Functional responses for zooplankton feeding on multiple resources: a review of assumptions and biological dynamics

Modelers often need to quantify the rates at which zooplankton consume a variety of species, size classes and trophic types. Implicit in the equations used to describe the multiple resource functional response (i.e. how nutritional intake varies with resource densities) are assumptions that are not often stated, let alone tested. This is problematic because models are sensitive to the details of these formulations. Here, we enable modelers to make more informed decisions by providing them with a new framework for considering zooplankton feeding on multiple resources. We define a new classification of multiple resource responses that is based on preference, selection and switching, and we develop a set of mathematical diagnostics that elucidate model assumptions. We use these tools to evaluate the assumptions and biological dynamics inherent in published multiple resource responses. These models are shown to simulate different resource preferences, implied single resource responses, changes in intake with changing resource densities, nutritional benefits of generalism, and nutritional costs of selection. Certain formulations are further shown to exhibit anomalous dynamics such as negative switching and sub-optimal feeding. Such varied responses can have vastly different ecological consequences for both zooplankton and their resources; inappropriate choices may incorrectly quantify biologically-mediated fluxes and predict spurious dynamics. We discuss how our classes and diagnostics can help constrain parameters, interpret behaviors, and identify limitations to a formulation's applicability for both regional (e.g. High-Nitrate-Low-Chlorophyll regions comprising large areas of the Pacific) and large-scale applications (e.g. global biogeochemical or climate change models). Strategies for assessing uncertainty and for using the mathematics to guide future experimental investigations are also discussed.     

A comparison of three marine ecosystems surrounding the Korean peninsula: Responses to climate change

This study uses a comparative approach to examine responses of marine ecosystems to climatic regime shifts. The three seas surrounding the Korean peninsula, the Japan/East Sea, the East China Sea and the Yellow Sea represent three contiguous but distinct ecosystems. Sampling has been carried out by the National Fisheries Research and Development Institute of South Korea since 1965, using the same methods in all three seas. Sampling was generally synoptic. Amplitude time series of 1st EOF modes for temperature, salinity, zooplankton biomass and concentrations of four major zooplankton taxa were used to determine whether the three marine ecosystems respond in a similar manner to climate variations. Temporal patterns of the variables were strongly similar among the three seas at decadal time scales, but very weakly similar at interannual scales. All three seas responded to a climatic regime shift that occurred in 1989. Temperature, zooplankton biomass and copepod concentrations increased in the late 1980s or early 1990s in all three seas. Concentrations of amphipods, chaetognaths and euphausiids also increased in the Japan/East Sea and the East China Sea, but not the Yellow Sea. The Yellow Sea ecosystem differs strongly from the other two seas, and water exchange between the Yellow Sea and the East China Sea is much weaker than that between the East China Sea and Japan/East Sea. Spatial patterns of zooplankton determined by the EOF analysis were closely related to currents and fronts in each of the three seas.     

台湾海峡中、北部及邻近海域夏季浮游动物总生物量的分布特点

台湾海峡中、北部夏季浮游动物（非胶质）总生物量较丰富，1983年夏季的总生物量高于其他季节，1987年7、8月和1988年7月则高于1983年同期的生物量，这3a的高生物量分布区，一般都连片出现在上升流锋面区，显然和上升流的出现有密切关系。构成高生物量的优势种以外海广高盐种为主，生物量和高生物量区的分布态势都存在年际变化，这可能和不同性质水系的消长，相互推移及被囊动物的摄食竞争有关。     

The Southern Antarctic Circumpolar Current Front: physical and biological coupling at South Georgia

The coupling of physics and biology was examined along a 160 km long transect running out from the north coast of South Georgia Island and crossing the Southern Antarctic Circumpolar Current Front (SACCF) during late December 2000. Surface and near surface potential TS properties indicated the presence of three water types: a near-shore group of stations characterised by water which became progressively warmer and fresher closer to South Georgia, an offshore grouping in which sea surface temperatures and those at the winter water level were relatively warm (1.8°C and 0.5°C, respectively), and a third in which surface and winter water temperatures were cooler and reflected the presence of the SACCF. The transect bisected the SACCF twice, revealing that it was flowing in opposite directions, north-westward closest to South Georgia and south-eastwards at its furthest point from the island. The innermost limb was a narrow intense feature located just off the shelf break in 2000–3500 m of water and in which rapid surface baroclinic velocities (up to 35 cm s⁻¹) were encountered. Offshore in the outermost limb, shown subsequently to be a mesoscale eddy that had meandered south from the retroflected limb of the SACCF, flow was broader and slower with peak velocities around 20 cm s⁻¹. Chlorophyll a biomass was generally low (<1 mg m⁻³) over much of the transect but increased dramatically in the region of the innermost limb of the SACCF, where a deepening of the surface mixed layer was coincident with a subsurface chlorophyll maximum (7.4 mg m⁻³) and elevated concentrations down to 100 m. The bloom was coincident with depleted nutrient concentrations, particularly silicate, nitrate and phosphate, and although ammonium concentrations were locally depleted the bloom lay within an elevated band (up to 1.5 mmol m⁻³) associated with the frontal jet. Increased zooplankton abundance, higher copepod body carbon mass and egg production rates all showed a strong spatial integrity with the front. The population structure of the copepods Calanoides acutus and Rhincalanus gigas at stations within the front suggested that rather than simply resulting from entrainment and concentration within the jet, increased copepod abundance was the result of development in situ. Estimates of bloom duration, based on silicate and carbon budget calculations, set the likely duration between 82 and 122 d, a figure supported by the development schedule of the two copepod species. Given this timescale, model outputs from FRAM and OCCAM indicated that particles that occurred on the north side of South Georgia in December would have been in the central-southern Scotia Sea 2–3 months earlier, probably in sea ice affected regions.     

Mesozooplankton biomass and grazing responses to Cyclone Opal, a subtropical mesoscale eddy

As part of E-Flux III cruise studies in March 2005, plankton net collections were made to assess the effects of a cyclonic cold-core eddy (Cyclone Opal) on the biomass and grazing of mesozooplankton. Mesozooplankton biomass in the central region of Cyclone Opal, an area of uplifted nutricline and a subsurface diatom bloom, averaged 0.80±0.24 and 1.51±0.59 g DW m⁻², for day and night tows, respectively. These biomass estimates were about 80% higher than control (OUT) stations, with increases more or less proportionately distributed among size classes from 0.2 to >5 mm. Though elevated relative to surrounding waters south of the Hawaiian Islands (Hawai’i lee), total biomass and size distribution in Cyclone Opal were almost exactly the same as contemporary measurements made at Stn. ALOHA, 100 km north of the islands, by the HOT (Hawaii Ocean Time-series) Program. Mesozooplankton biomass and community composition at the OUT stations were also similar to ALOHA values from 1994 to 1996, preceding a recent decadal increase. These comparisons may therefore provide insight into production characteristics or biomass gradients associated with decadal changes at Stn. ALOHA. Gut fluorescence estimates were higher in Opal than in ambient waters, translating to grazing impacts of 0.11±0.02 d⁻¹ (IN) versus 0.03±0.01 d⁻¹ (OUT). Over the depth-integrated euphotic zone, mesozooplankton accounted for 30% of the combined grazing losses of phytoplankton to micro- and meso-herbivores in Opal, as compared to 13% at control stations. Estimates of active export flux by migrating zooplankton averaged 0.81 mmol C m⁻² d⁻¹ in Cyclone Opal and 0.37 mmol C m⁻² d⁻¹ at OUT stations, 53% and 24%, respectively, of the carbon export measured by passive sediment traps. Migrants also exported 0.18 mmol N m⁻² d⁻¹ (117% of trap N flux) in Cyclone Opal compared to 0.08 mmol N m⁻² d⁻¹ (51% of trap flux) at control stations. Overall, the food-web importance of mesozooplankton increased in Cyclone Opal both in absolute and relative terms. Diel migrants provided evidence for enhanced export flux in the eddy that was missed by sediment trap and ²³⁴Th techniques, and migrant-mediated flux was the major export term in the observed bloom-perturbation response and N mass balance of the eddy.     

Zooplankton research off Peru: A review

A review of zooplankton studies conducted in Peruvian marine waters is given. After a short history of the development of zooplankton research off Peru, we review zooplankton methodology, taxonomy, biodiversity, spatial distribution, seasonal and interannual variability, trophodynamics, secondary production, and modelling. We review studies on several micro-, meso-, macro-, and meroplankton groups, and give a species list from both published and unpublished reports. Three regional zooplankton groups have been identified: (1) a continental shelf group dominated by Acartia tonsa and Centropages brachiatus; (2) a continental slope group characterized by siphonophores, bivalves, foraminifera and radiolaria; (3) and a species-rich oceanic group. The highest zooplankton abundances and biomasses were often found between 4–6°S and 14–16°S, where continental shelves are narrow. Species composition changes with distance from the shore. Species composition and biomass also vary strongly on short time scales due to advection, peaks of larval production, trophic interactions, and community succession. The relation of zooplankton to climatic variability (ENSO and multi-decadal) and fish stocks is discussed in the context of ecological regime shifts. An intermediate upwelling hypothesis is proposed, based on the negative effects of low upwelling intensity in summer or extremely strong and enduring winter upwelling on zooplankton abundance off Peru. According to this hypothesis, intermediate upwelling creates an optimal environmental window for zooplankton communities. Finally, we highlight important knowledge gaps that warrant attention in future.     

Elevated biomass of mesozooplankton and enhanced fecal pellet flux in cyclonic and mode-water eddies in the Sargasso Sea

Accumulating evidence points to the importance of mesoscale eddies in supplying nutrients to surface waters in oligotrophic gyres. However, the nature of the biological response and its evolution over time has yet to be elucidated. Changes in mesozooplankton community composition due to eddy perturbation also could affect biogeochemical cycling. Over the course of two summers we sampled seven eddies in the Sargasso Sea. We focused on and followed a post-phytoplankton bloom cyclonic eddy (C1) in 2004 and a blooming mode-water anticyclonic eddy (A4) in 2005. We collected zooplankton in all eddies using a Multiple Opening and Closing Net Environmental Sampling System (MOCNESS) and quantified biomass (>0.15 mm, in five size fractions) from 0 to 700 m over nine discrete depth intervals. Zooplankton biomass (>0.5 mm) in the upper 150 m was similarly enhanced at night for the periphery of C1 and the center of A4 at 0.514 g m⁻² and 0.533 g m⁻², respectively, compared to outside (0.183 g m⁻² outside C1 and 0.197 g m⁻² outside A4). Despite minimal chlorophyll a enhancement and dominance by picoplankton in C1, zooplankton biomass increased most for the largest size class (>5 mm). Gut fluorescence for euphausiids and large copepods was also elevated on the C1 periphery. In A4, peak biomass occurred at eddy center coincident with peak primary production, but was highly variable (changing by >3-fold) over time, perhaps resulting from the dense, but patchy distribution of diatom chains in this region. Shifts in zooplankton community composition and abundance were reflected in enhancement of fecal pellet production and active transport by diel vertical migration in eddies. Inside C1 the flux of zooplankton fecal pellets at 150 m in June 2004 was 1.5-fold higher than outside the eddy, accounting for 9% of total particulate organic carbon (POC) flux. The flux of fecal pellets (mostly from copepods) increased through the summer in eddy A4, matching concurrent increases in zooplankton <2 mm in length, and accounting for up to 12% of total POC flux. Active carbon transport by vertically migrating zooplankton was 37% higher on the periphery of C1 and 74% higher at the center of A4 compared to the summer mean at the Bermuda Atlantic Time-series Study (BATS) station. Despite contrasting responses by the phytoplankton community to cyclonic and mode-water eddies, mesozooplankton biomass was similarly enhanced, possibly due to differential physical and biological aggregation mechanisms, and resulted in important zooplankton-mediated changes in mesoscale biogeochemistry.     

福建湄洲岛周围海域浮游动物的分布

根据１９９０年５月至１９９１年１月采自湄洲岛海域的浮游动物样品,鉴定出１０种终生浮游动物和１２类阶段性浮游幼体,桡足类的种类和数量在群落中均占优势。中华哲水蚤、精致真刺水蚤、中华假磷虾、肥胖箭虫和短尾类溲状幼体为主要优势种。海龙箭虫、拟经车浅室水母、针刺真浮莹等１０种,四季都有分布,本文还描述了总生物量,总个体和主要种类的分布,并对夏季浮游动物的分布与上升流的关系,以及总生物量季节纱明显进行了讨论     

Zooplankton Feeding Ecology: Contents of Fecal Pellets of the Cyclopoid Copepods Oncaea venusta, Corycaeus amazonicus, Oithona plumifera, and O. simplex from the Northern Gulf of Mexico

Abstract. In situ feeding habits of the cyclopoid copepods Oncaea venusta, Corycaeus amazonicus, Oithona plumifera , and O. simplex were investigated by scanning electron microscope examination of fecal pellets, the contents of which reflected copepod gut contents upon capture. Peilet contents were compared with assemblages of phytoplankton present in surface waters at times of copepod collection. All samples were from the northern Gulf of Mexico. All four copepods fed upon phytoplankton and O. venusta also ingested other crustaceans. Dominant components of fecal pellets generally did not mirror those of available phytoplankton assemblages. In some cases, O. venusta ingested primarily larger-sized particles even when these were not most abundant, and in other cases it did not ingest large cells even when they were present in bloom concentrations. The presence of small (< 2–5 urn diameter) centric diatoms in O. venusta pellets suggests the possibility of feeding by mechanisms other than suspension or raptorial feeding. Limited observations suggest that C. amazonicus and O. plumifera may feed raptorially on larger particles even when these are not particularly abundant, and that the small O. simplex (< 500 nm total length) feeds mainly upon nanoplankton. It appears that cyclopoid feeding mechanisms are complex, and likely more so than those of many calanoids.     

The vertical distribution of zooplankton in the western Irish Sea

Zooplankton diel vertical migration is evident on the mixed isothermal side of the western Irish Sea frontal system but is often influenced by large tides and persistent geostrophic currents. On the stratified side of the front, temperature acts as a controlling factor with most of the zooplankton occurring above the thermocline and carrying out pronounced vertical migration when chlorophyll a levels are low and diffuse. At higher chlorophyll levels, when discrete chlorophyll a maxima form, zooplankton vertical movement may be greatly modified with a large number of species and stages concentrating within these maxima at all times of the diel light cycle.