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.     

Hebella parasitica (Cnidaria,Hydroida): A thecate polyp producing an anthomedusa

The thecate hydroid Hebella parasitica (Ciamician), reared under controlled conditions, produced an ephemerous medusa with gonads on the manubrium. This feature does not agree with the current classification of hydroids, according to which a thecate polyp is expected to produce a medusa with gonads on its radial canals. As no data on the reproduction of other species of Hebella are available, the systematic position of H. parasitica should be considered uncertain.Work done within the Finalized Project Oceanografia e i fondi marini, sub-project Risorse biologiche of the C.N.R.     

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.     

Complex reproductive cycle inEucheilota paradoxica (Hydrozoa: Leptomedusae): Medusae,polyps and frustules produced from medusa stage

Eucheilota paradoxica Mayer, 1900 is a small leptomedusa unknown in the Mediterranean Sea until 1977. Since then, it has become endemic and occurs in great abundance in autumn in the bay of Villefranche-sur-mer. Since no sexual stages have ever been collected, either from the Atlantic Ocean or from the Mediterranean Sea, we decided to investigate the still unknown life cycle of this species. Specimens were obtained in the bay of Villefranche-sur-mer from plankton hauls between 0 and 50 m depth during October and November 1986. At temperatures >18°C, the medusa reproduces intensively by asexual medusal budding (one young medusa liberated per two days). This asexual reproduction explains the rapid proliferation of the species and the origin of the large population every autumn. In cultures maintained at temperatures <18°C, two other reproduction modes, not yet reported in a leptomedusa, were observed: (1) At 16° to 17°C, frustules bud along the canals; these frustules are dormant, giving rise to a small polyp when the temperature increases once more above 18°C. (2) At temperatures close to 15°C each medusa, in place of the gonads, differentiate into one or two polyps; these polyps develop progressively, concomitant with gradual regression of the mother medusa.     

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 reconstitutional performances of the Leptomedusa Campanularia jonstoni

The reconstitutional abilities of the Leptomedusa Campanularia jonstoni (Alder) were studied by means of isolating various regions and sectors of the animals. All such fragments first restore the typical bell-shape by a rapid morphodynamic process, the mechanisms of which are not yet understood. In a second, much slower step, the missing organs such as manubrium, radial and circular canals, tentaces, statocysts and gonads are regenerated to a varying extent. The original radial symmetry is practically never restored. The manubrium plays a key role in these processes. When it is present in the fragment isolated, the regeneration of the peripheral components of the vascular system (radial canals) originates from the manubrium. When the fragment is devoid of a manubrium, the latter is first reconstituted by the subumbrellar tissues including ectoderm and entoderm. Interradial sectors lacking radial and circular canals, as well as a manubrium, are capable of reconstituting a small medusa using exumbrellar and subumbrellar tissues alone. Gonads are regenerated along with the reconstitution of radial canals. The histodynamics of these processes have, so far, not been investigated.     

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.     

Temperature,salinity, and prey effects on polyp versus medusa bud production by the invasive hydrozoan<Emphasis Type="Italic"> Moerisia lyonsi</Emphasis>

Hydrozoan species are renowned for flexible asexual reproduction, which may predispose them to be successful invaders. Polyps of the invasive hydrozoan Moerisia lyonsi (Boulenger, 1908) have very high rates of asexual production of both polyp and medusa buds. In order to determine how environmental factors affect asexual reproduction in M. lyonsi, the quantitative relationships between polyp bud and medusa bud production were studied in a 31-day laboratory experiment during August 2001. The combined effects of prey (4, 8, 12, 16 Acartia tonsa copepods polyp^–1 day^–1), temperature (20°C, 29°C), and salinity (5, 15, 25) were tested on the development times for polyp buds (DT_p) and medusa buds (DT_m), the total asexual reproduction rate (ARR, no. buds polyp^–1 day^–1), and the ratio of medusa bud to total bud production (R_m). Greater food consumption significantly and directly enhanced ARR and R_m and shortened DT_p and DT_m. A lower temperature (20°C) and higher salinity (25) reduced food consumption, lengthened development times, and decreased ARR and R_m, with opposite effects for the higher temperature (29°C) and lowest salinity (5). The patterns of variation of these reproductive parameters are more complex. DT_m was most sensitive and was significantly and directly affected by all three measured factors. In addition to food consumption, direct effects were seen by temperature on DT_p and by salinity on R_m. ARR was directly affected only by food consumption. Overall, DT_p, DT_m, and R_m were more sensitive to environmental differences than was ARR. More favorable conditions enhanced medusa bud production. The adaptive reproductive processes and their significance for the maintenance and dispersal of M. lyonsi are discussed.Communicated by J.P. Grassle, New Brunswick     

The life cycle of Eutima commensalis sp. nov. (Eutimidae,Hydromedusae)

Developmental stages of the medusa Eutima commensalis sp. nov. were obtained from plankton collections made at 2 fixed stations, one at Fareway Buoy and the other at Aroor in the Cochin backwater system. Six stages were arbitrarily assigned and described. The hydroid and the medusa of this form were found to occur in these waters throughout the year, except during the monsoon months from June to September. The hydroid is endocommensalic on wood-boring molluscs such as Nausitora hedleyi, Teredo furcifera and Martesia striata.     

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.     

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.     

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.     

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     

<Emphasis Type="Italic">Tiburonia granrojo</Emphasis> n. sp., a mesopelagic scyphomedusa from the Pacific Ocean representing the type of a new subfamily (class Scyphozoa: order Semaeostomeae: family Ulmaridae: subfamily Tiburoniinae subfam. nov.)

Submersible observations off Japan, Hawaii, and California, USA, at depths of 645–1497 m, have revealed a previously undescribed species of large semaeostome scyphomedusa. These observations were made from 1993 to 2002. The medusa, Tiburonia granrojo n. sp. is sufficiently different from other species in the family Ulmariidae to justify the creation of a new subfamily (Tiburoniinae). This subfamily and species are distinct in overall bell morphology and color, lacking any marginal tentacles, and having four to seven short, thick oral arms that extend beyond the bell margin. The entire medusa, including the mesoglea, is a deep red. A new key to the subfamilies of the Ulmaridae and large subunit rRNA sequence information for T. granrojo are provided. That new species of this size and mass are still being discovered in the deep waters of the world suggests that deep-water species remain undescribed. Additional information is available at http://www.mbari.org/midwater     

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.     

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.