skip to main content
US FlagAn official website of the United States government
dot gov icon
Official websites use .gov
A .gov website belongs to an official government organization in the United States.
https lock icon
Secure .gov websites use HTTPS
A lock ( lock ) or https:// means you've safely connected to the .gov website. Share sensitive information only on official, secure websites.


Title: Vertical trophic structure and niche partitioning of gelatinous predators in a pelagic food web: Insights from stable isotopes of siphonophores
Abstract Gelatinous zooplankton are increasingly recognized as key components of pelagic ecosystems, and there have been many recent insights into their ecology and roles in food webs. To examine the trophic ecology of siphonophores (Cnidaria, Hydrozoa), we used bulk (carbon and nitrogen) and compound‐specific (nitrogen) isotope analysis of individual amino acids (CSIA‐AA). We collected samples of 15 siphonophore genera using blue‐water diving, midwater trawls, and remotely operated vehicles in the California Current Ecosystem, from 0 to 3000 m. We examined the basal resources supporting siphonophore nutrition by comparing their isotope values to those of contemporaneously collected sinking and suspended particles (0–500 m). Stable isotope values provided novel insights into siphonophore trophic ecology, indicating considerable niche overlap between calycophoran and physonect siphonophores. However, there were clear relationships between siphonophore trophic positions and phylogeny, and the highest siphonophore trophic positions were restricted to physonects. Bulk and source amino acid nitrogen isotope (δ15N) values of siphonophores and suspended particles all increased significantly with increasing collection depth. In contrast, siphonophore trophic positions did not increase with increasing collection depth. This suggests that microbially reworked, deep, suspended particles with higher δ15N values than surface particles, likely indirectly support deep‐pelagic siphonophores. Siphonophores feed upon a range of prey, from small crustaceans to fishes, and we show that their measured trophic positions reflect this trophic diversity, spanning 1.5 trophic levels (range 2.4–4.0). Further, we demonstrate that CSIA‐AA can elucidate the feeding ecology of gelatinous zooplankton and distinguish between nutritional resources across vertical habitats. These findings improve our understanding of the functional roles of gelatinous zooplankton and energy flow through pelagic food webs.  more » « less
Award ID(s):
2048210 1829812
PAR ID:
10573373
Author(s) / Creator(s):
; ; ; ; ; ; ;
Publisher / Repository:
Wiley Periodicals LLC
Date Published:
Journal Name:
Limnology and Oceanography
Volume:
69
Issue:
4
ISSN:
0024-3590
Page Range / eLocation ID:
902-919
Format(s):
Medium: X
Sponsoring Org:
National Science Foundation
More Like this
  1. Abstract Eukaryotic microalgae play critical roles in the structure and function of marine food webs. The contribution of microalgae to food webs can be tracked using compound‐specific isotope analysis of amino acids (CSIA‐AA). Previous CSIA‐AA studies have defined eukaryotic microalgae as a single functional group in food web mixing models, despite their vast taxonomic and ecological diversity. Using controlled cultures, this work characterizes the amino acidδ13C (δ13CAA) fingerprints—a multivariate metric of amino acid carbon isotope values—of four major groups of eukaryotic microalgae: diatoms, dinoflagellates, raphidophytes, and prasinophytes. We found excellent separation of essential amino acidδ13C (δ13CEAA) fingerprints among four microalgal groups (mean posterior probability reclassification of 99.2 ± 2.9%). We also quantified temperature effects, a primary driver of microalgal bulk carbon isotope variability, on the fidelity ofδ13CAAfingerprints. A 10°C range in temperature conditions did not have significant impacts on variance inδ13CAAvalues or the diagnostic microalgalδ13CEAAfingerprints. Theseδ13CEAAfingerprints were used to identify primary producers at the base of food webs supporting consumers in two contrasting systems: (1) penguins feeding in a diatom‐based food web and (2) mixotrophic corals receiving amino acids directly from autotrophic endosymbiotic dinoflagellates and indirectly from water column diatoms, prasinophytes, and cyanobacteria, likely via heterotrophic feeding on zooplankton. The increased taxonomic specificity of CSIA‐AA fingerprints developed here will greatly improve future efforts to reconstruct the contribution of diverse eukaryotic microalgae to the sources and cycling of organic matter in food web dynamics and biogeochemical cycling studies. 
    more » « less
  2. Abstract Zooplankton contribute a major component of the vertical flux of particulate organic matter to the ocean interior by packaging consumed food and waste into large, dense fecal pellets that sink quickly. Existing methods for quantifying the contribution of fecal pellets to particulate organic matter use either visual identification or lipid biomarkers, but these methods may exclude fecal material that is not morphologically distinct, or may include zooplankton carcasses in addition to fecal pellets. Based on results from seven pairs of wild‐caught zooplankton and their fecal pellets, we assess the ability of compound‐specific isotope analysis of amino acids (CSIA‐AA) to chemically distinguish fecal pellets as an end‐member material within particulate organic matter. Nitrogen CSIA‐AA is an improvement on previous uses of bulk stable isotope ratios, which cannot distinguish between differences in baseline isotope ratios and fractionation due to metabolic processing. We suggest that the relative trophic position of zooplankton and their fecal pellets, as calculated using CSIA‐AA, can provide a metric for estimating the dietary absorption efficiency of zooplankton. Using this metric, the zooplankton examined here had widely ranging dietary absorption efficiencies, where lower dietary absorption may equate to higher proportions of fecal packaging of undigested material. The nitrogen isotope ratios of threonine and alanine statistically distinguished the zooplankton fecal pellets from literature‐derived examples of phytoplankton, zooplankton biomass, and microbially degraded organic matter. We suggest that δ15N values of threonine and alanine could be used in mixing models to quantify the contribution of fecal pellets to particulate organic matter. 
    more » « less
  3. Abstract Gelatinous zooplankton play a crucial role in pelagic marine food webs, however, due to methodological challenges and persistent misconceptions of their importance, the trophic role of gelatinous zooplankton remains poorly investigated. This is particularly true for small gelatinous zooplankton including the marine pelagic tunicate,Dolioletta gegenbauri.D. gegenbauriand other doliolid species occur persistently on wide subtropical shelves where they often produce massive blooms in association with shelf upwelling conditions. As efficient filter feeders and prodigious producers of relatively low‐density organic‐rich aggregates, doliolids are understood to contribute significantly to shelf production, pelagic ecology, and pelagic–benthic coupling. Utilizing molecular gut content analysis and stable isotope analysis approaches, the trophic interactions of doliolids were explored during bloom and non‐bloom conditions on the South Atlantic Bight continental shelf in the Western North Atlantic. Based on molecular gut content analysis, relative ingestion selectivity varied withD. gegenbaurilife stage. At all life stages, doliolids ingested a wide range of prey types and sizes, but exhibited selectivity for larger prey types including diatoms, ciliates, and metazoans. Experimental growth studies confirmed that metazoan prey were ingested, but indicated that they were not digested and assimilated. Stable isotopic composition (δ13C and δ15N) of wild‐caught doliolids, during bloom and non‐bloom conditions, were most consistent with a detrital‐supplemented diet. These observations suggest that the feeding ecology ofD. gegenbauriis more complex than previously reported, and have strong and unusual linkages to the microbial food web. 
    more » « less
  4. Abstract Compound‐specific stable isotope analysis of individual amino acids (CSIA‐AA) has emerged as a transformative approach to estimate consumer trophic positions (TPCSIA) that are internally indexed to primary producer nitrogen isotope baselines. Central to accurate TPCSIAestimation is an understanding of beta (β) values—the differences between trophic and source AA δ15N values in the primary producers at the base of a consumers’ food web. Growing evidence suggests higher taxonomic and tissue‐specificβvalue variability than typically appreciated.This meta‐analysis fulfils a pressing need to comprehensively evaluate relevant sources ofβvalue variability and its contribution to TPCSIAuncertainty. We first synthesized all published primary producer AA δ15N data to investigate ecologically relevant sources of variability (e.g. taxonomy, tissue type, habitat type, mode of photosynthesis). We then reviewed the biogeochemical mechanisms underpinning AA δ15N andβvalue variability. Lastly, we evaluated the sensitivity of TPCSIAestimates to uncertainty in meanβGlx‐Phevalues and Glx‐Phe trophic discrimination factors (TDFGlx‐Phe).We show that variation inβGlx‐Phevalues is two times greater than previously considered, with degree of vascularization, not habitat type (terrestrial vs. aquatic), providing the greatest source of variability (vascular autotroph = −6.6 ± 3.4‰; non‐vascular autotroph = +3.3 ± 1.8‰). Within vascular plants, tissue type secondarily contributed toβGlx‐Phevalue variability, but we found no clear distinction among C3, C4and CAM plantβGlx‐Phevalues. Notably, we found that vascular plantβGlx‐Lysvalues (+2.5 ± 1.6‰) are considerably less variable thanβGlx‐Phevalues, making Lys a useful AA tracer of primary production sources in terrestrial systems. Our multi‐trophic level sensitivity analyses demonstrate that TPCSIAestimates are highly sensitive to changes in bothβGlx‐Pheand TDFGlx‐Phevalues but that the relative influence ofβvalues dissipates at higher trophic levels.Our results highlight that primary producerβvalues are integral to accurate trophic position estimation. We outline four key recommendations for identifying, constraining and accounting forβvalue variability to improve TPCSIAestimation accuracy and precision moving forward. We must ultimately expand libraries of primary producer AA δ15N values to better understand the mechanistic drivers ofβvalue variation. 
    more » « less
  5. Midwater zooplankton are major agents of biogeochemical transformation in the open ocean; however their characteristics and activity remain poorly known. Here we evaluate midwater zooplankton biomass, amino acid (AA)-specific stable isotope composition (δ15N values) using compound-specific isotope analysis of amino acids (CSIA-AA), trophic position, and elemental composition in the North Pacific Subtropical Gyre (NPSG). We focus on zooplankton collected in the winter, spring, and summer to evaluate midwater trophic dynamics over a seasonal cycle. For the first time we find that midwater zooplankton respond strongly to seasonal changes in production and export in the NPSG. In summer, when export from the euphotic zone is elevated and this ‘summer pulse’ material is transported rapidly to depth, CSIA-AA indicates that large particles (> 53 μm) dominate the food web base for zooplankton throughout the midwaters, and to a large extent even into the upper bathypelagic zone. In winter, when export is low, zooplankton in the mid-mesopelagic zone continue to rely on large particle basal resources, but resident zooplankton in the lower mesopelagic and upper bathypelagic zones switch to include smaller particles (0.7–53 μm) in their food web base, or even a subset of the small particle pool. Midwater zooplankton migration patterns also vary with season, with migrants distributed more evenly at night through the euphotic zone in summer as compared to being more compressed in the upper mixed layer in winter. Deeper zooplankton migration within the mesopelagic zone is also reduced in late summer, likely due to the increased magnitude of large particle material available at depth during this season. Our observed seasonal change in activity and trophic dynamics drives modestly greater biomass in summer than winter through the mesopelagic zone. In contrast midwater zooplankton carbon (C), nitrogen (N), and phosphorus (P) composition does not change with season. Instead we find increasing C:N, C:P, and N:P ratios with greater depths, likely due to decreases in proteinaceous structures and organic P compounds and increases in storage lipids with depth. Our study highlights the importance and diversity of feeding strategies for small zooplankton in NPSG midwaters. Many small zooplankton, such as oncaeid and oithonid copepods, are able to access small particle resources at depth and may be an important trophic link between the microbial loop and deep dwelling micronekton species that also rely on small particle-based food webs. Our observed midwater zooplankton trophic response to export-driven variation in the particle field at depth has important implications for midwater metabolism and the export of C to the deep sea. 
    more » « less