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.     

Dissolved organic substances during a Phaeocystis pouchetii bloom in the German Bight (North Sea)

Dissolved organic substances (carbohydrates, organic nitrogen, free amino acids) were measured in the German Bight (North Sea) in June, 1981. During and before this survey, sea foam was observed in the east Frisian coastal water and it accumulated on the nearby beaches to an unusually high extent. In this coastal water area, a large Phaeocystis pouchetii Lagerheim bloom and very high concentrations of dissolved organic matter were found. The above dissolved organic substances were all positively correlated to a highly significant degree with P. pouchetii cell numbers in the bloom area. An influence of salinity (or river water) on this correlation could be excluded. Thus, exudation or decomposition products of P. pouchetii were most likely the cause of the unusually high concentrations of dissolved organic matter in the bloom area off the east Frisian coast, where P. pouchetii blooms have been reported for many years. Ammonia concentrations were very low in the P. pouchetii bloom area; this and the accumulation of dissolved organic substances might lead to speculation that decomposition of dissolved organic matter by bacteria could have been reduced due to antibiotic activity of P. pouchetii.     

Population dynamics of Eudendrium racemosum (Cnidaria, Hydrozoa) from the North Adriatic Sea

Benthic suspension feeders in shallow waters develop in relation to the food availability and the variation of physical parameters giving rise to complex communities that act as a control factor on the plankton biomass. The aim of the work is to establish the role of the hydrozoan Eudendrium racemosum in the energy transfer from the plankton to the benthos in marine food chains of the North Adriatic Sea. This study highlighted that the hydroid biomass changed over time in relation to temperature and irradiance, and the highest abundance was observed during summer with about 400,000 polyps m⁻² (about 19 g C m⁻²). The population suffered an evident summer decrease in relation to a peak of abundance of its predator, the nudibranch Cratena peregrina, whose adult specimens were able to eat up to 500 polyps day⁻¹ and reached an abundance of 10 individuals m⁻². The gut content analysis revealed that the hydroid diet was based on larvae of other benthic animals, especially bivalves and that the amount of ingested preys changed during the year with a peak in summer when it was estimated an average predation rate of 13.7 mg C m⁻² day⁻¹. In July, bivalves represented over 60 % of the captured items and about 18 mg C m⁻² day⁻¹. Values of biomass of E. racemosum are the highest ever recorded in the Mediterranean Sea, probably supported by the eutrophic conditions of the North Adriatic Sea. Moreover, our data suggest that settling bivalves provide the greater part of the energetic demand of E. racemosum.     

Overwintering strategies of dominant calanoid copepods in the German Bight, southern North Sea

Abundance, stage composition and reproductive parameters (egg production, egg viability, proportion of spawning females) of the four copepod species Acartia clausi, Centropages hamatus, C. typicus and Temora longicornis were measured at the long term sampling station Helgoland Roads (German Bight, southern North Sea) from September 2003 to May 2004 to study their overwintering strategies. A. clausi was overwintering as females with arrested reproduction from November to January. T. longicornis, which is known to produce resting eggs in the North Sea, had a pelagic population with all developmental stages present during winter and reproductive rates closely related to food concentrations. Although their females produced eggs in response to ambient food conditions, both C. hamatus and C. typicus were rare in the pelagic. The C. hamatus population returned in May, probably from resting eggs, whereas C. typicus depended on advection. The Centropages species seemed to be less adapted to pelagic life in winter than A. clausi and T. longicornis. Sporadic occurrence of large numbers of nauplii and young copepodids of A. clausi and Centropages spp. pointed to different overwintering strategies or more successful survival in adjacent regions and advection of them into the waters around Helgoland island. While A. clausi was decoupled from environmental conditions in late autumn and winter, the other species were able to respond to variations in the food environment. Thus, egg production of T. longicornis increased during an unusual autumn diatom bloom.     

Life history and settlement preferences of the edible jellyfish Catostylus mosaicus (Scyphozoa: Rhizostomeae)

 The life history and settlement preferences of larvae of Catostylus mosaicus (Scyphozoa: Rhizostomeae) (Quoy and Gaimard, 1824) were investigated in New South Wales, Australia, over a 2 mo period beginning in November 1998. The life history consisted of an alternation between a sexual, medusoid stage and an asexual, polypoid stage, and was similar to that described for other rhizostomes. Planula larvae were brooded by the adults. Approximately 4 d after collection, larvae settled on a variety of substrata including wood, sandstone, shell, seagrass and glass, and metamorphosed into four-tentacled polyps. The number of tentacles increased and polyps strobilated when they had between 12 and 20 tentacles. Strobilation occurred within 15 d of settlement, but only polyps that settled on the concave surfaces of the shells strobilated. Both monodisk and polydisk strobilation was observed. Ephyrae were raised for one month and were observed developing oral arms. Polyps reproduced asexually via the formation of podocysts, by production of buds, and by partial fission. Received: 30 April 1999 / Accepted: 27 August 1999     

Morphology and development of benthic and pelagic life stages of North Sea jellyfish (Scyphozoa, Cnidaria) with special emphasis on the identification of ephyra stages

Jellyfish blooms or invasions could be detected in an early phase of development if the youngest medusa stages (ephyrae) and their early growth stages (post-ephyrae) were identifiable in plankton samples but a useful identification key for ephyrae in early growth stages is lacking for most species. In the present study, the identification characteristics of adult North Sea scyphomedusae (Aurelia aurita, Cyanea capillata, Cyanea lamarckii, Chrysaora hysoscella, Rhizostoma octopus) collected around the island of Helgoland (German Bight) in July?CAugust 2003 and 2004 are described. Planula larvae were measured and reared to polyps in the laboratory. The process of ephyrae development asexually produced by the polyps (strobilation) was photo-documented. Photographs of the ephyrae growth stages were combined with drawings of features useful for the species identification. The provided identification key allows discrimination among post-ephyrae from plankton samples, probably leading to conclusions on the development of jellyfish blooms and their causes.     

Bioluminescence of deep-sea coronate medusae (Cnidaria: Scyphozoa)

Bioluminescence is the production of visible light by a living organism. The light commonly appears as flashes from point sources (involving one or more cells, usually described as photocytes) or as a glandular secretion. A visible flash usually involves synchronous light emission from a group of cells or, if from a single-celled organism such as a dinoflagellate, from a group of organelles. The number of cells (or organelles) responding synchronously is the main determinant of the flash intensity. Bioluminescence is a common phenomenon in many deep-sea animals and is widespread among the Cnidaria. In this paper, we compare and contrast in situ and laboratory recordings of the bioluminescent responses of specimens of the deep-sea scyphozoans Atolla wyvillei, Atolla vanhoffeni, Atolla parva, Nausithoe rubra, Paraphyllina intermedia, Periphyllopsis braueri and Periphylla periphylla. Displays in all seven species consist of localised flashes and propagated waves of light in the surface epithelium. The first few single waves propagate at rates of up to 60 cm s^-1 but subsequent ones in any sequence of stimuli gradually decrease in speed. After several single wave responses, a subsequent stimulus may elicit multiple waves that persist for several seconds. Following such a frenzy, the specimen becomes temporarily refractory to further stimuli, but if rested will recover its normal responses and may produce further frenzies. The dome area, situated above the coronal groove, of the genera Paraphyllina, Periphylla, and Nausithoe is covered with luminescent point sources. Such point sources are generally absent from the dome of species of Atolla. Captured specimens of A. parva also produce secretory bioluminescence, corroborating prior in situ observations of this ability. Secretory bioluminescence in P. periphylla takes the form of scintillating particles released from the lappet margins. We did not observe secretory displays in specimens of any other species in the laboratory, but one instance of apparent secretory luminescence was recorded in situ in a specimen of A. wyvillei.Communicated by J. P. Thorpe, Port Erin     

Requirement of exogenous inducers for metamorphosis of axenic larvae and buds of Cassiopea andromeda (Cnidaria: Scyphozoa)

Planula larvae and asexually-produced buds of the rhizostome scyphozoan Cassiopea andromeda (collected throughout the year in Eilat, Israel) have the ability, under axenic conditions, to attach to a substrate and undergo morphogenetic development to form a polyp (=scyphistoma) in: (1) the presence of unidentified inducers found in the adult habitat and (2) the presence of cefined organic compounds. Axenic planulae and buds were unable to settle and complete metamorphosis in autoclaved artificial or natural seawater from the North Sea when maintained without food, but continued swimming while decreasing in size and protein content, eventually dying within three months. When maintained in autoclaved seawater from the Red Sea, between 25 and 46% of the planulae and 4 and 11% of the buds metamorphosed within 30 d. Axenic solutions of cholera toxin, thyroid stimulating hormone, and pancreatic casein hydrolysate peptides in artificial seawater induced morphogenic development of 20 to 100% of planulae and buds within 2 to 18 d. The natural inducer(s) in Red Sea seawater, though unidentified, may have characteristics similar to the large proteins and small peptide inducers used in this study. Planulae and buds older than 20 d metamorphosed sooner and responded to lower concentrations of pancreatic casein hydrolysate peptides than younger individuals. This may be a physiological mechanism for enhancing metamorphosis and survival in nature. The data show that settlement and metamorphosis can be induced by solutions of cholera toxin and thyroid stimulating hormone, suggesting that, as in mammalian systems, the mechanism of action of these chemicals may involve cyclic adenosine monophosphate (cAMP) as an intermediate messenger. However, dibutyric cAMP, which is capable of passing through membranes and functioning normally inside the cell, did not induce metamorphosis of buds, and the levels of intracellular cAMP in buds and larvae typically increased slowly during induction of metamorphosis, unlike the high and rapid increases associated with cAMP-mediated biochemical events in mammalian cells. These results suggest that the observed cAMP changes seen were associated with metamorphic development, but not with the triggering mechanism.     

Substratum preferences in planula larvae of two species of scleractinian corals, Goniastrea retiformis and Stylaraea punctata

To test whether coral planulae recruit randomly to different coral reef habitats or have specific substratum preferences, the settling behavior of planulae from two shallow water coral species from Pago Bay, Guam (13°25.02N, 144°47.30E) were examined in the laboratory in June and July of 1995. Goniastrea retiformis is generally restricted to the shallow reef front (<10 m depth) in areas dominated by crustose coralline algae (CCA), while Stylaraea punctata is abundant on inner reef flats were CCA coverage is low and sand and carbonate rubble covered by biofilms is common. When presented with four substrata (1) carbonate rock scrubbed free of biofilm and dried as a control, (2) the CCA Hydrolithon reinboldii, (3) the CCA Peyssonelia sp., and (4) naturally conditioned carbonate rubble covered by a biofilm, G. retiformis larvae showed a significant preference for H. reinboldii, and S. punctata larvae for the carbonate biofilm treatment. The preference shown by S. punctata larvae for biofilmed surfaces did not diminish with increasing larval age up to 11 days. These results suggest that the larvae of both species are capable of habitat selection, and that the preferred substrata among those tested bears a relationship to the habitats in which adult colonies were found. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Organ-like gonads, complex oocyte formation, and long-term spawning in Periphylla periphylla (Cnidaria, Scyphozoa, Coronatae)

An account is given about the development of the gametes of the holopelagic coronate scyphomedusa Periphylla periphylla (Péron and Lesueur 1809). The gonads of the species are complex and differ from those thus far described in Scyphozoa in having this characteristic composition of trophocytes, follicle cells, gamete-releasing pores, mucus cells, and resorption cells. Our results differ from those of previous coronate studies with respect to the contact of the oocytes with the gonad tissue throughout the duration of development. Among the Medusozoa, follicle cells have thus far been considered as an apomorphy for the Staurozoa, but their presence in the Scyphozoa casts some doubts on this assumption. From morphological structure and examination of the gonads and gametes, it can be deduced that P. periphylla has true organs and that gamete release is continuous. These results supplement our knowledge of the reproductive biology of this mesopelagic species.     

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.     

Carbon metabolism and strobilation in Cassiopea andromedea (Cnidaria: Scyphozoa): Significance of endosymbiotic dinoflagellates

Scyphopolyps and scyphomedusae of Cassiopea andromeda Forskål (Cnidaria, Scyphozoa) containing dinoflagellate endosymbionts (zooxanthellae) were investigated for rates and pathways of carbon fixation. Photosynthesis by the algae, accounting for 80 and 15 mol C h^-1 on a dry weight basis in medusae and polyps, respectively, by far exceeds dark incorporation of inorganic carbon by the intact association. Photosynthetic carbon fixation is operated via C₃ pathway of carbon reduction. DCMU-treatment (1×10^-6 M and 1×10^-5 M) completely inhibits light-dependent carbon assimilation. Major photosynthates presumably involved in a metabolite flow from algal symbionts to animal tissue are glycerol and glucose. A total of 5–10% net algal photosynthate appears to be seleased in vivo to the host. This is probably less than the energy supply ultimately required for the nutrition of the polyps and medusae. The presence of zooxanthellae proved to be indispensable for strobilation in the scyphopolyps. However, photosynthesis by algal symbionts as well as photosynthate release is obviously not essential for the initiation of ephyrae as is shown by DCMU-treatment, culture in continous darkness, and aposymbiotic controls. It is therefore concluded that strobilation is supported, but not triggered by algal photosynthetic activity. The induction of strobilation thus seems to depend on a more complex system of regulation.     

Bud formation and metamorphosis inCassiopea andromeda (Cnidaria: Scyphozoa): A developmental and ultrastructural study

Asexual reproduction by formation of swimming buds which metamorphose directly into polyps plays a most important role in the propagation ofCassiopea andromeda (Cnidaria: Scyphozoa). (C. andromeda polyps, originally supplied by the Löbbecke Museum and Aquarium Düsseldorf, FRG, were cultured in our laboratories.) We have defined five budding stages and investigated epithelial recruitment and dynamics during bud formation using intracellular vital stains. The region of cell recruitment was found to encircle the budding site asymmetrically. The aboral side contributing considerably less to the developing bud than the oral and lateral sides. Furthermore, it was found that the epithelial flow involved in bud formation is part of a permanent apico-basal displacement of ectodermal cells. Light and electronmicroscopic investigations revealed that no drastic changes occur at the cellular level, neither in the ectoderm nor in the endoderm which both participate in bud formation. Scanning and transmissionelectron microscopic investigations of the swimming bud revealed that the ectoderm is composed of three, and the endoderm of two, cell types. Nerve elements have been detected near the mesoglea between both ecto- and endodermal cells. Amoebocytes are regularly found either at the basis of epidermal cells or within the mesoglea, whereas symbionts are located in the endoderm. Buds induced to metamorphose by a bacterial-inducing factor were used to investigate morphological and ultrastructural changes occurring during metamorphosis and scyphistoma morphogenesis. Metamorphosis starts with the settling of a bud, followed by the formation of the pedal disk in which desmocytes, as typical cnidarian adhesive structures, are differentiated. Metamorphosis is completed with the formation of the mouth and tentacles. Whereas metamorphosis of cnidarian planulae implies considerable changes at the cellular level, tissue remodeling processes prevail in bud metamorphosis ofC. andromeda.     

Temporal variation of Tubularia indivisa (Cnidaria, Tubulariidae) and associated epizoites on artificial habitat communities in the North Sea

We have analyzed the composition, diversity, density and biomass of a temporal series of samples taken in a Tubularia indivisa community, which dominates a shipwreck in the North Sea waters (N 51°23′,730–E 02°29′,790, 17 nautical miles from the coast, 30 m depth). This shipwreck has structures emerging up to 8 m above the seabed. Water temperature ranged from 4.2°C in March to 20.3°C in August. Salinity showed few variations around 33.9 psu. Bottom tidal currents followed a semi-diurnal cycle and were preferentially NE oriented with 84% of them in the range 0.25–0.75 m s⁻¹. The mean value for total suspended matter was 6.2 mg l⁻¹ with large variations on a monthly scale. The species richness of samples varied from 15 in October to 42 in August with a mean value of 33 species. Diversity indices were higher during autumn and winter because of the strong dominance of a few crustacean species during the warmer months. The total density of individuals ranged from 6,500 ind m⁻² in October to 445,800 ind m⁻² in July, most of these individuals belonging to the amphipod species Jassa herdmani. The biomass of the T. indivisa community varied from 9 g AFDW m⁻² in October to 1,106 g AFDW m⁻² in July, with T. indivisa itself constituting between 59 and 82% of the total biomass. The biomass of T. indivisa was positively correlated with species richness and with the density of 23% of the species identified on this community, suggesting that T. indivisa plays an important structural role in this habitat. This was further confirmed by the number of species associated with T. indivisa which was generally superior to 55% of the sorted species. Multivariate analysis indicated strong differences between spring/summer−autumn/winter assemblages mostly but not solely due to the abundance patterns of species. These findings support the conclusion that shipwrecks in Belgian waters allow the development of assemblages dominated by a high biomass of T. indivisa which in turn provides shelter for high densities and biomass of epizoites. These assemblages will further show large monthly variations in densities and composition due to large variation in T. indivisa biomass under an apparent repetitive annual cycle. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Predatory behavior ofPhacellophora camtschatica and size-selective predation uponAurelia aurita (Scyphozoa: Cnidaria) in Saanich Inlet,British Columbia

Predation uponAurelia aurita byPhacellophora camtschatica was studied by SCUBA divers in a fjord in British Columbia between July and September 1986. The behavior and size ofP. camtschatica affected their foraging success. Larger predators captured more and larger prey. The size and behavior of the prey also affected the probability of capture. Predators were found at densities which can affect both the size composition and the overall numbers of the prey population.     

Energetics,competency, and long-distance dispersal of planula larvae of the coral Pocillopora damicornis

Pocillopora damicornis (Linnaeus) were collected from Enewetak Atoll, Marshall Islands, in 1980–1981) and Kaneohe Bay, Hawaii, in 1982. Their planula larvae contained 17% protein, 70% lipid, and 13% carbohydrate by dry weight. Calculations based on stored energy reserves and daily metabolic expenditure indicate that planulae could survive approximately 100 d and still settle successfully. Competency experiments demonstrated that larvae settled and metamorphosed after 103 d. This period of time is sufficient to allow immigration of larvae from the Central Pacific to the eastern Pacific, and supports the hypothesis of long-distance dispersal of larvae for the origin of present eastern Pacific populations of P. damicornis.     

Diel variability of feeding activity in haddock (Melanogrammus aeglifinus) larvae in the East Shetland area, North Sea

Investigations of factors affecting feeding success in fish larvae require knowledge of the scales of variability of the feeding process itself and the indices used to assess this variability. In this study, we measured short-term (diel) variability in feeding rates of wild haddock (Melanogrammus aeglifinus) larvae four times per day during a 10-d cruise in the northern North Sea. Feeding activity was evaluated using indices of gut fullness, prey digestive state and biochemical measurements (tryptic enzyme activity). The gut fullness and the enzyme activity indices indicated moderate to high rates of food consumption throughout the cruise. Time series analysis of the three indices showed significant diel variability in all indices and enabled identification of significant lags between food uptake and peak digestive enzyme activity. The typical pattern of food consumption and digestion was characterized by maximal ingestion of prey early in the evening (19:00 hrs) and peak digestive enzyme activity at 01:00  hrs. The time scale over which enzyme activities reacted to prey ingestion was ca. 6 h, and is consistent with expectations from controlled laboratory experiments with other larval fish species. Significant diel variability in tryptic enzyme activity suggests that attempts to relate this measure of feeding success to other variables (e.g. food concentrations) should take care to accommodate natural cycles in feeding activity before making statistical comparisons. Received: 29 October 1998 / Accepted: 18 June 1999     

Feeding behaviour of North Sea turbot (Scophthalmus maximus) and Dover sole (Solea solea) larvae elicited by chemical stimuli

The effect of chemical stimuli on the feeding behaviour of turbot (Scophthalmus maximus L.) and sole (Solea solea L.) larvae was investigated under controlled laboratory conditions. Laboratory-reared flatfish larvae exposed to chemical stimuli showed significant differences in the frequency of various behaviour patterns related to feeding (such as swimming, snapping and darting) compared to larvae exposed to blanks of filtrated seawater. The chemical substances Il-asparagine, glycine, inosine 5-monophosphate and betaine evoked the strongest behavioural responses in turbot larvae. In sole larvae the most potent substances were Il-phenylalanine, Il-lysine, Il-asparagine, inosine 5-monophosphate and betaine. These results show that feeding of turbot and sole larvae is influenced by chemosensory processes at an early larval stage, and indicate that chemoreception may be an integrated part of turbot and sole larvae feeding strategy.     

Recent strandings of the giant jellyfish Rhizostoma luteum Quoy and Gaimard, 1827 (Cnidaria: Scyphozoa: Rhizostomeae) on the Atlantic and Mediterranean coasts

We present reports of sightings of living and stranded specimens of Rhizostoma luteum on the Atlantic coast of Morocco and along the south shore of the Iberian Peninsula in June–July 2012 and in January–February 2013. During summer 2012 and following the dominant currents, the jellyfish first appeared in the Gulf of Cadiz west of the Strait of Gibraltar. Subsequently, seven additional sightings were reported east of the Strait, in the Alboran Sea. In winter 2013, another event of stranded individuals of this species occurred in the Gulf of Cadiz. A phylogenetic analysis performed on the mitochondrial cytochrome c oxidase I (COI) gene sequence in specimens from both stranding events confirmed the morphological classification, ratifying that R. luteum differs from Rhizostoma octopus and Rhizostoma pulmo. This study records the presence of this species for the first time in 60 years.