Abundance,distribution and prey composition of scyphomedusae in the southern North Sea

The annual cycle of abundance and distribution of the scyphozoan medusae Aurelia aurita, Cyanea lamarckii, C. capillata and Chrysaora hysoscella were studied in the southern North Sea in 2004 and 2005. Three different patterns of seasonal occurrence of medusae were distinguished: (1) the early occurring C. lamarckii (February–August), (2) C. capillata and A. aurita (April–August) and (3) the late appearing C. hysoscella (July/August–September). Cyanea lamarckii was the most frequently encountered species in this study; its highest mean abundance was 1.8 ± 2.7 ind. 100 m⁻³. The prey spectra of C. lamarckii, C. capillata and C. hysoscella contained several copepod and other crustacean species and thus make them potential competitors with fish larvae. Medusae in this study also consumed fish eggs and larvae, including clupeids, in all months analysed. Although peak spawning of sprat (Sprattus sprattus) coincides with the maximum abundance of medusae (May–June) the relative low abundance of all medusae species in this study makes jellyfish predation unlikely to be a factor controlling sprat recruitment in the time frame investigated.     

A comparative ecological study on the scyphozoans Aurelia aurita,Cyanea capillata and C. lamarckii in the Gullmar Fjord,western Sweden, 1982 to 1986

During the years 1982 to 1986, the life cycles and population dynamics of three scyphozoans, Aurelia aurita (L.), Cyanea capillata (L.) and C. lamarckii (Person and Lesueur), were studied in the Gullmar Fjord on the Swedish west coast. The settling of planulae, strobilation of scyphistomae and release of ephyrae were followed on ceramic settling plates in the laboratory and in the field. Weekly to bi-weekly hauls with Bongo nets were used to study the abundance of ephyrae and medusae. The results show great differences in the life cycles and ecology of the three species. A. aurita utilizes the best season for scyphistoma growth (August to September) and strobilates during the highest zooplankton abundance in October. C. capillata strobilates during the spring (March to May), and the abundance of C. capillata medusae is more dependent on immigration from the North Sea than A. aurita. C. lamarckii does not reproduce at all in the Gullmar Fjord and is totally dependent on immigration from the North Sea. The possibility of interspecific competition between A. aurita and C. capillata is discussed. A preliminary experiment showed that scyphistomae of A. aurita eat planula larvae of C. capillata during the autumn.     

Morphology,fluid motion and predation by the scyphomedusa Aurelia aurita

Although medusan predators play demonstrably important roles in a variety of marine ecosystems, the mechanics of prey capture and, hence, prey selection, have remained poorly defined. A review of the literature describing the commonly studied medusa Aurelia aurita (Linnaeus 1758) reveals no distinct patterns of prey selectivity and suggests that A. aurita is a generalist and feeds unselectively upon available zooplankton. We examined the mechanics of prey capture by A. aurita using video methods to record body and fluid motions. Medusae were collected between February and June in 1990 and 1991 from Woods Hole, Massachusetts and Narragansett Bay, Rhode Island, USA. Tentaculate A. aurita create fluid motions during swimming which entrain prey and bring them into contact with tentacles. We suggest that this mechanism dominates prey selection by A. aurita. In this case, we predict that medusae of a specific diameter will positively select prey with escape speeds slower than the flow velocities at their bell margins. Negatively selected prey escape faster than the medusan flow velocity draws them to capture surfaces. Faster prey will be captured by larger medusac because flow field velocity is a function of bell diameter. On the basis of prey escape velocities and flow field velocities of A. aurita with diameters of 0.8 to 7.1 cm, we predict that A. aurita will select zooplankton such as barnacle nauplii and some slow swimming hydromedusae, while faster copepods will be negatively selected.     

Prey selection by the scyphomedusan predator Aurelia aurita

We describe feeding behavior of Aurelia aurita (Linnaeus) using gut content analyses of field-collected specimens and a mesocosm experiment. The field studies were conducted in Narragansett Bay, Rhode Island, USA from March to April 1988, and the mesocosm studies were done at the Marine Ecosystems Research Laboratory at the University of Rhode Island. Patterns of prey selection changed with medusa diameter. Smaller medusae (12 mm diameter) consumed mostly hydromedusan prey whereas larger medusae (up to 30 mm diameter) ingested greater numbers of copepod prey. While larger medusae did feed on copepods, their diet also contained more barnacle nauplii and hydromedusae than expected from the relative abundances of these prey types in plankton samples. A marginal flow mechanism of feeding by A. aurita provided an explanation for the patterns of prey selection we observed in medusae of different sizes and among widely divergent prey types. Our data indicated that large prey, with escape speeds slower than the marginal flow velocities around the bell margins of A. aurita, made up a substantial fraction of the daily ration when they were available. Such prey species may be more important to nutrition than the more abundant copepods and microzooplankton. Successful development of young medusae may depend upon an adequate supply of slowly escaping prey.     

Laboratory study of predation by Aurelia aurita on larvae of cod,flounder, plaice and herring: development and vulnerability to capture

Capture success of the medusa Aurelia aurita preying on various developmental stages of fish larvae was measured together with larval reactivity and escape speed after being stung. These experiments were conducted in the spring of 1983 with A. aurita medusae collected from Loch Etive, Scotland and laboratory-reared larvae of Gadus morhua L., Platichthys flesus L., Pleuronectes platessa L. and Clupea harengus L. Capture success of the medusae increased with medusa size, but decreased with advancing larval development. Smaller species of larvae were more vulnerable to capture. Larval reactivity to encounters with medusae increased with advancing development, and larger species of larvae were more reactive to encounters. Larval escape swimming speeds also increased with advancing larval development and size. These results indicate that earlier stages of larvae within a species and smaller species of larvae at a given stage are more vulnerable to predation by medusae since they are less reactive to encounters. Apparently they are more susceptible to the effects of neurotoxins. Predation rates on different developmental stages of herring larvae are documented and compared with rates predicted by a predation model. Predictions fell within the range of observed predation rates, but tended to overestimate rates by larger medusae feeding on larger herring larvae. This indicates the possibility of predator satiation and/or behavioural avoidance.     

A laboratory study of predation by Aurelia aurita on larval herring (Clupea harengus): Experimental observations compared with model predictions

Predation by the medusa Aurelia aurita L. on early first-feeding stage larvae of the herring clupea harengus L. was studied in the laboratory. The medusae were captured in Loch Etive, Scotland. Herring larvae were reared from the extificially fertilized eggs of spawning Clyde herring caught in March, 1982. Swimming speeds, volume searched”, capture efficiency and predation rates increased as medusa size increased. Predation rates on fish larvae increased with prey density, but appeared to approach a maximum at high prey densities; in 1 h experiments, a maximum rate of predation of 6.64 larvae h^-1 was estimated by fitting an Ivlev function. A model to predict predation rates was constructed from swimming speeds, sizes and densities of medusae and larvae, and capture efficiency. The rates of predation predicted from the model fell within the range of experimental data, but tended to underestimate rates and did not account for saturation of medusae. Swimming patterns of medusae changed after prey capture: (a) before capture, encounter rates were low and medusae were relatively less active; (b) after capture of 1 larva, encounter rates doubled, with the stimulated medusae exhibiting increased activity and an aftered “searching” path; and (c) after capture of many larvae, swimming speeds and encounter rates of medusae decreased.     

Ultrastructural observations on prey capture and digestion in the scyphomedusa Aurelia aurita

Aurelia aurita medusae are able to catch their prey with their entire body surface. Catch efficiency in medusae caught in Kiel harbour in May 1985 was found to be highest at the tentacles and lowest at the subumbrella. Surface structures of the medusa as well as the cnidom are described by SEM observations. Microbasic heterotrichous euryteles and atrichous isorhizas were found. Discharged nematocysts on the prey's skin indicate different functions of the two types. The villi in the gastral cavity show a characteristic morphological differentiation that consists of a ciliated distal and a basal area covered by vesicles. Four types of glandular cells were identified by TEM observations. Mucous cell types preferably occur in densely ciliated areas. The presence of serous cells is restricted to the basal region of the gastral villi and gastral cavity where the extracellular predigestion takes place. The time of food passage in young medusae of A. aurita decreases from 19 h at 4°C to 4 h at 22°C.     

Release of dissolved organic carbon (DOC) by the scyphozoan jellyfish Aurelia aurita and its potential influence on the production of planktic bacteria

A series of incubation experiments were made to measure the rate of release of dissolved organic carbon (DOC) by the jellyfish Aurelia aurita (L.) (collected in 1991–1992 in Gullmarsfjorden, Sweden). Release of DOC by medusae (9.5 to 18 cm in diameter) from the Skagerrak ranged from 0.70 to 1.6 mg C ind^-1 d^-1 with a mean of 1.2 mg C ind^-1 d^-1 (SD=0.29 mg C ind^-1 d^-1, n=10). Based on data from the literature for two medusa populations, this can be equivalent to 2.5 and 7.1% of the carbon assimilated in one season (June to September). This was similar to the amount of carbon allocated to reproduction. Bacterial abundance was monitored to evaluate the possible stimulating effect of the DOC released. Bacterial growth was stimulated by the presence of A. aurita. The importance of A. aurita as a source of DOC on a large scale is small compared to exudate from primary producers. The volume surrounding each medusa, wherein the DOC released may be enough to sustain a bacterial production such as the one reported from the Swedish west coast, is equivalent to a sphere with a diameter of 0.5 m. Thus, considering the patchy distribution of medusae, the local influence of their DOC release could be important.     

Sexual reproduction of the scyphomedusa Aurelia aurita in relation to temperature and variable food supply

The effects of food availability and temperature on sexual maturation and female reproductive output of the scyphomedusa Aurelia aurita was examined in two populations from the contrasting environments of Southampton Water and Horsea Lake, England. Trends in oogenesis and subsequent reproductive output differed markedly between the two populations. In Southampton Water, the onset of sexual maturation occurred earliest in the larger medusae, but eventually all females became ripe, the smallest being 45 mm bell diameter (BD). The decrease in minimum size at maturity was correlated with increasing temperature. In A. aurita from Horsea Lake, size at maturity varied on a seasonal basis, with the smallest ripe female being only 19 to 20 mm BD. There were spring and autumn periods of sexual maturation in this population. During the autumn period, it is likely that food limitation was playing a more critical role in determining medusa size, with decreasing temperature indirectly affecting A. aurita by limiting primary and secondary production. In similar-sized ripe medusae, fecundity was greater in Southampton Water, but the planula larvae produced were significantly smaller than those in Horsea Lake. It is suggested that in Horsea Lake, the quality of the larvae are greater in terms of biochemical content to ensure survival of the few gametes produced (i.e. K-strategy). Comparison of the reproductive effort of the two A. aurita populations revealed that medusae from Southampton Water, which experience greater food availability, are able to direct more energy to reproduction than Horsea Lake medusae. In the latter, A. aurita medusae appear to partition the available food resources into either somatic growth (and therefore increased future fecundity) when food is abundant, or reproductive growth when food is scarce. Received: 24 June 1997 / Accepted: 23 March 1998     

Intraguild predatory interactions between the jellyfish <Emphasis Type="Italic">Cyanea capillata</Emphasis> and <Emphasis Type="Italic">Aurelia aurita</Emphasis>

While qualitative observations of jellyfish intraguild predation abound in the literature, there are only few rate measurements of these interactions. We quantified predation rates among two common jellyfish in northern boreal waters, Cyanea capillata and its prey Aurelia aurita, both of which also feed on crustacean zooplankton and fish larvae. A series of incubation experiments using a wide range of prey concentrations (0.38–3.8 m⁻³) in large containers (2.6 m³) was carried out. By replenishing the prey continuously as they were captured we maintained a nearly constant prey concentrations. Ingestion rates increased linearly up to prey concentrations of 1.92 m⁻³, yielding maximum clearance rates of ∼2.37 ± 0.39 m³ predator⁻¹ h⁻¹ for C. capillata predators 16 ± 2.3 cm in diameter. Mean ingestion rate at saturated prey concentrations (1.92–3.85 m⁻³) was 4.01 ± 0.78 prey predator⁻¹ h⁻¹. Behavioral observations suggested that predators did not alter their swimming behavior during meals, and thus that feeding rates were generally handling limited rather than encounter limited. Predators captured more prey than needed, and semi-digested prey was often discarded when fresh prey was encountered.     

An individual-based numerical model of medusa swimming behavior

Scyphomedusae are ubiquitous in marine and estuarine systems, where they frequently play an important role in trophodynamics. Many scyphomedusae are cruising predators, and feeding rates depend, in part, on swimming behavior. Yet, no model of medusa swimming exists. An individual-based correlated random walk (CRW) model of medusa swimming behavior in three dimensions was developed. The model was validated using a previously published dataset of the swimming of 19 Chrysaora quinquecirrha (Desor, 1848) medusae that were observed in the presence or absence of zooplankton prey in laboratory mesocosms in August–October 1998 (Matanoski et al. in Mar Biol 139:191–200, 2001). In the presence of prey, medusae swam at a constant moderate rate in looping trajectories. In the absence of prey, medusae alternated periods of slow and fast swimming in more linear trajectories. In the model, looping trajectories were reproduced only when changes in movement by a medusa were oriented to its current position and orientation; more linear trajectories were reproduced by movement oriented to a fixed framework. This suggests that medusae change from swimming behavior oriented to local stimuli (e.g., contact with prey) to long-range stimuli (e.g., gravity) depending on the availability of prey. The model reproduced cyclical changes in swimming speeds by medusae in the absence of prey by simulating switching in the behavior controlling the strength of swimming bell pulsations using a probabilistic function. Model results also demonstrated that medusae tend to swim toward the surface, avoid contact with the bottom, increase time spent in prey patches if they alter swimming patterns in the presence of prey, and exhibit significant periodicities in swimming patterns that are the result of deterministic behavior. The model will permit the simulation of the complex behavior of medusae.     

Occurrence and public perception of jellyfish along the German Baltic coastline

Jellyfish accumulations are often problematic for tourism and fisheries both along European coastlines and overseas. They damage the reputation of several seaside resorts and cause serious economic losses. On the German Baltic coast, jellyfish accumulations are well known, too. Besides population increase due to anthropogenic effects, normal hydrodynamic processes are notable key factors when it comes to jellyfish accumulations. Along German Baltic coastlines, moon jellyfish (Aurelia aurita) occur most frequently but also stinging lion??s mane jellyfish (Cyanea capillata) appear regularly, however not as abundant. The objectives of our study were a) to screen when, where, how many and which jellyfish species occur on German Baltic coasts, b) to find the causes for jellyfish accumulations and c) to evaluate how beach visitors perceive native medusae and whether information can influence their perception. Different methods where combined: systematic jellyfish observations in cooperation with lifeguards, investigations of jellyfish abundance in shallow water zones, and interviews with beach visitors. Our results suggest that jellyfish occurrence along the German Baltic coast correlates with offshore wind or shore-parallel wind, which cause upwelling events. In contrast, there is no evidence that frequency of jellyfish occurrence was dependent on water temperature in summer. In regard to tourism, we found that beach visitors who received information about jellyfish stated to feel significantly less bothered by medusae. Overall, this article addresses different methods to learn more about jellyfish accumulations and it shows information strategies in terms of improved beach management.     

Enhanced survival of 0-group gadoid fish under jellyfish umbrellas

Young (0-group) gadoid fish, which have been observed sheltering beneath jellyfish (Scyphozoa and Hydrozoa) umbrellas, may find refugia from predation by retreating among medusan tentacles. The survival of juvenile gadoids may therefore be improved by high abundances of medusae. Jellyfish (including Cyanea lamarckii and C. capillata) were caught in the North Sea during routine sampling for 0-group gadoids (cod Gadus morhua, haddock Melanogrammus aeglefinus, Norway pout Trisopterus esmarkii and whiting Merlangius merlangus) between 1971 and 1986, and considerable overlaps have been shown between the spatial distributions of these fish and medusae. Here correlations are made between the abundance of medusae in the North Sea and the residual survival of 0-group fish, as quantified by the deviation in 1-group recruitment from the expected Ricker modelled estimate. Significant positive correlations between the residual survival of whiting and medusa abundance are evident for each individual Cyanea sp. and for the combined Cyanea spp. ln(maximum) abundance (all R ≥ 0.60, P < 0.01, N = 15). The abundance of jellyfish may thus be an important factor influencing the mortality of whiting in the North Sea, and as such should be considered in the development of ‘ecosystem-based’ management of whiting stocks.     

The common jellyfish Aurelia aurita: standing stock,excretion and nutrient regeneration in the Kiel Bight,Western Baltic

The population dynamics, ammonia and inorganic phosphate excretion, and nutrient regeneration of the common jellyfish Aurelia aurita was investigated from 1982 to 1984 in the Kiel Bight, western Baltic Sea. During summer 1982, medusae abundance ranged between 14 and 23 individuals 100 m^-3, biomass was estimated at about 5 g C 100 m^-3 and the mean final diameter of individuals was 22 cm. Abundance, based on numbers, in 1983 and 1984 was an order of magnitude lower; biomass was less than 2 g C 100 m^-3 and jellyfish grew to 30 cm. During the summers of 1983 and 1984, A. aurita biomass constituted roughly 40% of that of the total zooplankton>200 m. In 1982, for which zooplankton data were lacking, it was assumed that medusae biomass was greater than that of all other zooplankton groups. Total ammonia excretion ranged between 6.5 and 36 mol h^-1 individual^-1, whereas inorganic phosphate release was 1.4 to 5.7 mol h^-1 individual^-1. Allometric equations were calculated and exponents of 0.93 for NH₄–N release and 0.87 for PO₄–P excretion were determined. Nitrogen and phosphorus turnover rates were 5.4 and 14.6% d^-1, respectively. In 1982, the medusae population released 1 100 mol NH₄–N m^-2 d^-1, about 11% of the nitrogen requirements of the phytoplankton. The inorganic phosphate excretion (150 mol m^-2 d^-1) sustained 23% of the nutrient demands of the primary producers. In the other two years the nutrient cycling of the medusae was much less important, and satisfied only 3 to 6% of the nutrient demands. It is suggested that in some years A. aurita is the second most important source of regenerated nutrients in Kiel Bight, next to sediment.     

Morphology and development of benthic and pelagic life stages of North Sea jellyfish (Scyphozoa, Cnidaria) with special emphasis on the identification of ephyra stages

Jellyfish blooms or invasions could be detected in an early phase of development if the youngest medusa stages (ephyrae) and their early growth stages (post-ephyrae) were identifiable in plankton samples but a useful identification key for ephyrae in early growth stages is lacking for most species. In the present study, the identification characteristics of adult North Sea scyphomedusae (Aurelia aurita, Cyanea capillata, Cyanea lamarckii, Chrysaora hysoscella, Rhizostoma octopus) collected around the island of Helgoland (German Bight) in July?CAugust 2003 and 2004 are described. Planula larvae were measured and reared to polyps in the laboratory. The process of ephyrae development asexually produced by the polyps (strobilation) was photo-documented. Photographs of the ephyrae growth stages were combined with drawings of features useful for the species identification. The provided identification key allows discrimination among post-ephyrae from plankton samples, probably leading to conclusions on the development of jellyfish blooms and their causes.     

Quantitative feeding ecology of the hydromedusan Nemopsis bachei in Chesapeake Bay

We determined feeding rates of the hydromedusan Nemopsis bachei L. Agassiz in the mesohaline region of Chesapeake Bay, USA during the spring of 1989 and 1990 from gut contents, digestion rates and abundances of medusae and zooplankton. The medusae consumed primarily copepodites of Acartia tonsa, selecting against naupliar stages. The peak abundance of N. bachei medusae was in April to May, when densities averaged more than 10 m^-3. Medusa densities were similar in both years, but were greatest (maximum of 132 medusae m^-3) along a southern transect sampled only in 1990. At peak densities, N. bachei medusae consumed 30% d^-1 of the copepodite standing stocks, but they consumed <1% d^-1 at the lower densities typical of late May or early June. The predation effects were generally greater than those reported for other hydromedusan species. But even at peak predation, N. bachei medusae could not have controlled or reduced A. tonsa copepod populations, which had a production rate of 85% d^-1 at that time. Medusa feeding rates were highest at nighttime, and were correlated with prey density in the field, but not in the laboratory.Communicated by J. Grassle, New Brunswick     

Trophic ecology and the related functional morphology of the deepwater medusa <Emphasis Type="Italic">Periphylla periphylla</Emphasis> (Scyphozoa,Coronata)

Remotely operated vehicle (ROV)-based field studies on the distribution and behaviour of Periphylla periphylla Péron and Lesueur (Ann Mus Hist Nat Marseille 14:316–366, 1809), from three Norwegian fjords have been combined with on-board experiments and morphological and histological studies in order to understand the trophic ecology of this species. Field studies from one of the fjords showed that the zooplankton biomass was negatively related with P. periphylla abundance, indicating a predatory effect. The majority of zooplankton biomass tended to be distributed above the aggregation of P. periphylla, which in turn showed highest abundance at 100–200 m depth. Observation on the orientation of medusae passing the ROV when descending down in the water column at dawn and dusk, showed no consistency with the theory of diel vertical migration. Estimated metabolic demand of P. periphylla indicated a daily predation impact on the prey assemblage of 13% as an average for the fjord. In situ behavioural observations showed that the dominant tentacle posture of large medusae was straight upward, with tentacles extended to the oral–aboral body axis. The hunting mode alternates between ambush and ramming, whereby tentacle posture minimises the water turbulence that may otherwise alarm the prey. The musculature of the tentacles is well developed, with an especially strong longitudinal muscle on the oral side, facilitating fast movement of the tentacle towards the mouth. In addition, ring-, radial-, and diagonal musculatures are also present. The diagonal is probably most important for the corkscrew retraction of the tentacle, used at the moment of prey capture. Results from laboratory experiments show that different body-parts of P. periphylla vary in sensitivity for chemical and mechanical stimuli, including hydrodynamic disturbance and vibration in the surrounding water. Feeding success is facilitated by combining the vibration-sense on the tentacle tips and the marginal lappets, the touch-sense on the tentacle bases and marginal lappets, and a taste control of the captured prey at the mouthlips.     

Feeding rates of the jellyfish Aurelia aurita on fish larvae

We quantified feeding rates of field caught Aurelia aurita feeding on yolk sac cod (Gadus morhua) larvae in a series of incubation experiments. A short-time (~1 h) functional response experiment with a wide range of prey concentrations (0.5–16 prey l⁻¹, initial concentration) revealed that ingestion rates increased linearly over this range, such that clearance rates were similar between the different prey concentrations. This suggests that A. aurita is capable of efficiently utilizing dense prey patches. This indication was further supported by a linear increase of prey captured by A. aurita during 2.5 h of feeding at extremely high prey concentration (>200 prey l⁻¹). Clearance rate in darkness scaled with jellyfish diameter to a power of ~1.7 for jellyfish 3.9–9.5 cm in diameter. The jellyfish did not alter their umbrella pulse frequency in response to presence of fish larvae. There were no significant differences between A. aurita feeding rates in light and darkness for yolk sac prey ages 0–7 days (at 7.5°C). Although prey vision and escape abilities of fish may develop rapidly during early larval ontogeny, these factors apparently have little impact on interactions with predators such as A. aurita during the yolk sac stage.     

Diversity of swimming behavior in hydromedusae as related to feeding and utilization of space

Feeding behaviors of the following 4 species of hydromedusae are described from field and laboratory observations: Probosidactyla flavicirrata, Stomotoca atra, Phialidium gregarium and Polyorchis penicillatus. Feeding efficiency of medusae has previously been considered equivalent to fishing with a given amount (combined tentacle length) of adhesive fishing line; however, detailed observation shows that behavior of medusae greatly modifies the fishing capacity of each species. It is hypothesized that in addition to (1) tentacle number and length, the following factors strongly influence feeding efficiency: (2) tentacle posture, (3) velocity of tentacles moving through water (4) swimming pattern of medusa, (5) streamlining effects of medusa bell on water flow, (6) diameter of prey, (7) swimming pattern and velocity of prey. Each species of hydromedusa utilizes the above factors in different combinations.Mailing address