelf from overwintering sources along the Gulf of Alaska and Bering Sea shelf breaks, under a variety of environmental conditions. Our float tracking experiments suggest that the timing of on-shelf transport and the distribution of oceanic zooplankton on the shelf can vary substantially between one year and another. The Bering, Pribilof, and Zhemchug Canyons and Cape Navarin were all regions of elevated on-shelf float transport. Wind direction was the primary factor controlling inter-annual variability in the timing, amount, and location of the on-shelf transport of our Neocalanus floats. Float transport across the northern and southern shelves responded in opposite directions to inter-annual differences in wind forcing: southeasterly wind enhanced on-shelf transport of the Neocalanus floats along the southern shelf but suppressed on-shelf transport over the northern shelf. Conversely, northwesterly wind suppressed on-shelf zooplankton transport onto the southern shelf but promoted enhanced transport around Cape Navarin on the northern shelf. Transport of the Neocalanus floats onto the shelf can be very episodic, reflecting the short duration of winds that promote on-shelf transport. Relatively short (days to weeks) periods of southeasterly wind between March and April significantly impacted the number of floats transported onto the shelf. The relative importance of different source areas to supplying oceanic zooplankton to the Bering Sea shelf does not appear to vary much from year to year. Our model results suggest that the Neocalanus found on the southern shelf most likely originate from overwintering sites in the Alaska Stream or the Eastern Bering Sea shelf break south of the Pribilof Islands, while Neocalanus found on the northern shelf most likely originate from sites north of the Pribilof Islands.