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.     

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.     

Comparison of the statolith structures of <Emphasis Type="Italic">Chironex fleckeri</Emphasis> (Cnidaria,Cubozoa) and <Emphasis Type="Italic">Periphylla periphylla</Emphasis> (Cnidaria,Scyphozoa): a phylogenetic approach

The rhopalia and statocysts of Periphylla periphylla (Péron and Lesueur in Ann Mus Hist Nat Marseille 14:316–366,1809) and Chironex fleckeri Southcott (Aust J Mar Freshw Res 7(2):254–280 1956) were examined histologically and showed several homologous characteristics. Differences in sensory area distribution could be connected to a slightly different functionality of equilibrium sensing. In P. periphylla, the statoliths (crystals) grow independently of each other; whereas in C. fleckeri, one large crystal covers the smaller ones. The structures of both statoliths were examined in detail with single-crystal diffraction, microtomography and diffraction contrast tomography. The single compact statolith of C. fleckeri consisted of bassanite as was previously known only for other rhopaliophoran medusae. An origin area with several small oligocrystals was located in the centre of the cubozoan statolith. The origin areas and the accretion of statoliths are similar in both species. Our results lead to the assumption that the single bassanite statolith of C. fleckeri (Cnidaria, Cubozoa) is a progression of the scyphozoan multiplex statolith. It is therefore suggested that the Cubozoa are derived from a scyphozoan ancestor and are a highly developed taxa within the Rhopaliophora.     

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 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.     

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     

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.     

Larval metamorphosis of the sea urchins,<Emphasis Type="Italic"> Pseudocentrotus depressus</Emphasis> and<Emphasis Type="Italic"> Anthocidaris crassispina</Emphasis> in response to microbial films

In Japan, mass-production of sea urchin juveniles involves the culture of periphytic diatom films on plastic plates in 5- to 15-tonne tanks for the induction of larval metamorphosis. This study focused on the larval response of sea urchins, Pseudocentrotus depressus and Anthocidaris crassispina, to natural microbial films in the sea and diatom-based films formed in the tanks. The effect of diatoms and bacteria on larval metamorphosis was also examined using laboratory-cultured diatom-based films in the presence of germanium dioxide and antibiotics during film culture. Furthermore, the nature of the cue of the cultured diatom-based film was also investigated. Results showed that P. depressus and A. crassispina metamorphosed both on natural microbial films and diatom-based films in a tank. In the sea, the metamorphosis (%) of P. depressus increased gradually in accordance with the immersion period of film formed on glass slides, whereas the larval metamorphosis of A. crassispina had a bell-shaped response curve. In the tank, although the diatom-based films showed a low inducing activity for larval metamorphosis of A. crassispina, the metamorphosis of P. depressus larvae increased linearly in accordance with the diatom density. These results suggest that diatom-based films could promote the larval metamorphosis of P. depressus, but are less important in A. crassispina. In a simultaneous larval assay (May), P. depressus showed a higher percentage of metamorphosis than A. crassispina. We concluded that the former is more sensitive to diatom-based film than the latter and that this is due to differences in their natural habitats. For laboratory-cultured diatom-based film, both species of sea urchins showed a similar response, in which reduction in diatom and bacteria density resulted in a decrease in the original inducing activity. There seems to be a synergistic effect between diatom and bacteria in inducing larval metamorphosis. Films subjected to treatment with 0.1 N HCl were no longer inductive for either sea urchin, while those films treated with 40^°C heat or EtOH (5% and 10% EtOH) showed a significant reduction in the inducing activity. Thus the surface-associated cues may be highly susceptible to the above treatments.Communicated by T. Ikeda, Hakodate     

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.     

Medusa buds on Eirene elliceana gonads (Leptomedusae,Eirenidae)

The Indo-Pacific leptomedusa Eirene elliceana (Agassiz and Mayer, 1902) is known to produce blastostyles budded from the gonads of the adult medusa. Three specimens out of 5, taken during the International Indian Ocean Expedition (IIOE) in the Bay of Bengal, showed medusa buds growing on young gonads whose sex could not be determined. The new findings are illustrated.Contribution No. 320 of the Instituto Oceanográfico da Universidade de São Paulo, S. Paulo, Brazil.     

<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     

Larval metamorphosis of the mussel <Emphasis Type="Italic">Mytilus galloprovincialis</Emphasis> in response to <Emphasis Type="Italic">Alteromonas</Emphasis> sp. 1: evidence for two chemical cues?

Bacterial isolates from multi-species biofilms were identified by 16S rDNA gene sequences and investigated for their inductive effects as monospecific biofilms on larval metamorphosis of Mytilus galloprovincialis. Alteromonas sp. 1 biofilm was found to have inductive activity, which increased with increasing cell density. The cue(s) of Alteromonas sp. 1 biofilm responsible for inducing larval metamorphosis was further investigated. Treatment of the biofilm with formalin, ethanol, heat or ultraviolet irradiation resulted in a significant reduction in the inductive activity of Alteromonas sp. 1, and the crude extract of surface-bound products of the biofilm showed no activity. These results indicated that if the cue was a surface-bound chemical cue, it was unstable, or susceptible to the treatments or the extraction process. On the other hand, the inductive activity of treated biofilms had a linear regression to the cell survival of bacteria, indicating a metabolically active biofilm was a requirement for larval metamorphosis. Conditioned water of the biofilm did not induce larvae to metamorphose. However, larval crawling behavior in the conditioned water was the same as that in the biofilm prior to larval metamorphosis, and significantly different to larval behavior in seawater. This indicated that a potential or partial waterborne cue existed, but remained inactive when alone. A synergistic effect of the conditioned water with formalin-fixed Alteromonas sp. 1 biofilm resulted in a significant increase in larval metamorphosis. Heat treatment and fractionation of the conditioned water demonstrated that the waterborne cue was heat-stable and <3,000 Da in molecular weight. Platinum-coating, Lentil Lectin and Wheat Germ Agglutinin treatments of the formalin-fixed biofilm significantly reduced its synergistic effect with the conditioned water, suggesting that a surface-bound cue was present on the biofilm and that the cue might be associated with the bacterial exopolysaccharide or glycoprotein. Evidence presented here suggests that two chemical cues derived from bacteria act synergistically on larval metamorphosis of Mytilus galloprovincialis.     

Correlating gene expression with larval competence,and the effect of age and parentage on metamorphosis in the tropical abalone<Emphasis Type="Italic"> Haliotis asinina</Emphasis>

The non-geniculate crustose coralline alga (CCA) Mastophora pacifica can induce the metamorphosis of competent Haliotis asinina (Vetigastropoda) larvae. The ability to respond to this natural cue varies considerably with larval age, with a higher proportion of older larvae (e.g. 90 h) able to metamorphose in response to M. pacifica than younger larvae (e.g. 66 h). Here we document the variation in time to acquisition of competence within a larval age class. For example, after 18 h of exposure to M. pacifica, approximately 15 and 36% of 84 and 90-h-old H. asinina larvae had initiated metamorphosis, respectively. This age-dependent response to M. pacifica is also observed when different aged larvae are exposed to CCA for varying periods. A higher proportion of older larvae require shorter periods of exposure to CCA than younger larvae in order to initiate metamorphosis. In this experiment, as in the previous, a small proportion of young larvae were able to respond to brief periods of CCA exposure, suggesting that they had developed the same state of competency as the majority of their older counterparts. Comparisons of the proportions of larvae undergoing metamorphosis between families reveals that parentage also has a significant (P<0.05) affect on whether an individual will initiate metamorphosis at a given age. These familial differences are more pronounced when younger, largely pre-competent larvae (i.e. 66 h old) are exposed to M. pacifica, with proportions of larvae undergoing metamorphosis differing by as much as 10 fold between families. As these data suggest that variation in the rate of development of the competent state has a genetic basis, and as a first step towards identifying the molecular basis to this variation, we have identified numerous genes that are differentially expressed later in larval development using a differential display approach. Spatial expression analysis of these genes suggests that they may be directly involved in the acquisition of competence, or may play a functional role in the postlarva following metamorphosis.Communicated by M.S. Johnson, Crawley     

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.     

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.     

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.     

Growth and differentiation during delayed metamorphosis of feeding gastropod larvae: signatures of ancestry and innovation

Extent of larval growth among marine invertebrates has potentially profound implications for performance by benthic recruits because body size influences many biological processes. Among gastropods, feeding larvae often attain larger size at metamorphic competence than non-feeding larvae of basal gastropod clades. Delay of metamorphosis can further influence size at recruitment if larvae continue to grow during the delay. Some caenogastopod larvae grow during delayed metamorphosis, but opisthobranch larvae do not. Data on larval growth of neritimorph gastropods are needed to help determine which of these growth patterns for planktotrophic gastropod larvae is more derived. We cultured planktotrophic larvae from all three major gastropod clades with feeding larvae through delays of metamorphosis of 3–10 weeks. Larvae of the caenogastropod Euspira lewisii and the euthyneurans Haminoea vesicula (Opisthobranchia) and Siphonaria denticulata (Pulmonata) conformed to previously described growth patterns for their respective major clades. Furthermore, the caenogastropod continued to lengthen the prototroch (ciliary band for swimming and feeding) and to differentiate prospective post-metamorphic structures (gill filaments and radular teeth) during delayed metamorphosis. Larvae of the neritimorph Nerita atramentosa arrested shell growth during delayed metamorphosis but the radula continued to elongate, a pattern most similar to that of non-feeding larvae of Haliotis, a vetigastropod genus. Character mapping on a phylogenetic hypothesis suggests that large larval size and capacity for continued growth during delayed metamorphosis, as exhibited by some caenogastropods, is a derived innovation among feeding gastropod larvae. This novelty may have facilitated post-metamorphic evolution of predatory feeding using a long proboscis.