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.     

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.     

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.     

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     

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.     

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.     

Reproductive periodicity,spawning and development of the deep-sea scleractinian coral <Emphasis Type="Italic">Flabellum angulare</Emphasis>

Increasing threats to deep-sea corals highlight the need to expand knowledge of these taxa so that conservation measures can be developed. The present study focused on the reproductive patterns of the deep-sea solitary coral Flabellum angulare. A series of samples (n = 398) collected in 2006–2008 in the northwest Atlantic at depths of 925–1,430 m revealed that gametogenesis was synchronous among males and females and fluctuated seasonally. Initiation of gamete synthesis was estimated to be in August–September and spawning in June. Further analysis and daily monitoring of 30–60 individuals maintained in a flow-through mesocosm showed that gamete release occurred in March–June with a peak in May. Release of oocytes coincided with rising seawater temperatures and high deposition rates indicative of elevated water column productivity. Oocytes (900–1,200 μm diameter) were released through the oral cavity, generally in bundles of 3–5 surrounded by mesenterial filaments and attached to a thread (30–50 mm long). As oocytes became free in the water column, ovulation occurred followed by fertilization. Eggs/embryos initially remained on the tentacles of the spawner before either falling onto the substratum or floating to the surface. The embryos developed into planula larvae measuring 2–3 mm in length within about 24 h. Together, these findings shed new light on the strategies used by deep-sea corals to maximize their reproductive success.     

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.     

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.     

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.     

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.     

The utility of statoliths and bell size to elucidate age and condition of a scyphomedusa (Cassiopea sp.)

Scyphomedusae play important roles in marine ecosystems and are of economic significance. However, no reliable techniques for estimating scyphomedusa age have been documented. This study focused on the utility of Cassiopea sp. (Cnidaria: Scyphozoa) statoliths, statocysts, and body size as proxies for age of medusae. Reared medusae of known age and a manipulative experiment were used to assess the accuracy and reliability of four measures of age: number of statoliths, size (diameter) of statoliths, area of statocyst (housing statoliths), and bell diameter. Bell diameter provided the most accurate measure of age under constant conditions, but was increasingly inaccurate under varying environmental conditions. In contrast, the average number of statoliths per medusa reflected age with relatively low accuracy, but did not vary with changes in food availability and salinity. Only temperature influenced the average number of statoliths. Comparisons of bell diameter to the number of statoliths in medusae under low food availability to those fed well showed that the ratio of medusa size to the number of statoliths can be used to recognise medusae that are relatively poorly conditioned. Statoliths, therefore, provide a tool for studying both population ecology and the influence of environmental variation on medusa growth.     

Mechanics of prey selection by ephyrae of the scyphomedusa Aurelia aurita

In situ feeding patterns of ephyrae of the jellyfish Aurelia aurita (Linnaeus) revealed the importance of relatively large (>1 mm) prey in the diet of these scyphozoan predators. These studies were carried out in Narragansett Bay, Rhode Island, USA in March and April, from 1993 through 1996. Rotifers were the only small prey ingested in quantity, and then only when they were unusually abundant in the plankton. Copepod nauplii, similar in size to rotifers and equally abundant, were rarely consumed. Since copepods evince rapid escape responses, this observation suggested a role for prey escape in determining prey vulnerability, while the predominance of large prey in the diet suggested a role for prey size. Using two dimensional video observations of free-swimming ephyrae and their prey in the laboratory we tested hypotheses about the mechanisms underlying these dietary patterns, comparing mechanisms for capture of large versus small prey and for prey of equal size but differing escape behaviors. Capture efficiencies of ephyrae feeding on large prey were 4 to 12 times greater than for small prey taxa. Capture efficiencies for prey of equal size also differed significantly, indicating that other factors influence the outcome of predator–prey interactions. Most prey captures occurred while the ephyrae were swimming and creating fluid flows that entrained prey into the subumbrellar region. Even copepod nauplii were frequently drawn into the subumbrella of swimming ephyrae despite average potential escape velocities (25.7 mm s⁻¹) that exceeded mean maximum velocity of fluid flows around the ephyrae (13.1 mm s⁻¹). Large prey were more likely than small prey to contact nematocyst-bearing surfaces both before and after entrainment in flow fields. With regard to behavior, prey escape speeds were not the only predictor of prey vulnerability. Prey that continued swimming after entrainment (rotifers and brine shrimp) were captured more often than prey of equal size that ceased normal swimming (copepod nauplii and barnacle nauplii). Copepod nauplii were the prey least likely to be captured because they either “played dead” and were expelled from the subumbrella of the ephyrae before contacting a surface, or they eventually escaped at high velocity. These observations indicate that size-selective predation by ephyrae of A. aurita can be influenced by a variety of behavioral responses of the prey. Received: 9 April 1997 / Accepted: 5 September 1997     

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.     

A floating mid-water coral nursery as larval dispersion hub: testing an idea

The global decline in reef health has prompted the need for effective management methodologies, including the development of active restoration measures. One such approach is the ‘gardening concept’ that involves use of underwater nurseries where coral fragments are farmed before their transplantation into denuded reefs. Here we document enhanced sexual reproduction in colonies of the coral Stylophora pistillata cultured in mid-water floating nursery situated in nutrient enriched water, near the fish farms in Eilat, Red Sea. We found that after 2 years of nursery, the average number of oocytes per polyp in farmed colonies was ca. 35% higher than in corresponding naturally growing colonies. Small branches in the nursery developed gravid colonies that released equal (or more) numbers of planula larvae as compared to similar size, 5-year old naturally growing colonies. These nursery-borne planulae possessed more zooxanthellae and contained more chlorophyll per larva. While settled and metamorphosed in equal rates compared to planulae originated from reef-grown colonies, the nursery borne planulae developed faster growing young colonies. We estimate that a coral nursery could generate, during the reproductive season, tens of millions of planula larvae and therefore should be regarded as a ‘larval dispersion hub’ that can be used as a management tool for natural recruitment enhancement.     

Bioenergetics of ephyra larvae of the scyphozoan jellyfish Aurelia aurita in relation to temperature and salinity

The body mass of Aurelia aurita ephyrae was better correlated with the diameter of the central disc than with the distance between opposite rhopaliae or distance between opposite lappet tips. Body dry weight (y, in μg) related to the disc diameter (x, in mm) through the equation y = 22.33 x ^1.99. The exponent 1.99 was significantly lower than that for the medusa stage, indicating a tendency to grow in diameter rather than in weight through the ephyra stage. The average ash-free dry weight (AFDW) of ephyrae was 38.0% of the dry weight. The AFDW/diameter relationship was used to convert measured diameters to body AFDW and calculate succession in body mass, daily ration, daily growth rate and gross growth efficiency. Effects of temperature (6, 9.5, 12, 15 and 18 °C) and salinity (17.5, 22, 26, 30.5 and 35 PSU) on these parameters and feeding were studied at saturated prey concentration (222 Artemia nauplii l⁻¹ initial concentration) by daily measurements over 10 d. There was a strong effect of temperature for total ingestion, growth rate, growth efficiency and final body mass of individual ephyrae, whereas the daily ration was not significantly different between the different temperatures. The experimental group kept at the highest temperature (18 °C) diverged the most, and ephyrae at this temperature ingested 2.7 times more and increased in weight 5.4 times more than at 6 °C. The average daily growth rate and gross growth efficiency of these ephyrae were 34.5% and 25.1%, respectively, significantly higher than at 6, 9.5 and 15 °C. Significant effects of salinity were shown for total ingestion, daily ration, daily growth rate and final weight, although only total ingestion and daily ration diverged sufficiently to show effects in a post-hoc test. This test showed that total ingestion was significantly different for all salinities except between 22 and 35 PSU and between 17.5 and 26 PSU. The daily ration for 35 PSU diverged from all other salinities, whereas none of the other salinities showed any significant differences. Thus, provided food in excess A. aurita can double its weight every 2 to 4 d, dependent on temperature and can therefore develop to the medusa stage in short time. Differences in environmental salinity in the range 17.5 to 35 PSU have little or no effect on growth rate and growth efficiency, whereas our results indicate that the full seawater salinity (35 PSU) causes significantly higher ingestion rate compared to lower salinities. Received: 11 January 1999 / Accepted: 11 May 1999     

Seasonal development of scyphozoan medusae and the predatory impact of Aurelia aurita on the zooplankton community in the Bornholm Basin (central Baltic Sea)

The annual cycle of abundance and distribution of the scyphozoan medusae Aurelia aurita and Cyanea capillata was studied in the Bornholm Basin (central Baltic Sea) in 2002. Seasonal changes in prey composition and predatory impact were investigated by analyzing stomach contents. A. aurita occurred from July to November, with a maximum mean abundance of 2.3 ind. per 100 m³ in August, whereas C. capillata was caught in much smaller numbers from July to September. No ephyrae of either species were found; therefore, advection of medusae from the western Baltic Sea is assumed. From July to October, ~80% of A. aurita medusae was distributed in the upper 20 m above the thermocline, whereas C. capillata occurred only in the halocline below 45 m. A. aurita did not migrate vertically and fed mainly on the most abundant cladoceran species Bosmina coregoni maritima. Further prey organisms were the cladocerans Evadne nordmanni and Podon spp., mollusk larvae and copepods. Copepod nauplii and copepodite stages I–III were not eaten by the medusae, neither were fish eggs and larvae used as prey. Based on mean medusa and zooplankton abundance from the upper 20 m, the predatory impact was very low. In August, when mean abundance of A. aurita was highest, only 0.1% of the copepod and 0.5% of the cladoceran standing stock were eaten per day. However, in regions with higher medusa or lower zooplankton abundance, up to 7.9% of the cladoceran standing stock was consumed per day. Hence, A. aurita did not regulate the zooplankton community in the Bornholm Basin, and fish larvae did not suffer from competition with and predation by the medusae.Communicated by O. Kinne, Oldendorf/Luhe     

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

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

Reproductive biology of the deep-sea octocoral Drifa glomerata in the Northwest Atlantic

The present study examined the mode and timing of reproduction of poorly understood deep-water octocorals and the environmental factors that may influence their reproductive patterns. Data on reproductive characteristics of the octocoral Drifa glomerata (Alcyonacea: Nephtheidae) collected between 2004 and 2007 at ca. 100–330 m depth off Newfoundland and Labrador (eastern Canada) were compared among years, months and depth ranges. No male gonad was ever observed during the study. The ratio of fertile colonies possessing large pinkish polyps with oocytes/planulae was >50% throughout the year. The number of brooded planula larvae within a single fertile polyp varied between 1 and 10 for a total of approximately 40–3,000 in a whole colony. The size of oocytes and/or planulae was consistently greater in the polyps than in the branchlets, indicating that the development pathway of oocytes to planulae is from the branchlets to the polyps. Although larval production seemed to persist year round, the onset of major planulation events occurred in December–January of two consecutive years, when large mature planulae were released in correlation with the first increase of photoperiod and maximum temperature at 150 m. A second peak in planulation between April and early June followed the phytoplankton bloom. Seasonal trends were more apparent in colonies from <200 m, and the planula index varied among sampling depths and years. Larval release in a live colony under laboratory conditions occurred between January and June 2008, closely following predictions based on field samples.