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     

Polyp and medusa of Dipurena bicircella n. sp. (Hydrozoa: Corynidae) from Northern California

The polyp and medusa of Dipurena bicircella n. sp., a metagenic hydrozoan from Northern California, USA, is described. The arrangement of capitate tentacles of the polyp, in two alternating cycles or circlets, is unique to the genus Dipurena as well as to the family Corynidae. Medusae released in the laboratory attained sexual maturity in 2 weeks. The adult medusa can be distinguished from other medusae of the genus by means of 2 gonad rings, a slight terminal nematocyst cluster on each tentacle, and a relatively small size (about 2 mm in bell height). The polyp was not found in association with sponges, as is the case with some of the other polyps of the genus Dipurena. The medusa is not yet known from the plankton.     

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.     

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     

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.     

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.     

Life cycle of the rhizostome jellyfish <Emphasis Type="Italic">Rhizostoma octopus</Emphasis> (L.) (Scyphozoa,Rhizostomeae), with studies on cnidocysts and statoliths

Stages in the life cycle of the rhizostome jellyfish Rhizostoma octopus (L.) were reared in the laboratory from planula to young medusa and are described here. Fertilization of eggs was external, and planulae, 110–150 μm in length, appeared after 2 days. These settled and developed into the scyphistoma stage. Scyphistomae reached a maximum size of 2.3 mm, bore up to 24 tentacles, and had a large manubrium. Polyps reproduced asexually mainly by podocysts. Strobilation was induced by temperature change. Each strobila released up to five ephyrae that measured 2.7–5.8 mm in diameter at liberation. During transformation from newly released ephyra to young medusa, velar lappets appeared and increased in number, the manubrium developed eight branched oral arms with epaulettes, and a marginal gastric network arose. Only one cnidocyst type, the “a”-atrichous haploneme, was present in the planula. In addition to these haplonemes, heterotrichous microbasic euryteles were observed in polyps, ephyrae, and medusae. Statoliths, located in the rhopalia, had a characteristic compact shape. These sensory structures increased in number and size with the growth of ephyrae and medusae.     

The life cycle of the commensal hydromedusa Eutima sapinhoa n. sp.

Eutima sapinhoa n. sp. is a small hydroid, which lives attached by a basal disc to the mantle, foot, gills and labial palps of the bivalve Tivela mactroides (Born, 1778), found on the northern littoral of São Paulo State, Brazil. This bivalve, locally known as “sapinhoá”, is used as food by the coastal inhabitants. The hydroid produces a leptomedusa which belongs to the family Eutimidae. The medusa bud develops as a small knob-like protuberance near the base of the polyp and, after 61/2 days, reaches the free-swimming stage in covered preparation dishes containing about 200-ml filtered sea water at a temperature of 20° to 22°C. After 25 days the medusa is 4 mm high, 4-mm in diameter, and possesses 4 radial tentacles —each about 2 to 3 times as long as the bell diameter, 8 marginal vesicles — 2 in each quadrant, and 28 small marginal swellings — 7 in each quadrant. The peduncle is as long as the height of the bell, 4 single long gonads develop on the subumbrella on each radial canal. The nematocysts are small atrichous isorhizae, with capsules averaging 7 μm in length.     

Derivation of the reduced life cycle of <Emphasis Type="Italic">Thecoscyphus zibrowii</Emphasis> Werner, 1984 (Cnidaria,Scyphozoa)

The species Thecoscyphus zibrowii Werner, 1984 has an exceptional life cycle, which lacks a medusa stage but develops an extraordinary structure (egg sac) for reproduction. Investigation of the life cycle, as well as anatomical and histological studies of the different developmental stages of T. zibrowii were performed to provide evidence for a possible homology of the egg sac with the medusa stage and to determine whether the reduced metagenesis of T. zibrowii is derived from strobilation. The egg sac showed several characteristics, which were compared to those of coronate medusae. The ectodermis of the egg sac had a plate-like appearance and was completely ciliated as is typical for coronate medusae. The number and the location of the gonads were similar to those of coronate medusae. The cnidocysts were significantly larger in the egg sac than in the polyp. A size difference of cnidocysts in the medusa and the polyp stage is known for several Coronatae. Characteristics of egg sac formation were compared to characteristics of strobilation. The formation of the early operculum was similar in T. zibrowii and N. eumedusoides. The constriction of egg sac and strobila occurred in the same mode and the gastric cavities of two egg sacs stayed in contact in a similar fashion to the gastric cavities of the strobila discs. The developmental zones of cnidoblasts of the egg sac and polyp were separated during the formation of the egg sac which showed a similar developmental gradient to a strobila. The existence of all of these consistent characteristics makes it very likely that the egg sac structure was homologous to a medusa. The species T. zibrowii would therefore be derived from a metagenetic ancestor. This species has reduced the medusa generation to the greatest extent within the Nausithoidae and has demonstrated thus far the endpoint of a regressive evolution of the medusa generation.     

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.     

Comparison of telomere length among different life cycle stages of the jellyfish Cassiopea andromeda

The polyp (scyphistoma) of the jellyfish Cassiopea andromeda reproduces asexually repeatedly, while the medusa, the sexually reproducing stage, exhibits a relatively shorter life span. As a first step to understand the mechanism behind the differences in the life spans of the polyp and medusa stages of the jellyfish, we compared the lengths of the telomere region of one targeted chromosome between the polyp and medusa stages using a modified single telomere length assay (STELA). The double-stranded regions of the telomeres were amplified by PCR, and the average length of the PCR products was estimated by densitometry analysis of the gel smear. Chromosomes within cells of the bell region of the medusa were characterized by longer telomeres than those of polyps, asexual propagules, or other regions of the medusa. This is the first study to estimate the telomere lengths of targeted chromosomes in a cnidarian and opens a way to understand the mechanism underlying different life spans of the polyp and medusa stages.     

The life cycle ofClytia attenuata (calyptoblastea: Campanulariidae)

The life cycle of the hydroidClytia attenuata (Calkins) (Calyptoblastea: Campanulariidae) has been completed in the laboratory including development of the medusa, previously described asPhialidium lomae Torrey (Leptomedusae: Campanulariidae). Under laboratory conditions, the hydroid exhibits some morphological variation. Characteristic branching of the hydroid occurs at temperatures between 17° to 19°C. At 13° to 15°C the colonies are unbranched and cannot be distinguished fromClytia cylindrica L. Agassiz. Young medusae are similar to other young species ofPhialidium. Development to the adult form requires 25 to 30 days at 17° to 24°C. The adult medusae are 6 to 10 mm in diameter, watch-glass shaped, and have 20 to 28 tentacles. Based on the adult medusa,Clytia attenuata is maintained as a valid species.     

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.     

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.     

Determination of δ13C and δ15N and trophic fractionation in jellyfish: implications for food web ecology

Application of stable isotope analysis (SIA) in jellyfish allows definition of trophic patterns not detectable using gut content analysis alone, but analytical protocols require standardization to avoid bias in interpreting isotopic data. We determined δ¹³C and δ¹⁵N in Aurelia sp. from the northern Gulf of Mexico (30°00′N, 89°00′W–30°24′N, 88°00′W) to define differences in stable isotope composition between body parts and whole body, the effect of lipid extraction on δ¹³C in tissues, and fractionation values from medusa to prey. The isotopic composition of bell and whole Aurelia sp. was not different. The increase in δ¹³C values after lipid removal suggested a correction is needed. To aid future analyses, we derived a correction equation from empirical data for jellyfish samples. Laboratory feeding experiments indicated medusae increased +4 ‰ in δ¹³C and +0.1 ‰ in δ¹⁵N compared to their diet. These results suggest protocols commonly applied for other species may be inaccurate to define Aurelia sp. trophic ecology. Because Aurelia spp. are commonly found in marine ecosystems, accurately defining their trophic role by use of SIA has implications for understanding marine food webs worldwide.     

Food availability,intrusion pressure and territory size: an experimental study of Anna's hummingbirds (Calypte anna)

Theoretical considerations implicate food availability and intrusion pressure as important determinants of territory size, but empirical studies have led to contradictory conclusions about cause-and-effect relationships among these three variables. To investigate this problem, we provided patches of electronically controlled artificial flowers, which were defended by male Calypte anna. Food availability was experimentally manipulated, and intrusion rate and territory size were calculated from behavioral observations of the territory owner. Changes in both food availability and intrusion rate were found to be significantly correlated with changes in territory size under certain conditions. Intrusion rate, which was influenced by food availability, was negatively associated with territory size so long as food availability was high. This association persisted even after possible effects of food availability were controlled statistically. Food availability was negatively correlated with territory size only when intrusion rates were high and after owners had been defending territories for 3 days. As food availability and intrusion rate increased, owners increasingly restricted their defense to the patch itself; partial regressions revealed a significant association for intrusion rate but not food availability. When intrusion rate was low and food availability varied from low to high levels, no relationship was observed between food availability and territory size, apparently because of opposing influences of food abundance on territory size. Correspondence to: P.W. Ewald     

Swimming and feeding by the scyphomedusa Chrysaora quinquecirrha

The semaeostome scyphomedusa, Chrysaora quinquecirrha (Desor, 1848), is an abundant and important planktonic predator in estuaries and coastal waters of the eastern USA during the summer. We videotaped free-swimming medusae in the laboratory and in the field in order to determine the relationship between swimming motions and prey encounter with capture surfaces. Medusae were collected from the Choptank River (Chesapeake Bay) in September 1992 and in the Niantic River, Connecticut, USA in July 1994. We used newly hatched Artemia sp. nauplii and fluorescein dye to trace water motions around swimming medusae. Swimming results in a pulsed series of toroids which travel along the medusan oral arms and tentacles. Prey are entrained in this flow and the location of naupliar encounter was influenced by the phase of the pulsation cycle during which entrainment occurred. Flow-field velocities, measured by tracking particles adjacent to the bell margin during contraction, increased with bell diameter. Received: 29 March 1997 / Accepted: 11 April 1997     

Prey, feeding rates, and asexual reproduction rates of the introduced oligohaline hydrozoan Moerisia lyonsi

Moerisia lyonsi Boulenger (Hydrozoa) medusae and benthic polyps were found at 0 to 5‰ salinity in the Choptank River subestuary of Chesapeake Bay, USA. This species was introduced to the bay at least 30 years before 1996. Medusae and polyps of M. lyonsi are very small and inconspicuous, and may occur widely, but unnoticed, in oligohaline waters of the Chesapeake Bay system and in other estuaries. Medusae consumed copepod nauplii and adults, but not barnacle nauplii, polychaete and ctenophore larvae or tintinnids, in laboratory experiments. Predation rates on copepods by medusae increased with increasing medusa diameter and prey densities. Feeding rates on copepod nauplii were higher than on adults and showed no saturation over the range of prey densities tested (1 to 64 prey l⁻¹). By contrast, predation on copepod adults was maximum (1 copepod medusa⁻¹ h⁻¹) at 32 and 64 copepods l⁻¹. Unexpectedly, M. lyonsi colonized mesocosms at the Horn Point Laboratory during the spring and summer in 4 years (1994 to 1997), and reached extremely high densities (up to 13.6 medusae l⁻¹). Densities of copepod adults and nauplii were low when medusa densities were high, and estimated predation effects suggested that M. lyonsi predation limited copepod populations in the mesocosms. Polyps of M. lyonsi asexually produced both polyp buds and medusae. Rates of asexual reproduction increased with increasing prey availability, from an average total during a 38 d experiment of 9.5 buds polyp⁻¹ when each polyp was fed 1 copepod d⁻¹, to an average total of 146.7 buds polyp⁻¹ when fed 8 copepods d⁻¹. The maximum daily production measured was 8 polyp buds and 22 medusae polyp⁻¹. The colonizing potential of this hydrozoan is great, given the high rates of asexual reproduction, fairly wide salinity tolerance, and existence of a cyst stage. Received: 29 October 1998 / Accepted: 3 March 1999     

Life cycle of the jellyfish <Emphasis Type="Italic">Rhizostoma pulmo</Emphasis> (Scyphozoa: Rhizostomeae) and its distribution,seasonality and inter-annual variability along the Catalan coast and the Mar Menor (Spain,NW Mediterranean)

Rhizostoma pulmo is one of the most abundant scyphomedusae along the Mediterranean coasts. To understand changes in the population densities of the medusa stage and its relationship with the benthic stage, we describe all developmental stages from the life cycle of R. pulmo, from the scyphistoma stage to young medusae reared in the laboratory. Mature scyphistomae showed a mean total body length of 1.7 mm and the average tentacle number was 16. Asexual reproduction occurred by lateral budding, podocyst and pedalocyst production. Each strobila released up to 8 ephyrae that had a mean total body diameter of 3.5 mm. Moreover, we also present data on the temporal and spatial distribution of the species in the Catalan Sea and the coastal lagoon on Mar Menor (NW Mediterranean) during the years 2007–2009. In the Catalan Sea, the presence of adult R. pulmo was recorded as extended along the whole coast, but most of the observation was concentrated in the central area; the highest abundances were recorded during the months of July and August. The highest number of observations was detected in 2008 in coincidence with the inter-annual variation observed in the Mar Menor lagoon.     

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.