Life cycle of <Emphasis Type="Italic">Carybdea marsupialis</Emphasis> Linnaeus, 1758 (Cubozoa,Carybdeidae) reveals metamorphosis to be a modified strobilation

Two types of metamorphosis occur in the life cycle of Carybdea marsupialis Linnaeus 1758 (Cubozoa, Carybdeidae). In addition to the metamorphosis described several times over, during which the entire polyp transforms into one medusa, we investigated a second type of metamorphosis which leaves a regenerative remnant. This mode of medusa formation seems to be derived from the strobilation known from Scyphozoa. Both types are described and the implications these results have on the idea about the origin of the class Cubozoa, are discussed.     

Rate-temperature responses in scyphozoan medusae and polyps

The effects of temperature on oxygen consumption and spontaneous rhythmic activity have been investigated in various stages of the life histories of 3 species of jellyfish from the Chesapeake Bay, USA. All 3 species clearly show the ability to acclimate positively to temperature change. Thermal sensitivity of metabolism in the winter medusa Cyanea capillata fulva is fairly low at temperature intervals which are experienced in nature. Polyps of the two summer medusae, Chrysaora quinquecirrha and Aurelia aurita, show reduced metabolic sensitivity at temperatures normally accompanying high developmental activity and the onset of strobilation.     

The morphogenesis of ephyra in Coronatae (Cnidaria,Scyphozoa)

The morphogenesis of ephyrae of Atorella vanhoeffeni Bigelow, 1909 (Werner, 1967) and of Nausithoe maculata Jarms, 1990 during strobilation is described. We found differences in the developmental pattern of marginal structures and structural changes of longitudinal muscle tubes in particular. During strobilation the polyp’s tetraradiate symmetry is passed to the ephyra of both taxa as a tetraradiate central symmetry that we consider a major plesiomorphic character. The present results also indicate divergent patterns of ephyra development during strobilation that lead to variable marginal symmetries of ephyrae and thus of medusae. Therefore, the marginal symmetry of medusae of N. maculata is octoradiate and of A. vanhoeffeni is hexaradiate. We conclude the latter is stated as derived pattern. These results lead us to maintain both families Nausithoidae and Atorellidae.     

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.     

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.     

Life cycle ofRhopilema nomadica: a new immigrant scyphomedusan in the Mediterranean

Rhopilema nomadica is an Indopacific scyphomedusan, which has migrated into the eastern Mediterranean in recent years. Large aggregations of the medusae were recorded in Haifa Bay, Israel, reaching 5.5×10⁵ medusae per square nautical mile during summer 1989. The life cycle ofR. nomadica from planula to young medusa is described. Fertilization is external and planulae are formed within a few hours at 20°C. After settlement, polyps were fed withArtemia sp. nauplii and developed into polydisc strobilae within 45 d. The strobilation process was completed within 7 d, and the liberated ephyrae developed into young medusae within 2 mo. Asexual reproduction occurred mainly via podocyst formation. The population explosion ofR. nomadica could be attributed to its high reproductive potential.     

Life cycle of the jellyfish <Emphasis Type="Italic">Lychnorhiza lucerna</Emphasis> (Scyphozoa: Rhizostomeae)

The life cycle of Lychnorhiza lucerna (Scyphozoa: Rhizostomeae) and the settlement preferences of its larvae were studied using laboratory-based rearing experiments. Mature medusae of L. lucerna were collected from the beach of the Río de la Plata estuary, Argentina. This species displayed the typical metagenetic, (i.e. medusoid and polypoid), life cycle reported for other rhizostomes. The fertilized eggs developed into motile and short lived planulae. The majority of planulae settled on the air-water interface (p < 0.001). Of those that settled on the settlement plates provided, no significant differences were observed between styrene slides, glass slides and shells of the bivalve Mactra isabelleana (p > 0.05). No planulae settled on stones. Several hours after planulae settled, they metamorphosed into sessile four-tentacled scyphistomae. Most scyphistomae attached onto the air-water interface. At 19–22°C, the scyphistomae grew up to 22 tentacles and reached 1,500 μm height. The scyphistomae increased their numbers by means of formation of podocysts from which new polyps emerged and strobilated. Strobilation occurred 46 days after settlement. Only polydisk strobilation was observed and each strobila always produced three ephyrae. After releasing ephyrae, strobilae returned to normal scyphistomae and were capable of repeating strobilation. A single founder polyp was estimated to produce up to 60 ephyrae over 4 months. Ephyrae developed into metephyrae 15 days after release at 19–22°C. In this paper we describe the morphological and some behavioural features of L. lucerna in the polypoid and early medusoid stages.     

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.     

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.     

Development and biology of <Emphasis Type="Italic">Periphylla</Emphasis><Emphasis Type="Italic">periphylla</Emphasis> (Scyphozoa: Coronatae) in a Norwegian fjord

The globally distributed coronate scyphomedusa Periphylla periphylla (Peron and Lesueur, 1809) occurs in permanent and extremely high abundance in some Norwegian fjords. Studies on the species in Lurefjorden, 40 km north of Bergen, have revealed a holopelagic life cycle with direct development. We distinguished 14 successive developmental stages, characterised by size and morphology. Eggs and early stages are non-mobile, neutrally buoyant, and found mainly at intermediate depths. Rearing studies indicated a development time of 2-3 months from fertilisation to stage 9, when the medusa starts feeding and becomes motile. The fjord population usually shows a strong diel vertical migration, with aggregations at shallower depths during the night. The development includes a gradual increase in pigmentation (porphyrins), starting with the stomach, and thereafter extending to tentacles and the whole exumbrella. This pigment is photodegraded by natural light. Rearing experiments in the laboratory have shown a lethal effect of light. Here, development stops at stage 5. Pigments become visible in stage 7. Older medusae are severely damaged and killed within a few days by exposure to daylight. All stages of P. periphylla are bioluminescent, with this capacity increasing with growth, probably in connection with the development of the nervous net. The light reaction starts at the point of stimulation and spreads at two different speeds corresponding to the two nerve nets of the medusae. The coronal furrow has a central function in transmission of the reaction. It is hypothesised that the bioluminescence is used mainly as a warning to some potential predators, signalling that the porphyrin-containing medusa is unpalatable. P. periphylla has the same two categories of nematocysts as other coronates, with a total of six different types. Nematocyst abundance, distribution and morphology indicate their function. The giant euryteles (capsule up to 100 µm and tubule up to 1160 µm) are the largest ones known among Scyphozoa, and are unique in that this size increases during medusa development.     

Invertebrate introductions in marine habitats: two species of hydromedusae (Cnidaria) native to the Black Sea,Maeotias inexspectata and Blackfordia virginica,invade San Francisco Bay

The hydrozoans Maeotias inexspectata Ostroumoff, 1896 and Blackfordia virginica Mayer, 1910, believed to be native to the Black Sea (i.e. Sarmatic) and resident in a variety of estuarine habitats worldwide, were found as introduced species in the Petaluma River and Napa River, California, in 1992 and 1993. These rivers are mostly-estuarine tributaries that flow into north San Francisco Bay. Both species appeared to be well-established in this brackishwater habitat. Salinities at the collection sites were about 11 during the summer, rising to nearly 20 in the early autumn and falling to near 0 in the winter. Large numbers of all sizes of both species of medusae were observed and collected, indicating that the hydroid stages of the life cycles of the two are also well-established in these rivers. In the Petaluma River, populations of both species were at maximum in late July, with numbers of individuals declining through August and into September; the Napa River was sampled only in October, and at that time only B. virginica was found. Examination of full guts of M. inexspectata and B. virginica medusae revealed that both species had fed nearly exclusively on small crustaceans, principally barnacle nauplii, copepods and their eggs and nauplii, and crab zoea larvae (M. inexspectata only). All the M. inexspectata medusae were males, indicating that the population has probably developed from the introduction of perhaps only a single male polyp or polyp bud. In spite of its inability to reproduce sexually, this population appears to be maintained by the prodigious ability of the polyp to bud and reproduce asexually, and is fully capable of invading additional low-salinity habitats from its present Petaluma River site. Male and female B. virginica medusae were collected in both the Petaluma River and the Napa River, indicating that B. virginica may have been introduced by either the polyp or medusa stage (or both), but that multiple individuals (of both sexes) must have arrived from another port in one or more invasions. As indicated for M. inexspectata, the B. virginica population will also probably seed new populations in San Francisco Bay and elsewhere. Based on its cnidome as well as the morphology of both medusa and polyp, M. inexspectata has been reclassified by moving it from the family Olindiidae, Limnomedusae, to the family Moerisiidae, Anthomedusae.     

Swimming and feeding in Periphylla periphylla (Scyphozoa, Coronatae)

Some western Norwegian fjords host extraordinarily abundant and persistent populations of the mesopelagic, coronate scyphomedusa, Periphylla periphylla. In these environments, from late autumn to spring, the medusae undertake regular diel vertical migrations into surface waters. From unique observations obtained with a remotely operated vehicle (ROV), including observations made without artificial light, we observed that 90% of the medusae swam with their tentacles in aboral position. Stomach content analyses of surface-collected specimens revealed that the medusae ate mainly calanoid copepods, but ostracods and large euphausiids were also prominent components of their diets. The clearance rate potential of P. periphylla, assessed from in situ observations and stomach contents, was comparable to that of similar-sized, epipelagic gelatinous species. Our findings suggest that P. periphylla behave as active predators in surface waters.     

Laboratory observations on the life history of Rhopilema verrilli (Scyphozoa: Rhizostomeae)

The life history of the scyphozoan Rhopilema verrilli is described from the planula to the young medusa stages. Planulae are retained within the gonadal tissue of the medusa until fully developed. On liberation, most planulae set and developed into small scyphistomae within 7 to 10 days. The scyphistoma differs from those of other described species in having an unusually large, clavate manubrium. The only means of asexual reproduction observed in the scyphistoma cultures involved the formation of podocysts. Strobilation was usually of the monodisc variety, although polydics strobilation was not infrequent. The process of strobilation was completed within 7 days at 20°C. Newly liberated ephyrae typically had 8 pairs of lappets, and 8 rhopalia. Ephyral development resembled that of the closely related rhizostome Rhizostoma pulmo. The cnidome of the planula and scyphistoma consisted of atrichous isorhizas (a atrichs) and microbasic heterotrichous euryteles, while that of the strobila and ephyra consisted of a atrichs, euryteles, and holotrichous haplonemesContribution No. 546 from the Virginia Institute of Marine Science, Gloucester Point, Virginia 23062, USA.     

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     

Strobilation,budding and initiation of scyphistoma morphogenesis in the rhizostome Cassiopea andromeda (Cnidaria: Scyphozoa)

Scyphistomae of Cassiopea andromeda Forskål, 1775 containing symbiontic zooxanthellae did not develop medusae at a constant temperature of 20°C, but monodisc strobilation was initiated after transfer of the polyps to 24°C. After release of the ephyrae and regeneration of the hypostome and tentacular region, the recovered polyps either produced vegetative buds or entered a new strobilation phase. Formation of motile, planula-like buds was not found to be indicative of unfavourable environmental conditions. Intensity of budding was positively correlated with available food and with increase of temperature. Budding was negatively correlated with the number of polyps maintained per dish and with the conditioning of the sea water. Under optimal feeding and temperature conditions, polyps could simultaneously produce chains of buds at 2 to 4 budding regions. Settlement and development of buds into scyphistomae was suppressed in pasteurized sea water and in pasteurized sea water containing antibiotics, but polyps developed from buds in the presence of algal material taken from the aquarium, debris or egg shells of Artemia salina, or on glass slides which had been incubated in used A. salina culture medium. Several species of marine bacteria were detected after staining these slides. One, a Gramnegative coccoid rod, which was identified as a nonpathogenic Vibrio species, was isolated, cultivated as a pure strain, and was proved to induce the development of C. andromeda buds into polyps. Millipore filter-plates coated with Vibrio sp. cells grown in suspension culture were ineffectual, but diluted filtrate initiated polyp morphogenesis. The inducing factor is obviously not a constituent of the bacterial cell surface, but is a product of growing Vibrio sp. cells released into the medium. This product was found to be relatively heat-stable and dialyzable. As to the basic mechanism involved in the induction of polyp formation, it is suggested that the inducing factor (s) acts bimodally by inducing pedal disc development and by eliminating a head inhibitor originating from the basal end of the bud. The life history, and various aspects of medusa-formation and of vegetative reproduction in scyphozoans are reviewed and discussed with particular reference to rhizostome species. Special attention has been paid to some reports of larval metamorphosis controlled by marine bacteria.     

A commensal relationship between the scyphozoan medusae<Emphasis Type="Italic"> Catostylus mosaicus</Emphasis> and the copepod<Emphasis Type="Italic"> Paramacrochiron maximum</Emphasis>

The copepod Paramacrochiron maximum was found in high numbers (up to 5,675 copepods/medusa) on the oral arms of the scyphozoan Catostylus mosaicus. This association was considered to be commensalism for the following reasons: P. maximum (Lichomolgidae) was abundant on the medusae (approximately 805 copepods/kg of medusae) and very rare in the water column (approximately 5.99×10^-4 copepods/kg of water); copepodites and adults of the symbiont were present on the host; the copepods were on the medusae both day and night, at different times (nine occasions between March 1999 and May 2000) and different locations (Botany Bay and Lake Illawarra, NSW, Australia). Over 40 taxa of plankton were found on the oral arms of C. mosaicus (including protists, cnidarians, polychaetes, molluscs, a wide range of holoplanktonic and meroplanktonic crustaceans, chaetognaths and fish eggs). These taxa were abundant in the water column and we concluded that they were prey. Symbiotic amphipods and carangid fishes were found with medusae. We conclude that there is a symbiotic association between P. maximum and C. mosaicus and care should be taken not to confound these copepods with the prey of C. mosaicus. Poecilostomid copepods are well known for consuming mucus and feeding is likely to be a major reason for the association.Communicated by G.F. Humphrey, Sydney     

Spermatozeugmen und Paarungsverhalten bei Tripedalia cystophora (Cubomedusae)

New observations on the complete life cycle of Tripedalia cystophora Conant have revealed that—because of the radialsymmetrically constructed body of the polyp and its complete metamorphosis into one medusa—the “Cubozoa” are unique in their development, systematics and evolution. the same is true for the sexual biology of their medusae. It has long been known that the female T. cystophora are larviparous; fertilized eggs develop within the gastral pockets into free-swimming planulae. How do spermatozoa approach the eggs inside the female? In laboratory cultures, young medusae could be raised to maturity. Rearing experiments and observations yielded surprising results, revealing a situation which is new for Cnidaria. In T. cystophora, ripe medusae of both sexes can be identified easily by the different colour and structure of the gonads. In the gonads of the ripe male, numerous spermatozoa are joined to form spermatozeugmata of a simple type. Numerous spermatozeugmata form big globular bodies (spermatophores), which develop in small grooves on the inside surface of the stomach. The spermatophores are transferred directly from the male to the female during successive steps of special mating behaviour. The observations confirm that Cubomedusae have reached the peak of medusan specialization.     

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.     

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.