Biology subjects

Stukel, M. R.

Publications and source records attributed to Stukel, M. R..

17 recordsLinked to original sources

bioRxiv2026

Comparative food-web analysis of bluefin tuna spawning habitats in the eastern Indian Ocean and Gulf of Mexico

Using linear inverse ecosystem modeling as a data assimilation tool, we compare spawning grounds of Atlantic and Southern Bluefin Tuna (ABT and SBT, respectively) based on results from field campaigns in the Gulf of Mexico (GoM) and eastern Indian Ocean off northwest Australia (Argo Basin). Both regions are warm, stratified, low-nutrient waters dominated by cyanobacteria (Prochlorococcus). Despite these similarities, the Argo Basin is more productive, with [~]1.5X higher net primary production and nearly 2X higher production of top trophic levels in the model (tuna larvae, planktivorous fish, and predatory gelatinous zooplankton). Higher primary production in the Argo Basin is mainly driven by higher N2 fixation and storm mixing of new nutrients in the upper and lower euphotic zone, respectively. Increased ecosystem efficiency (secondary production of top trophic levels / primary production) results from differences in plankton food web organization. In the GoM, protistan zooplankton are the direct consumers of nearly all phytoplankton production. In contrast, higher rates of herbivory by crustaceans feeding on nanophytoplankton combines with a higher impact of appendicularians on cyanobacteria to convert plankton production into larval tuna prey more efficiently in the Argo Basin. Despite similarities in the proportions of phytoplankton production mediated by cyanobacteria and other picoplankton in both systems, food web pathways to larval tuna and other planktivorous fish are substantially shorter in the Argo Basin. Our results highlight the impact of distinct zooplankton ecological niches on ecosystem efficiency and suggest a need for better inclusion of plankton food-web structure in models simulating climate impacts on fisheries production. HIGHLIGHTSO_LIDeveloped food web models of tuna spawning habitat (Indian Ocean & Gulf of Mexico) C_LIO_LISpawning habitats in the Argo Basin and Gulf of Mexico (GoM) are both oligotrophic C_LIO_LIArgo Basin had higher net primary production in part as a result of nitrogen fixation C_LIO_LIArgo Basin had higher rates of direct herbivory by metazoan zooplankton C_LIO_LIThis resulted in greater ecosystem efficiency in the Argo Basin. C_LI

ecology↗

bioRxiv2025

The ecology of carbon sequestration in coastal upwelling ecosystems

The biological carbon pump (BCP) transports carbon fixed by phytoplankton into the deep ocean via multiple pathways and leads to net carbon dioxide sequestration. The BCP is very active in highly productive eastern boundary upwelling systems (EBUSs), however, the extent to which climate change will impact this key ecosystem service remains uncertain. While this review and synthesis focuses on results from the California Current Ecosystem (the most extensively studied EBUS), similarities and differences are also noted with the Benguela, Canary, and Humboldt ecosystems. We focus on vertical carbon export mediated by sinking particles (responsible for >half of the BCP in EBUSs), as well as subduction of organic matter and vertically migrating zooplankton and fish. We suggest that a plug-flow-reactor conceptual model can be used to link BCP results with physical circulation changes within EBUSs to predict whether climate change will lead to increased or decreased biologically mediated CO2 uptake.

ecology↗

bioRxiv2025

Analysis of environmental data during a lagrangian experiment: The influence of vertical movements.

Investigating the time evolution of physical and biochemical properties of the ocean with in situ sampling can follow two approaches: Eulerian and Lagrangian. In the Eulerian approach, repeated measurements are taken at fixed locations, whereas the sampling point moves with the displacement of a water parcel in the Lagrangian approach. During the BLOOFINZ-IO cruise off northwest Australia, four Lagrangian experiments ("cycles") were conducted with multidisciplinary sampling done at regular intervals for several days following a satellite tracked drifter with mixed-layer drogue. To test the Lagrangian nature of these experiments, we adapted the Bindoff and McDougall (1994) approach for decomposing observed changes between adjacent CTD profiles into components due to vertical movement (heaving) and those occurring along isopycnal surfaces. Profile depth variability was mainly driven by vertical displacements of isopycnals (internal waves), while temperature, chlorophyll, oxygen and salinity were relatively stable when observed on isopycnal surfaces across all casts within the same cycle, and different on average between cycles. Our analysis clearly indicated that density surface was a more appropriate vertical coordinate than physical depth for assessing real environmental variability during each cycle and confirmed the Lagrangian character of the experiments. While the BLOOFINZ cruise serves as a case study, the methodology can be readily extended to analyze other biochemical variables in different ocean regions.

ecology↗

bioRxiv2025

Primary productivity and N2-fixation in the eastern Indian Ocean: bottom-up support for an ecologically and economically important ecosystem

Oligotrophic regions of the global ocean are characterized by strong nutrient limitation, low standing phytoplankton biomass, and highly efficient nutrient recycling. We quantified nutrient inventories, primary productivity and N2 fixation during the BLOOFINZ-IO expedition (February 2022) in the Argo Basin located in the eastern Indian Ocean, the sole known spawning ground for Southern Bluefin Tuna. Surface nitrate concentrations were near depletion (<0.02 {micro}mol L-1), with low but persistent residual phosphate (P) concentrations suggesting nitrogen as the major limiting nutrient. Depth-integrated net primary production (NPP), from 14C-based in-situ incubations during 4 Lagrangian cycles, averaged [~]460 mg C m-2d-1, in good agreement with satellite-based NPP estimates. Nitrogen fixation provided a consistent new nitrogen source, contributing [~]16% to local NPP in the upper euphotic zone. Gross primary production (GPP), derived from fast-repetition-rate-fluorometry-based electron transport estimates, revealed significant autotrophic respiration losses, with GPP:NPP ratios averaging [~]1.8, consistent with metabolic costs under nutrient limitation. Net community production (NCP), estimated from O2/Ar ratios, remained positive across all cycles, averaging [~]20% of NPP in the upper 30 m. This result, in combination with N2 fixation measurement indicates that N2 fixation supports most of the export production in this region. Together, the multi-method approach revealed a recycling-dominated ecosystem affected by episodic mixing events, where primary productivity is maintained primarily through efficient nitrogen recycling and physiological photoacclimation. These results provide a comprehensive baseline of bottom-up support on ecosystem productivity for the Argo Basin for assessing future climate-driven changes in stratification, nutrient cycling, and food-web dynamics.

ecology↗

bioRxiv2025

Microbial food web dynamics in tropical waters of the bluefin tuna spawning region off northwestern Australia

Whereas recruitment success for many fisheries depends on coincident timing of larvae with abundance peaks of their prey, less can be more in the tropical/subtropical spawning areas of bluefin tunas if lower but steady food resources are offset by reduced larval vulnerability to pelagic predators. To understand larval habitat characteristics for Southern Bluefin Tuna (SBT), we quantified microbial community carbon flows based on growth and grazing rates from depth profiles of dilution incubations and carbon biomass assessments from microscopy and flow cytometry (FCM) during their peak spawning off NW Australia (Indian Ocean) in February 2022. Two Chla-based estimates of phytoplankton production gave differing offsets due to cycling or mixotrophy, exceeding 14C net community production on average (677 {+/-} 98 versus 447 {+/-} 43 mg C m-2 d-1). Productivity was higher than in the Gulf of Mexico spawning area for Atlantic Bluefin Tuna but less than similar studies of oceanic upwelling regions. Microzooplankton grazing averaged 482 {+/-} 63 mg C m- 2 d-1 (71 {+/-} 13% of production). Two measurement variables for Prochlorococcus gave average production and grazing rates of 282 {+/-} 36 and 248 {+/-} 32 mg C m-2 d-1 (86 {+/-} 6% grazed). Prochlorococcus comprised almost half of production and grazing fluxes in the upper (0-25 m) euphotic zone where SBT larvae reside. Prochlorococcus declined and eukaryotic phytoplankton and heterotrophic bacteria increased in relative importance in the lower euphotic zone. These results describe relatively classic open-ocean oligotrophic conditions as the food web base for nutritional flows to SBT larvae.

ecology↗

bioRxiv2025

Sinking particle export within and beneath the euphotic zone in the eastern Indian Ocean

The eastern Indian Ocean is substantially under sampled with respect to the biological carbon pump - the suite of processes that transport the carbon fixed by phytoplankton into the deeper ocean. Using sediment traps and other ecosystem measurements, we quantified sinking organic matter flux and investigated the characteristics of sinking particles in waters overlying the Argo Abyssal Plain directly downstream of the Indonesian Throughflow off northwest Australia. Carbon export from the euphotic zone averaged 7.0 mmol C m-2 d-1, which equated to an average export efficiency (export / net primary production) of 0.17. Sinking particle flux within the euphotic zone (beneath the mixed layer, but above the deep chlorophyll maximum) averaged slightly higher than flux at the base of the euphotic zone, suggesting that the deep euphotic zone was a depth stratum of net particle remineralization. Carbon flux attenuation continued into the twilight zone with a transfer efficiency (export at euphotic depth + 100m / export at euphotic depth) of 0.62 and an average Martins b-value of 1.1. Within the euphotic zone, fresh phytoplankton (chlorophyll associated with sinking particles, possibly contained within appendicularian houses) were an important component of sinking particles, but beneath the euphotic zone the fecal pellets of herbivorous zooplankton (phaeopigments) were more important. Changes in carbon and nitrogen isotopic composition with depth further reflected remineralization processes occurring as particles sank. We show similarities with biological carbon pump functioning in a similar semi-enclosed oligotrophic marginal sea, the Gulf of Mexico, including net remineralization across the deep chlorophyll maximum. Submitted to: Deep-sea Research II HighlightsDespite low productivity, export efficiency was 17% of primary production Flux attenuation beneath the euphotic zone (EZ) was low for a tropical region Sinking particle flux from the upper to lower EZ exceeded export from lower EZ The deep EZ was a stratum of net particle remineralization (and net heterotrophy)

ecology↗

bioRxiv2025

Export dynamics of protists across the southern subtropical frontal zone reveal taxon-specific patterns

Gravitational particle sinking is the main mechanism for carbon export in the biological carbon pump. However, the export dynamics of the particle-associated protist community are not fully understood. We used 18S rRNA gene metabarcoding to characterise the exported protist community within sinking particles and bathypelagic surficial sediments in oligotrophic subtropical and high-nutrient, low-chlorophyll subantarctic waters. Sinking particles were collected with formalin-fixed and preservative-free particle interceptor traps (fixed and live traps, respectively) to identify the community involved in particle export (fixed) and protist loss from remineralisation (live). We paired this with community analysis of the upper and lower water column (mixed layer and below mixed layer to mesopelagic, respectively) to compare the relative sources of exported protists. Amplicon sequences variants (ASVs) from upper water column samples accounted for 2 to 4-fold higher proportion of reads and ASV rich-ness compared to lower water column samples in fixed trap and sediment samples, suggesting low influence of the suspended protist community from the lower water column on export. We further traced the export patterns of upper water column protist taxa by analysing the change in taxa relative abundance across the mixed layer to mesopelagic depths. Export patterns differed between taxa, which is similarly suggested by taxa-specific loss of ASV richness between fixed and live traps, but remained the same across biogeochemically-contrasting water masses. This could imply that the drivers for protist loss during export are related to characteristics consistent across environmental conditions, such as specific microbial interactions or inherent cell properties.

microbiology↗

bioRxiv2024

Phago-mixotrophy of small eukaryotic phytoplankton might alleviate iron limitation in HNLC Southern Ocean

Small phytoplankton, consisting of pico and nano size fractions, are diverse in size and taxonomy. Yet, the differences in their productivity and taxonomic diversity are poorly described. Here, we measured the cell-specific carbon fixation rates of picocyanobacteria Synechococcus, picoeukaryote and nanoeukaryote populations while unveiling their taxonomic composition in oligotrophic subtropical (ST) and high-nutrient low-chlorophyll subantarctic (SA) waters. We coupled 24 h in-situ radiolabelled 14C incubations to flow cytometry sorting (FCM-sorting) and DNA metabarcoding from the same incubated samples, offering a direct account of the community associated with the carbon fixation rates measured. In both water masses, nanoeukaryotes had the highest cell-specific carbon fixation rate, followed by picoeukaryotes and Synechococcus (2.24 {+/-} 29.99, 2.18 {+/-} 2.08 and 0.78 {+/-} 0.55 fgC cell-1 h-1, respectively). The cell-specific carbon fixation rates and growth rates of Synechococcus were 3-fold higher in ST compared to SA waters, while the rates of picoeukaryotes and nanoeukaryotes had no significant difference between the biogeochemically-contrasting water masses. Despite significant differences in their taxonomic composition, the FCM-sorted picoeukaryote and nanoeukaryote populations in SA waters were dominated by taxa with reported phago-mixotrophic strategies (Chrysophyceae, Dinophyceae and Prymnesiophyceae), suggesting phago-mixotrophy might alleviate nutrient stress in iron-limited conditions for discrete small photosynthetic eukaryote populations.

microbiology↗

bioRxiv2022

Prey Size Spectra and Predator:Prey ratios of 7 Species of New Zealand Salps

Salp grazing is important in shaping planktonic food-web structure. However, little is known about the size ranges of their prey in the field or how grazing impacts size structure. This study investigated the feeding habits of 7 different species of salps, representing a variety of sizes and life stages across subtropical and subantarctic waters east of New Zealand. Scanning electron microscopy was used to examine the gut contents of 58 salps, which were then compared to water column plankton communities characterized via epifluorescence microscopy, FlowCam, and flow cytometry. While most of the gut contents resembled ambient waters, substantial differences were found amongst some co-occurring species, such as increased retention of submicron bacteria amongst smaller salps like Thalia democratica. We found that even for those salps capable of feeding on bacteria efficiently, nanoplankton and small microplankton still made up the majority of gut biomass. Larger microplankton were rarer in the guts than in the water column, potentially suggesting an upper size-threshold in addition to the lower size-threshold that has been the focus of most previous work. Salp carbon-weighted predator to prey size ratios were variable, with the majority falling between 1,000:1 and 10,000:1 depending largely on the size of the salp. Taken together our results indicate that despite being able to feed on submicron particles, picoplankton make up at most 26.4% (mean = 6.4%) of salp gut carbon and are relatively unimportant to the energetics of most salps in this region compared to nanoplankton such as small dinoflagellates and diatoms.

ecology↗

bioRxiv2022

Salp blooms increase carbon export 5-fold in the Southern Ocean

The Southern Ocean (SO) contributes substantially to the global biological carbon pump (BCP). Salps in the SO, in particular Salpa thompsoni, are keystone grazers that produce large, fast-sinking fecal pellets with high export potential. In a first study of this kind, we conducted Lagrangian experiments to quantify the salp bloom impacts on export pathways by contrasting locations differing in salp bloom presence/absence. We show that blooms increased particle export by ~5-fold, and exported up to 46% of net primary production out of the euphotic zone. BCP efficiency increased from 5% in non-salp areas to up to 28% in salp areas, which is among the highest recorded in the global ocean. Using SO salp abundances from KRILLBASE, we estimate they can consume ~ 13% of regional production, mediating 13-40% of the SO BCP. Consideration in models forecasting the SO BCP is recommended considering long-term increases in SO salp abundances.

ecology↗

bioRxiv2022

Quantifying biological carbon pump pathways with a data-constrained mechanistic model ensemble approach

The ability to constrain the mechanisms that transport organic carbon into the deep ocean is complicated by the multiple physical, chemical, and ecological processes that intersect to create, transform, and transport particles in the ocean. In this manuscript we develop and parameterize a data-assimilative model of the multiple pathways of the biological carbon pump (NEMUROBCP). The mechanistic model is designed to represent sinking particle flux, active transport by vertically migrating zooplankton, and passive transport by subduction and vertical mixing, while also explicitly representing multiple biological and chemical properties measured directly in the field (including nutrients, phytoplankton and zooplankton taxa, carbon dioxide and oxygen, nitrogen isotopes, and 234Thorium). Using 30 different data types (including standing stock and rate measurements related to nutrients, phytoplankton, zooplankton, and non-living organic matter) from Lagrangian experiments conducted on 11 cruises from four ocean regions, we conduct an objective statistical parameterization of the model and generate one million different potential parameter sets that are used for ensemble model simulations. The model simulates in situ parameters that were assimilated (net primary production and gravitational particle flux) and parameters that were withheld (234Thorium and nitrogen isotopes) with reasonable accuracy. Model results show that gravitational flux of sinking particles and vertical mixing of organic matter from the surface ocean are more important biological pump pathways than active transport by vertically-migrating zooplankton. However, these processes are regionally variable, with sinking particles most important in oligotrophic areas of the Gulf of Mexico and California, sinking particles and vertical mixing roughly equivalent in productive regions of the CCE and the subtropical front in the Southern Ocean, and active transport an important contributor in the Eastern Tropical Pacific. We further find that mortality at depth is an important component of active transport when mesozooplankton biomasses are high, but that it is negligible in regions with low mesozooplankton biomass. Our results also highlight the high degree of uncertainty, particularly amongst mesozooplankton functional groups, that is derived from uncertainty in model parameters, with important implications from results that rely on non-ensemble model outputs. We also discuss the implications of our results for other data assimilation approaches.

ecology↗

bioRxiv2022

Seasonal variability in carbon:234thorium ratios of suspended and sinking particles in coastal Antarctic waters: Field data and modeling synthesis

238U-234Th disequilibrium is a powerful tool for investigating particle cycling and carbon export associated with the oceans biological carbon pump. However, the interpretation of this method is complicated by multiple processes that can modify carbon:thorium ratios over small spatial scales. We investigated seasonal variability in the thorium and carbon cycles at a coastal site in the Western Antarctic Peninsula. Throughout the ice-free summer season, we quantified carbon and 234Th vertical flux, total water column 234Th, particulate 234Th, and the C:234Th ratios of sinking material and bulk suspended material. Simultaneous identification and separation of fecal pellets from sinking material showed that fecal pellets (primarily from krill) contributed 56% of carbon flux and that as a result of lower C:234Th ratios than suspended particles, these fecal pellets were primary drivers of variability in the C:234Th ratios of sinking material. Bulk suspended particles had highly variable C:234Th ratios and were consistently elevated in the euphotic zone relative to deeper waters. The fraction of 234Th adsorbed onto particles was positively correlated with chlorophyll and particulate organic carbon (POC) concentrations. The C:234Th ratios of suspended particles were positively correlated with POC, although during the spring diatom bloom C:234Th ratios were lower than would have been predicted based on POC concentrations alone. We hypothesize that diatom production of transparent exopolymers may have led to enhanced rates of thorium adsorption during the bloom, thus decreasing the C:234Th ratios. We used a Bayesian model selection approach to develop and parameterize mechanistic models to simulate thorium sorption dynamics. The best model incorporated one slowly-sinking POC pool and rapidly-sinking fecal pellets, with second-order sorption kinetics. The model accurately simulated temporal patterns in the C:234Th ratios of sinking and suspended particles and the fraction of 234Th adsorbed to particles. However, it slightly over-estimated C:234Th ratios during the spring (diatom-dominated) bloom and underestimated C:234Th ratios during the fall (mixed-assemblage) bloom. Optimized model parameters for thorium sorption and desorption were 0.0047 {+/-} 0.0002 m3 mmol C-1 d-1 and 0.017 {+/-} 0.008 d-1, respectively. Our results highlight the important role that specific taxa can play in modifying the C:234Th ratio of sinking and suspended particles and provide guidance for future studies that use 234Th measurements to investigate the functional relationships driving the efficiency of the biological pump. HIGHLIGHTSInvestigated thorium and carbon cycling over full ice-free season C:234Th ratios of sinking particles were controlled by low C:234Th of fecal pellets C:234Th ratios of suspended particles were correlated with chlorophyll and POC Diatom abundance may have led to high particulate thorium during spring bloom Second-order thorium sorption kinetics model accurately simulates C:234Th ratios

ecology↗

bioRxiv2021

Taxon-specific phytoplankton growth, nutrient utilization, and light limitation in the oligotrophic Gulf of Mexico

The highly stratified, oligotrophic regions of the oceans are predominantly nitrogen limited in the surface ocean and light limited at the deep chlorophyll maximum (DCM). Hence, determining light and nitrogen co-limitation patterns for diverse phytoplankton taxa is crucial to understanding marine primary production throughout the euphotic zone. During two cruises in the deep-water Gulf of Mexico, we measured primary productivity (H13CO3-), nitrate uptake (15NO3-), and ammonium uptake (15NH4+) throughout the water column. Primary productivity declined with depth from the mixed-layer to the DCM, averaging 27.1 mmol C m-2 d-1. The fraction of growth supported by NO3- was consistently low, with upper euphotic zone values ranging from 0.01 to 0.14 and lower euphotic zone values ranging from 0.03 to 0.44. Nitrate uptake showed strong diel patterns (maximum during the day), while ammonium uptake exhibited no diel variability. To parameterize taxon-specific phytoplankton nutrient and light utilization, we used a data assimilation approach (Bayesian Markov Chain Monte Carlo) including primary productivity, nutrient uptake, and taxon-specific growth rate measurements. Parameters derived from this analysis define distinct niches for five phytoplankton taxa (Prochlorococcus, Synechococcus, diatoms, dinoflagellates, and prymnesiophytes) and may be useful for constraining biogeochemical models of oligotrophic open-ocean systems.

plant biology↗

bioRxiv2020

Trade-offs between risks of predation and starvation in larvae make the shelf break an optimal spawning location for Atlantic Bluefin tuna

Atlantic Bluefin tuna (ABT) (Thunnus thynnus) travel long distances to spawn in oligotrophic regions of the Gulf of Mexico. To estimate regional larval ABT mortality, we developed a spatially-explicit, Lagrangian, individual-based model that simulates dispersal, growth, and mortality within realistic predator and prey fields during the spawning periods from 1993-2012. Modelled larval ABT experience high mortality in the first week of feeding with an average mortality rate of 0.53 {+/-} 0.26 d-1 prior to postflexion. Survival ranged from 0.12%-0.32% suggesting that recruitment may vary by a factor of 2.7 due to early life stage mortality alone. Starvation is the dominant source of mortality driven by the early critical period; however, survival is ultimately limited by predation on older individuals. As a result, first-feeding larvae survive better in the more food-rich areas on the shelf, while larger larvae survive better in the open ocean with fewer predators, making the shelf break an optimal spawning area. Our findings support the hypothesis that ABT spawn in oligotrophic regions to minimize predation on their larvae. Ocean modeling tools presented in this study may help facilitate an ecosystem-based management approach to improve future stock assessment models by better resolving the stock-recruitment relationship.

ecology↗

bioRxiv2020

Investigating equations for measuring dissolved inorganic nutrient uptake in oligotrophic conditions

Multiple different equations have been used to quantify nutrient uptake rates from stable isotope tracer label incorporation experiments. Each of these equations implicitly assumes an underlying model for phytoplankton nutrient uptake behavior within the incubation bottle and/or pelagic environment. However, the applicability of different equations remains in question and uncertainty arising from subjective choices of which equation to use is never reported. In this study, I use two approaches to investigate the conditions under which different nutrient uptake equations should be used. First, I utilized a moderate-complexity pelagic ecosystem model that explicitly models the {delta}15N values of all model compartments (NEMURO+15N) to conduct simulated nitrate uptake and ammonium uptake incubations and quantify the accuracy of different nutrient uptake equations. Second, I used results of deckboard diel nutrient uptake experiments to quantify the biases of 24-h incubations relative to six consecutive 4-h incubations. Using both approaches, I found that equations that account for nutrient regeneration (i.e., isotope dilution) are more accurate than equations that do not, particularly when nutrient concentrations are low but uptake is relatively high. Furthermore, I find that if the goal is to estimate in situ uptake rates it is appropriate to use an in situ correction to standard equations. I also present complete equations for quantifying uncertainty in nutrient uptake experiments using each nutrient uptake equation and make all of these calculations available as Excel spreadsheets and Matlab scripts.

ecology↗

bioRxiv2020

Plankton food webs of the Gulf of Mexico spawning grounds of Atlantic Bluefin tuna

We used linear inverse ecosystem modeling techniques to assimilate data from extensive Lagrangian field experiments into a mass-balance constrained food web for the Gulf of Mexico open-ocean ecosystem. This region is highly oligotrophic, yet Atlantic Bluefin Tuna (ABT) travel long distances from feeding grounds in the North Atlantic to spawn there. Our results show that the food web is dominated by the microbial loop (>80% of net primary productivity is respired by heterotrophic bacteria and protists that feed on them). In contrast, herbivorous food web pathways from phytoplankton to metazoan zooplankton process <4% of net primary production in the mixed layer. Nevertheless, ABT larvae feed preferentially on calanoid copepods and other suspension-feeding zooplankton that in turn derive much of their nutrition from diatoms and mixotrophic flagellates. This allows ABT larvae to maintain a comparatively low trophic level ([~]4.0 for pre-flexion larvae; [~]4.2 for post-flexion larvae) that increases trophic transfer from phytoplankton to larval fish.

ecology↗

bioRxiv2019

The Importance of Mesozooplankton Diel Vertical Migration for Sustaining a Mesopelagic Food Web

We used extensive ecological and biogeochemical measurements obtained from quasi-Lagrangian experiments during two California Current Ecosystem Long-Term Ecosystem Research cruises to analyze carbon fluxes between the epipelagic and mesopelagic zones using a linear inverse ecosystem model (LIEM). Measurement constraints on the model include 14C primary productivity, dilution-based microzooplankton grazing rates, gut pigment-based mesozooplankton grazing rates (on multiple zooplankton size classes), 234Th:238U disequilibrium and sediment trap measured carbon export, and metabolic requirements of micronekton, zooplankton, and bacteria. A likelihood approach (Markov Chain Monte Carlo) was used to estimate the resulting flow uncertainties from a sample of potential flux networks. Results highlight the importance of mesozooplankton active transport (i.e., diel vertical migration) for supplying the carbon demand of mesopelagic organisms and sequestering carbon dioxide from the atmosphere. In nine water parcels ranging from a coastal bloom to offshore oligotrophic conditions, mesozooplankton active transport accounted for 18% - 84% (median: 42%) of the total carbon supply to the mesopelagic, with gravitational settling of POC (12% - 55%; median: 37%) and subduction (2% - 32%; median: 14%) providing the majority of the remainder. Vertically migrating zooplankton contributed to downward carbon flux through respiration and excretion at depth and via consumption loses to predatory zooplankton and mesopelagic fish (e.g. myctophids and gonostomatids). Sensitivity analyses showed that the results of the LIEM were robust to changes in nekton metabolic demands, rates of bacterial production, and mesozooplankton gross growth efficiency. This analysis suggests that prior estimates of zooplankton active transport based on conservative estimates of standard (rather than active) metabolism should be revisited.\n\nContribution to the FieldUnderstanding the flows of carbon within the ocean is important for predicting how global climate will shift; yet even after decades of research, the magnitude with which the ocean sequesters carbon is highly uncertain. One reason behind this uncertainty is that a variety of mechanisms control the balance between carbon input and carbon output within the ocean. The topic of this work is to inspect the role of biological organisms in physically transferring organic carbon from the surface to the deep ocean. As opposed to other mechanisms--such as sinking particles, the biological transfer of carbon is difficult to measure directly and is often quite variable, leading to large uncertainties. Here we use an extensive set of in situ observations off the coast of southern California to model the flow of carbon through the ecosystem. The model determined that in our study area nearly half of the total transfer of carbon from the surface ocean to deep was carried out by zooplankton that swim up to the surface each night to feed. This finding has direct implications for global carbon budgets, which often underestimate this transfer of carbon.

ecology↗