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.     

Rapid scavenging of jellyfish carcasses reveals the importance of gelatinous material to deep-sea food webs

Jellyfish blooms are common in many oceans, and anthropogenic changes appear to have increased their magnitude in some regions. Although mass falls of jellyfish carcasses have been observed recently at the deep seafloor, the dense necrophage aggregations and rapid consumption rates typical for vertebrate carrion have not been documented. This has led to a paradigm of limited energy transfer to higher trophic levels at jelly falls relative to vertebrate organic falls. We show from baited camera deployments in the Norwegian deep sea that dense aggregations of deep-sea scavengers (more than 1000 animals at peak densities) can rapidly form at jellyfish baits and consume entire jellyfish carcasses in 2.5 h. We also show that scavenging rates on jellyfish are not significantly different from fish carrion of similar mass, and reveal that scavenging communities typical for the NE Atlantic bathyal zone, including the Atlantic hagfish, galatheid crabs, decapod shrimp and lyssianasid amphipods, consume both types of carcasses. These rapid jellyfish carrion consumption rates suggest that the contribution of gelatinous material to organic fluxes may be seriously underestimated in some regions, because jelly falls may disappear much more rapidly than previously thought. Our results also demonstrate that the energy contained in gelatinous carrion can be efficiently incorporated into large numbers of deep-sea scavengers and food webs, lessening the expected impacts (e.g. smothering of the seafloor) of enhanced jellyfish production on deep-sea ecosystems and pelagic–benthic coupling.     

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.     

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.     

Effects of large gut volume in gelatinous zooplankton: ingestion rate, bolus production and food patch utilization by the jellyfish Sarsia tubulosa

Many gelatinous zooplankton consume a large amount of prey andhave stomach volumes much greater than the volume of individualprey. We suggest that jellyfish can use their voluminous stomachas a buffering food-accumulating organ that allows the organismto feed at maximum clearance rate in a wide range of fluctuatingfood concentrations. The food accumulation capability was confirmedfor the hydromedusa Sarsia tubulosa feeding on copepods. Starvedjellyfish feeding in high prey concentrations for 1 h displayedmuch higher average ingestion rates compared with jellyfishfeeding for 20 h or with jellyfish that were pre-adjusted tothe food concentration before incubation. The findings haveimplications for design and interpretation of experiments. Thepossibility for jellyfish to feed at maximum clearance ratein either very high prey concentration for a short time or lowprey concentration for a long time was illustrated with calculationsof prey uptake by S. tubulosa feeding in prey concentrationsof variable heterogeneity. The ability of jellyfish to captureprey at maximum clearance rate under different prey concentrations,and to accumulate relatively large amounts of food in theirguts, suggests that they would thrive in both homogenous andpatchy food distributions. This property may have contributedto the evolutionary and ecological success of the medusoid ‘bauplan’.     

Collection and culture techniques for gelatinous zooplankton

Gelatinous zooplankton are the least understood of all planktonic animal groups. This is partly due to their fragility, which typically precludes the capture of intact specimens with nets or trawls. Specialized tools and techniques have been developed that allow researchers and aquarists to collect intact gelatinous animals at sea and to maintain many of these alive in the laboratory. This paper summarizes the scientific literature on the capture, collection, and culture of gelatinous zooplankton and incorporates many unpublished methods developed at the Monterey Bay Aquarium in the past 15 years.     

Extension of methods for jellyfish and ctenophore trophic ecology to large-scale research

Science has rapidly expanded its frontiers with new technologies in the 20th Century. Oceanography now is studied routinely by satellite. Predictive models are on global scales. At the same time, blooms of jellyfish and ctenophores have become problematic, especially after 1980. Although we have learned a great deal about gelatinous zooplankton ecology in the 20th Century on local scales, we generally have not scaled-up to estimate the extent, the causes, or effects of large blooms. In this age of global science, research on gelatinous zooplankton needs to utilize large-scale approaches and predictive equations. Some current techniques enable jellyfish populations (aerial, towed cameras), feeding (metabolic rates, stable isotopes), and dynamics (predictive modeling) to be studied over large spatial and temporal scales. I use examples of scyphomedusae (Aurelia spp., Cyanea capillata, Chrysaora quinquecirrha) and Mnemiopsis leidyi ctenophores, for which considerable data exist, to explore expanding from local to global scales of jellyfish trophic ecology. Regression analyses showed that feeding rates of Aurelia spp. (FR in copepods eaten medusa⁻¹ d⁻¹) generally could be estimated ±50% from in situ data on medusa wet weight (WW) and copepod density; temperature was not a significant factor. FR of C. capillata and C. quinquecirrha were similar to those of Aurelia spp.; the combined scyphomedusa regression underestimated measured FR of C. quinquecirrha and Aurelia spp. by 50% and 180%, respectively, and overestimated measured FR of C. capillata by 25%. Clearance rates (CR in liters cleared of copepods ctenophore⁻¹ d⁻¹) of M. leidyi were reduced in small containers (≤20 l), and a ratio of container-volume to ctenophore-volume of at least 2,500:1 is recommended for feeding experiments. Clearance rates were significantly related to ctenophore WW, but not to prey density or temperature, and estimated rates within 10–159%. Respiration rates of medusae and ctenophores were similar across habitats with greatly ambient different temperatures (10–30°C), and can be predicted from regressions using only mass. These regressions may permit estimation of feeding effects of gelatinous predators without exhaustive collection of feeding data in situ. I recommend that data on feeding and metabolism of jellyfish and ctenophores be entered in a database to allow generalized predictive relationships to be developed to promote inclusion of these important predators in ecosystem studies and models. Guest editors: K. A. Pitt & J. E. Purcell Jellyfish Blooms: Causes, Consequences, and Recent Advances     

A test of the cross-scale resilience model: Functional richness in Mediterranean-climate ecosystems

Ecological resilience has been proposed to be generated, in part, in the discontinuous structure of complex systems. Environmental discontinuities are reflected in discontinuous, aggregated animal body mass distributions. Diversity of functional groups within body mass aggregations (scales) and redundancy of functional groups across body mass aggregations (scales) has been proposed to increase resilience. We evaluate that proposition by analyzing mammalian and avian communities of Mediterranean-climate ecosystems. We first determined that body mass distributions for each animal community were discontinuous. We then calculated the variance in richness of function across aggregations in each community, and compared observed values with distributions created by 1000 simulations using a null of random distribution of function, with the same n, number of discontinuities and number of functional groups as the observed data. Variance in the richness of functional groups across scales was significantly lower in real communities than in simulations in eight of nine sites. The distribution of function across body mass aggregations in the animal communities we analyzed was non-random, and supports the contentions of the cross-scale resilience model.     

Micro- and macrorheology of jellyfish extracellular matrix

Mechanical properties of the extracellular matrix (ECM) play a key role in tissue organization and morphogenesis. Rheological properties of jellyfish ECM (mesoglea) were measured in vivo at the cellular scale by passive microrheology techniques: microbeads were injected in jellyfish ECM and their Brownian motion was recorded to determine the mechanical properties of the surrounding medium. Microrheology results were compared with macrorheological measurements performed with a shear rheometer on slices of jellyfish mesoglea. We found that the ECM behaved as a viscoelastic gel at the macroscopic scale and as a much softer and heterogeneous viscoelastic structure at the microscopic scale. The fibrous architecture of the mesoglea, as observed by differential interference contrast and scanning electron microscopy, was in accord with these scale-dependent mechanical properties. Furthermore, the evolution of the mechanical properties of the ECM during aging was investigated by measuring microrheological properties at different jellyfish sizes. We measured that the ECM in adult jellyfish was locally stiffer than in juvenile ones. We argue that this stiffening is a consequence of local aggregations of fibers occurring gradually during aging of the jellyfish mesoglea and is enhanced by repetitive muscular contractions of the jellyfish.     

Long-Term Fluctuations in Circalunar Beach Aggregations of the Box Jellyfish Alatina moseri in Hawaii,with Links to Environmental Variability

The box jellyfish Alatina moseri forms monthly aggregations at Waikiki Beach 8–12 days after each full moon, posing a recurrent hazard to swimmers due to painful stings. We present an analysis of long-term (14 years: Jan 1998– Dec 2011) changes in box jellyfish abundance at Waikiki Beach. We tested the relationship of beach counts to climate and biogeochemical variables over time in the North Pacific Sub-tropical Gyre (NPSG). Generalized Additive Models (GAM), Change-Point Analysis (CPA), and General Regression Models (GRM) were used to characterize patterns in box jellyfish arrival at Waikiki Beach 8–12 days following 173 consecutive full moons. Variation in box jellyfish abundance lacked seasonality, but exhibited dramatic differences among months and among years, and followed an oscillating pattern with significant periods of increase (1998–2001; 2006–2011) and decrease (2001–2006). Of three climatic and 12 biogeochemical variables examined, box jellyfish showed a strong, positive relationship with primary production, >2 mm zooplankton biomass, and the North Pacific Gyre Oscillation (NPGO) index. It is clear that that the moon cycle plays a key role in synchronizing timing of the arrival of Alatina moseri medusae to shore. We propose that bottom-up processes, likely initiated by inter-annual regional climatic fluctuations influence primary production, secondary production, and ultimately regulate food availability, and are therefore important in controlling the inter-annual changes in box jellyfish abundance observed at Waikiki Beach.