Gelatinous zooplankton (ctenophores,salps and medusae): an important food resource of fishes in the temperate SW Atlantic Ocean

This study quantifies the occurrence of gelatinous zooplankton in the stomach contents of fishes from the southwest Atlantic Ocean (33°–55°S). More than 69,000 fish stomachs belonging to 107 species were examined. A total of 39 fishes were documented as consumers of gelatinous zooplankton, 23 of which were newly discovered. Three gelatinous organism consumption categories are recognized: (1) very frequent consumers (10 species, six of which were exclusive); frequent consumers (five species); and occasional consumers (26 species). Three types of gelatinous prey (ctenophores, salps and medusae) were found in the stomach contents of fishes. Ctenophores were consumed at high levels across almost the entire continental shelves of Argentina and Uruguay. Salps were frequent prey on the slope and southern shelf. In contrast, medusae were consumed in coastal areas, slopes and the southern shelf. Classification methods (group average sorting of the Bray–Curtis similarity measures based on log (X?+?1)-transformed percentage data) determined six areas where fishes predated on gelatinous organisms. SIMPER (similarity percentages) analysis determined which fishes contributed more to the consumption of gelatinous organisms. Results revealed that two fish species (Stromateus brasiliensis and Squalus acanthias) had high gelatinous zooplankton predation rates throughout the entire study area, while another six species (Patagonotothen ramsayi, Helicolenus dactylopterus lahillei, Macrourus holotrachys, Merluccius hubbsi, Schroederichthys bivius, and Macruronus magellanicus), while widely distributed, seemed to have specific areas where consumption occurred. This study not only provides new knowledge about the importance of gelatinous zooplankton in the diet of numerous fishes, but might also be valuable for planning and managing local fisheries.     

A physical context for gelatinous zooplankton aggregations: a review

The magnitude and extent of jellyfish blooms are influenced not only by the biology and behavior of the animal, but also by the geographic setting and physical environment. Hydrography alone is often thought to cause or favor gelatinous zooplankton aggregations, however, it is clear that interactions between biology of the animal and physics of the water are very important sources of population variations, especially at local scales. We summarize the role of physical processes and phenomena that promote aggregations of gelatinous zooplankton. We have identified and discussed a suite of physical gradients that can be perceived by gelatinous zooplankton. These include light, gravity, temperature, salinity, pressure and turbulence. A recurring theme is accumulation of jellyfish around physical discontinuities such as fronts (shelf-break, upwelling, tidal and estuarine) and pycnoclines (thermoclines and haloclines). Interestingly, there are few data to suggest that large-scale, quasi-stationary features, such as the largest oceanic fronts, serve to physically aggregate gelatinous animals at a similar scale. Rather, examples of local aggregations appear to dominate the literature. We also discuss various jellyfish behaviors that are theorized to promote aggregation, feeding and reproduction in relation to physical discontinuities.     

Resource Requirements of the Pacific Leatherback Turtle Population

The Pacific population of leatherback sea turtles (Dermochelys coriacea) has drastically declined in the last 25 years. This decline has been linked to incidental capture by fisheries, egg and meat harvesting, and recently, to climate variability and resource limitation. Here we couple growth rates with feeding experiments and food intake functions to estimate daily energy requirements of leatherbacks throughout their development. We then estimate mortality rates from available data, enabling us to raise food intake (energy requirements) of the individual to the population level. We place energy requirements in context of available resources (i.e., gelatinous zooplankton abundance). Estimated consumption rates suggest that a single leatherback will eat upward of 1000 metric tonnes (t) of jellyfish in its lifetime (range 924–1112) with the Pacific population consuming 2.1×10⁶ t of jellyfish annually (range 1.0–3.7×10⁶) equivalent to 4.2×10⁸ megajoules (MJ) (range 2.0–7.4×10⁸). Model estimates suggest 2–7 yr-old juveniles comprise the majority of the Pacific leatherback population biomass and account for most of the jellyfish consumption (1.1×10⁶ t of jellyfish or 2.2×10⁸ MJ per year). Leatherbacks are large gelatinous zooplanktivores with consumption to biomass ratios of 96 (up to 192 if feeding strictly on low energy density Cnidarians); they, therefore, have a large capacity to impact gelatinous zooplankton landscapes. Understanding the leatherback''s needs for gelatinous zooplankton, versus the availability of these resources, can help us better assess population trends and the influence of climate induced resource limitations to reproductive output.     

A comparison of carbon-specific respiration rates in gelatinous and non-gelatinous zooplankton: A search for general rules in zooplankton metabolism

Using 470 data from the literature the dry weight-specific respiration rates of gelatinous zooplankton (cnidarians, ctenophores and salps) and non-gelatinous zooplankton (mainly crustacea) were converted to carbon-specific values. The resulting carbon-specific respiration rates showed no significant differences between the two groups of zooplankton, indicating similar oxygen requirements per gram of carbon biomass. From this finding, it can be suggested that the differences in the rates of oxygen consumption measured in the two types of zooplankton in the sea can be explained by the carbon biomass ratio between gelatinous and non-gelatinous zooplankton. Furthermore, the low rate of metabolism of gelatinous species compared with that of non-gelatinous animals of the same volume can be attributed predominantly to the relatively low organic matter content in the former. It is recommended that all weight-specific metabolism rates be expressed using carbon as body mass unit (e.g. mg O₂ gC⁻¹ d⁻¹) which enables more accurate comparisons between individuals exhibiting different dry weight/carbon ratios.     

Gelatinous zooplankton of the Arctic Ocean: in situ observations under the ice

The gelatinous zooplankton of the Canada Basin were observed with a deep-diving remotely operated vehicle (ROV) during August–September, 2002. Taxa observed fell into four main groups: cnidarians, ctenophores, chaetognaths, and pelagic tunicates. We provide detailed data on the vertical distributions of many taxa from three sites which span the Canada Basin. The most common gelatinous organisms in the surface waters were the ctenophores Mertensia ovum and Bolinopsis infundibulum. These two species were found in very large numbers in the near-surface mixed layer. In the mesopelagic zone, below the transition from the Pacific water layer and the Atlantic water layer, the most common species was Sminthea arctica. Surprising numbers of the scyphomedusa Atolla tenella were found in the deep waters of the basin, along with an undescribed species of narcomedusae. The vertical distributions of the gelatinous zooplankton observed with the ROV show several trends related to the physical properties of the water and geographic location within the basin.     

New records of Amphipoda Hyperiidea in associations with gelatinous zooplankton

Associations between Amphipoda Hyperiidea specimens and gelatinous zooplankton such as cnidarians, ctenophores or salps are known in the literature. It is well established that many hyperiids are intimately associated with gelatinous zooplankton during part or all of their life-cycles, and that the associations could be of commensal or of parasitic nature, sometimes very highly species-specific. Although conventional net sampling usually does not preserve evidence of these associations, 12 instances of association (I.A.) were found in nine of the 49 samples collected during the 'FINEP XI Cruise', carried out on the Southeast coast of Brazil (23–28 °S), in January 1980. Lestrigonus bengalensisGiles, 1887 (11 specimens) and Simorhynchotus antennarius(Claus, 1871) (one specimen) were found inside salps and siphonophores. The study of the nature of these associations is essential to the understanding of the evolution of the group.     

Filter-feeding gelatinous macrozooplankton response to climate change and implications for benthic food supply and global carbon cycle

It is often suggested that gelatinous zooplankton may benefit from anthropogenic pressures of all kinds and in particular from climate change. Large pelagic tunicates, for example, are likely to be favored over other types of macrozooplankton due to their filter-feeding mode, which gives them access to small preys thought to be less affected by climate change than larger preys. In this study, we provide model-based estimate of potential community changes in macrozooplankton composition and estimate for the first time their effects on benthic food supply and on the ocean carbon cycle under two 21st-century climate-change scenarios. Forced with output from an Earth System Model climate projections, our ocean biogeochemical model simulates a large reduction in macrozooplankton biomass in response to anthropogenic climate change, but shows that gelatinous macrozooplankton are less affected than nongelatinous macrozooplankton, with global biomass declines estimated at −2.8% and −3.5%, respectively, for every 1°C of warming. The inclusion of gelatinous macrozooplankon in our ocean biogeochemical model has a limited effect on anthropogenic carbon uptake in the 21st century, but impacts the projected decline in particulate organic matter fluxes in the deep ocean. In subtropical oligotrophic gyres, where gelatinous zooplankton dominate macrozooplankton, the decline in the amount of organic matter reaching the seafloor is reduced by a factor of 2 when gelatinous macrozooplankton are considered (−17.5% vs. −29.7% when gelatinous macrozooplankton are not considered, all for 2100 under RCP8.5). The shift to gelatinous macrozooplankton in the future ocean therefore buffers the decline in deep carbon fluxes and should be taken into account when assessing potential changes in deep carbon storage and the risks that deep ecosystems may face when confronted with a decline in their food source.     

A golden age of gelata: past and future research on planktonic ctenophores and cnidarians

Hydrobiologia - The study of the natural history of gelatinous zooplankton (‘gelata’) reached a high point at the end of the 19th century, when scientists first began to understand the...     

Associations between gelatinous zooplankton and hyperiid amphipods (Crustacea: Peracarida) in the Gulf of California

Hyperiid amphipods are pelagic crustaceans that live associated with gelatinous zooplankton including medusae, ctenophores, siphonophores, and salps. Standard plankton sampling disrupts natural associations, so the most reliable way to determine an association is through direct observation of the organisms in their environment. The planktonic fauna of the Gulf of California dwelling between 10 and 3000 m was surveyed using SCUBA diving and a remotely operated submersible (ROV) during March 2003. Here we report our observations on a total of 14 symbiotic associations found between the hyperiid amphipods and various taxa of gelatinous zooplankton. We found parental care behavior in a group of amphipods (Oxycephalidae) in which this phenomenon has not been previously reported. For two hyperiid species, Euthamneus rostratus and Vibilia australis, we present the first information on their symbiotic relations. Additional hosts were discovered for other well-known and widely distributed hyperiid species (i.e. Brachyscelus crusculum, Hyperoche medusarum). Photographic evidence of some of these interactions is included in this contribution. This is the first survey of these relationships in the Gulf of California, and many aspects of the ecology and biology of these symbioses remain to be studied.     

High occurrence of jellyfish predation by black‐browed and Campbell albatross identified by DNA metabarcoding

Gelatinous zooplankton are a large component of the animal biomass in all marine environments, but are considered to be uncommon in the diet of most marine top predators. However, the diets of key predator groups like seabirds have conventionally been assessed from stomach content analyses, which cannot detect most gelatinous prey. As marine top predators are used to identify changes in the overall species composition of marine ecosystems, such biases in dietary assessment may impact our detection of important ecosystem regime shifts. We investigated albatross diet using DNA metabarcoding of scats to assess the prevalence of gelatinous zooplankton consumption by two albatross species, one of which is used as an indicator species for ecosystem monitoring. Black‐browed and Campbell albatross scats were collected from eight breeding colonies covering the circumpolar range of these birds over two consecutive breeding seasons. Fish was the main dietary item at most sites; however, cnidarian DNA, primarily from scyphozoan jellyfish, was present in 42% of samples overall and up to 80% of samples at some sites. Jellyfish was detected during all breeding stages and consumed by adults and chicks. Trawl fishery catches of jellyfish near the Falkland Islands indicate a similar frequency of jellyfish occurrence in albatross diets in years of high and low jellyfish availability, suggesting jellyfish consumption may be selective rather than opportunistic. Warmer oceans and overfishing of finfish are predicted to favour jellyfish population increases, and we demonstrate here that dietary DNA metabarcoding enables measurements of the contribution of gelatinous zooplankton to the diet of marine predators.     

Associations between gelatinous zooplankton and hyperiid amphipods (Crustacea: Peracarida) in the Gulf of California

Hydrobiologia - Hyperiid amphipods are pelagic crustaceans that live associated with gelatinous zooplankton including medusae, ctenophores, siphonophores, and salps. Standard plankton sampling...     

Selective and differential feeding on marine prokaryotes by mucous mesh feeders

Microbial mortality impacts the structure of food webs, carbon flow, and the interactions that create dynamic patterns of abundance across gradients in space and time in diverse ecosystems. In the oceans, estimates of microbial mortality by viruses, protists, and small zooplankton do not account fully for observations of loss, suggesting the existence of underappreciated mortality sources. We examined how ubiquitous mucous mesh feeders (i.e. gelatinous zooplankton) could contribute to microbial mortality in the open ocean. We coupled capture of live animals by blue-water diving to sequence-based approaches to measure the enrichment and selectivity of feeding by two coexisting mucous grazer taxa (pteropods and salps) on numerically dominant marine prokaryotes. We show that mucous mesh grazers consume a variety of marine prokaryotes and select between coexisting lineages and similar cell sizes. We show that Prochlorococcus may evade filtration more than other cells and that planktonic archaea are consumed by macrozooplanktonic grazers. Discovery of these feeding relationships identifies a new source of mortality for Earth's dominant marine microbes and alters our understanding of how top-down processes shape microbial community and function.     

Massive consumption of gelatinous plankton by Mediterranean apex predators

Stable isotopes of carbon and nitrogen were used to test the hypothesis that stomach content analysis has systematically overlooked the consumption of gelatinous zooplankton by pelagic mesopredators and apex predators. The results strongly supported a major role of gelatinous plankton in the diet of bluefin tuna (Thunnus thynnus), little tunny (Euthynnus alletteratus), spearfish (Tetrapturus belone) and swordfish (Xiphias gladius). Loggerhead sea turtles (Caretta caretta) in the oceanic stage and ocean sunfish (Mola mola) also primarily relied on gelatinous zooplankton. In contrast, stable isotope ratios ruled out any relevant consumption of gelatinous plankton by bluefish (Pomatomus saltatrix), blue shark (Prionace glauca), leerfish (Lichia amia), bonito (Sarda sarda), striped dolphin (Stenella caerueloalba) and loggerhead sea turtles (Caretta caretta) in the neritic stage, all of which primarily relied on fish and squid. Fin whales (Balaenoptera physalus) were confirmed as crustacean consumers. The ratios of stable isotopes in albacore (Thunnus alalunga), amberjack (Seriola dumerili), blue butterfish (Stromaeus fiatola), bullet tuna (Auxis rochei), dolphinfish (Coryphaena hyppurus), horse mackerel (Trachurus trachurus), mackerel (Scomber scombrus) and pompano (Trachinotus ovatus) were consistent with mixed diets revealed by stomach content analysis, including nekton and crustaceans, but the consumption of gelatinous plankton could not be ruled out completely. In conclusion, the jellyvorous guild in the Mediterranean integrates two specialists (ocean sunfish and loggerhead sea turtles in the oceanic stage) and several opportunists (bluefin tuna, little tunny, spearfish, swordfish and, perhaps, blue butterfish), most of them with shrinking populations due to overfishing.     

Chaoborus americanus predation on various zooplankters; Functional response and behavioral observations

Summary Predation rates for Chaoborus americanus on different types of noncyclomorphic zooplankton prey were determined in the laboratory as a function of both prey species and density. The sequence of events leading to consumption of prey was also observed, and probabilities determined for the various components.Predation rates generally reached an upper limit as the density of prey increased to high levels. Larger prey were always less vulnerable to Chaoborus predation. Among the species of zooplankton prey examined Holopedium gibberum suffered the lowest rates of predation, while those of the other plankters (Daphnia pulex, D. rosea, Moina hutchinsoni, Diaptomus birgei, D. leptopus, Cyclops vernalis) were approximately equivalent for any given size category.Observation of the predatory sequence indicates that for all the plankters except H. gibberum there were relatively high rates of evasion of attack by C. americanus and low rates of escape after capture. For H. gibberum the situation was reversed, and escape after capture was particularly important in determining the predation rate.The gelatinous envelope of Holopedium gibberum seems to operate as an effective anti-predator device which reduces the likelyhood of retention by Chaoborus after contact has been made.     

Stable isotopes challenge the perception of ocean sunfish Mola mola as obligate jellyfish predators

Evidence is provided from stable isotope analysis that aggregations of small ocean sunfish Mola mola (total length <1 m) feed broadly within coastal food webs and their classification as obligate predators of gelatinous zooplankton requires revision.     

Evidence for Changes in Estuarine Zooplankton Fostered by Increased Climate Variance

Estuaries are among the most valuable aquatic systems in terms of their services to human welfare. They offer an ideal framework to assess multiscale processes linking climate and food web dynamics through the hydrological cycle. Resolving food web responses to climate change is fundamental to resilience management of these threatened ecosystems under global change scenarios. Here, we examined the temporal variability of the plankton food web in the Mondego Estuary, central Iberian Peninsula, over the period 2003 to 2012. The results pointed out a cascading effect from climate to plankton communities that follow a non-stationary behavior shaped by the climate variance envelope. Concurrent changes in hydrographic processes at the regional, that is, upwelling intensity, and local, that is, estuarine hydrology, scales were driven by climatic forcing promoted by the North Atlantic Oscillation; the influence of which permeated the physical environment in the estuary affecting both autotrophic and heterotrophic communities. The most conspicuous change arose around 2008 and consisted of an obvious decrease in freshwater taxa along with a noticeable increase in marine organisms, mainly driven by gelatinous zooplankton. The observed increase in small-sized cosmopolitan copepods, that is, Clausocalanus arcuicornis, Oithona plumifera, thermophilic species, that is, Penilia avirostris, and gelatinous zooplankton suggests a structural change in the Mondego plankton community. These results provide empirical support to the expectation that expanding climate variance changes plankton structure and functioning, likely fostering trophic interactions in pelagic food webs.     

The influence of turbidity on growth and survival of fish larvae: a numerical analysis

Growth and survival through the early larval phase probably limit the production potential of many commercially important fish stocks. Attempts to increase the production of these stocks by restocking of juveniles have generally failed. Here, we analyse how enhanced concentrations of phytoplankton and zooplankton affect the survival of fish larvae during their early life stages. The analysis is developed for larvae feeding on copepod eggs and nauplii, with fish and invertebrates as major predators. A model of feeding and growth of fish larvae is applied to assess the benefit of enhanced phytoplankton and zooplankton abundance. The analysis shows that the shading effect of higher phytoplankton concentration may reduce predation rates on fish larvae substantially. This `top-down' effect may be more important for the cohort survivorship than the `bottom-up' mechanism in situations when larval food is sufficiently abundant. However, while increased algal biomass will improve recruitment at high zooplankton concentration, it may also reduce recruitment at low zooplankton concentrations and shallow mixing depths. Both the larvae and their vertebrate predators are dependent on light to detect their prey, and the longer reactive distance of the predators make them more susceptible than the larvae to reduced light levels and increased turbidity. We discuss the implications of reduced predation and increased zooplankton abundance on recruitment and production of fish larvae, and point at environmental conditions where changing algal biomass is likely to affect recruitment success.     

Vampire squid: detritivores in the oxygen minimum zone

Vampire squid (Vampyroteuthis infernalis) are considered phylogenetic relics with cephalopod features of both octopods and squids. They lack feeding tentacles, but in addition to their eight arms, they have two retractile filaments, the exact functions of which have puzzled scientists for years. We present the results of investigations on the feeding ecology and behaviour of Vampyroteuthis, which include extensive in situ, deep-sea video recordings from MBARI''s remotely operated vehicles (ROVs), laboratory feeding experiments, diet studies and morphological examinations of the retractile filaments, the arm suckers and cirri. Vampire squid were found to feed on detrital matter of various sizes, from small particles to larger marine aggregates. Ingested items included the remains of gelatinous zooplankton, discarded larvacean houses, crustacean remains, diatoms and faecal pellets. Both ROV observations and laboratory experiments led to the conclusion that vampire squid use their retractile filaments for the capture of food, supporting the hypothesis that the filaments are homologous to cephalopod arms. Vampyroteuthis'' feeding behaviour is unlike any other cephalopod, and reveals a unique adaptation that allows these animals to spend most of their life at depths where oxygen concentrations are very low, but where predators are few and typical cephalopod food is scarce.