Reproduction of Dendronephthya hemprichi (Cnidaria: Octocorallia): year-round spawning in an azooxanthellate soft coral

Sexual reproduction in the azooxanthellate octocoral Dendronephthya hemprichi Klunzinger, 1877 was studied in Eilat (Red Sea) for 2 years beginning March 1989. D. hemprichi is a gonochoric broadcasting species. Gonads at all developmental stages were found throughout the year. Small-sized oocytes and sperm sacs, 51 to 100 μm in diameter, are highly abundant, accompanied by numerous smaller primordial gonads. These features result from continuous gametogenesis, and lead to year-round oocyte and sperm maturation and release. Gamete release was observed in the laboratory on all monitoring dates, and most of the female colonies spawned repeatedly for several successive nights. The reproductive features of D. hemprichi constitute an exception to the generalization that free-spawning reef-corals have brief and synchronized broadcast-spawning episodes. This species demonstrates early onset of first reproduction, which further increases its reproductive output. Received: 7 May 1997 / Accepted: 15 May 1997     

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     

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.     

Hydrolithon spp. (Rhodophyta, Corallinales) overgrow live corals (Cnidaria, Scleractinia) in Yemen

In Yemen, off the northwestern coast in the Gulf of Aden, the coralline algae Hydrolithon rupestre (Foslie) Penrose 1996 and H. murakoshii Iryu and Matsuda 1996 have been observed to overgrow and kill living Porites lutea Milne-Edwards and Haime, 1860. Similarly, Hydrolithon onkodes (Heydrich) Penrose and Woelkerling 1992 and H. rupestre were observed overgrowing Stylophora pistillata (Esper, 1797). Competitive interactions between P. lutea and H. murakoshii were monitored from 2006 to 2009 at two sites and showed an average linear growth of 8.3 (±1.9 SD) mm?year^?1 over the coral. The small polyps of S. pistillata and P. lutea combined with putative chemical compounds produced by Hydrolithon spp. are likely to allow the coralline overgrowth. Although corallines can locally kill coral tissues, the CCA/coral interactions do not seem to affect the overall live coral cover at the study sites.     

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.     

Sexual reproduction in the coral Pocillopora verrucosa (Cnidaria: Scleractinia) in KwaZulu-Natal, South Africa

The mode of reproduction in Pocillopora verrucosa Ellis and Solander, 1786 varies between geographically isolated regions. This scleractinian coral is common along the KwaZulu-Natal coast, and its reproductive mode and period of reproduction were assessed using histological preparations. The study was undertaken from 1992 to 1994 and showed that P. verrucosa is a simultaneous hermaphrodite and broadcast spawner in KwaZulu-Natal. Gametogenesis occurs from October to January, with the gametes maturing simultaneously so that spawning can be synchronised at new moon in January (mid-summer). Zooxanthellae are present in the mature oocytes. Received: 10 September 1997 / Accepted: 6 July 1998     

Population dynamics of Eudendrium glomeratum (Cnidaria: Anthomedusae) on the Portofino Promontory (Ligurian Sea)

Eudendrium glomeratum Picard, in the Ligurian Sea, is one of the major components of hard-bottom sessile zoobenthos in the cold season. It settles mainly between 10 and 40 m depth, forming a seasonal facies. The presence of E. glomeratum has been evaluated by measuring in situ the height of the colonies present within a standard surface of 1 m². Observations were carried out every other week for a whole year. The height of the colonies was correlated with their biomass (wet weight) and trophic functionality (number of polyps). The quantitative evaluation of the population shows that the situation is stable, with almost identical values at the extremes of the observed yearly time series. The species is present from October to April–May, and it is absent in the summer. The formation and disappearance of the E. glomeratum population are sudden. The colonies are sexually mature from their appearance in October to December–January. Different categories of colonies are recognizable; and are essentially referable to two groups: (1) colonies deriving from regeneration of colonies present in the previous year (45%); (2) colonies deriving from planula settlement (43%). A third group (12%) is of uncertain origin and may belong to either of the abovementioned categories. Mortality is 60% in the first group, and 82% in the second one. By dead colonies is meant those which do not reappear after a yearly cycle.     

Foraging, prey capture, and gut contents of the mesopelagic narcomedusa Solmissus spp. (Cnidaria: Hydrozoa)

Narcomedusae are the most common group of medusae in the mesopelagic depths of Monterey Bay, California. Numerous capture events of various prey taxa were recorded in situ and analyzed using the Monterey Bay Aquarium Research Institute's remotely operated vehicle "Ventana". In situ video analysis of the stomach contents of 82 Solmissus incisa and S. marshalli revealed 88 identifiable prey items. Most (88%) were gelatinous animals. Of these gelatinous prey, 60.3% were ctenophores, 20.5% were cnidarians, 12.8% were salps, 3.8% were chaetognaths, and 2.6% were polychaetes. Euphausiids accounted for 11.4% of the diet, but they were probably captured adventitiously, as the gut contents of ctenophore prey. The tentacle-first foraging behavior of the narcomedusae is an effective way to capture large, comparatively fast-moving prey, because the fluid disturbance caused by swimming is decoupled from the area of prey capture. This behavior contrasts with the prevailing models of feeding behavior in medusae. Stealth predation may be the dominant mode of capturing prey by medusae in the mesopelagic depths of the oceans.     

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.     

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.     

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.     

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.     

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.     

Effects of temperature on success of (self and non-self) fertilization and embryogenesis in Diploria strigosa (Cnidaria, Scleractinia)

The effects of seawater temperature on the degree of success of self-fertilization versus cross-fertilization and on embryogenesis were investigated in the scleractinian coral Diploria strigosa (Dana 1846). Gametes from nine colonies were collected from the Flower Garden Banks reefs, a set of coral reefs in the northern Gulf of Mexico, ~110 nm offshore from Texas, United States. Gametes from the coral colonies were combined in all possible combinations and placed in 30°, 31°, and 32°C water baths on successive nights, respectively. Each cross, whether selfed or out-crossed, and each temperature treatment, was replicated twice. A high frequency of successful self- and cross-fertilization was observed at all temperature treatments and in both blocks. Higher temperatures, however, commonly produced numerous developmental aberrations during embryogenesis of the larvae. Thus, although fertilization rates can remain high under high temperature conditions, if temperatures remain high for several days, embryonic development and larval viability may be expected to decrease dramatically. We propose that the success of coral larval development may be diminished in areas where abnormally high sea surface temperatures occur during the spawning season. We also propose that highly successful selfing may enhance the abundance of locally adapted genotypes, which in turn may be advantageous where reefs, such as the Flower Garden Banks, are relatively geographically isolated.     

Isolation and cloning of DNA from somatic tissue of soft corals (Cnidaria: Octocorallia)

A method is described for isolating high molecular weight DNA from somatic tissue of soft corals. The method avoids problems associated with the presence of nucleases, pigments and other secondary metabolites in soft corals. Tissue is frozen in liquid nitrogen, pulverized with a pestle and mortar, and the powder extracted with buffered sodium dodecyl sulphate at 4°C in the presence of phenol/chloroform. This technique has been applied toAlcyonium, Sinularia, Sarcophyton andLobophytum species. Repeated sequences have been cloned fromAlcyonium sp., and used to probe slot-blots of genomic DNA and Southern blots of restriction digests. Homologous repeated sequences were detected in threeAlcyonium sp., but not in threeSinularia sp., despite these genera being closely related.     

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.     

Use of electrical stimulation for discharging cnidom components of a species of Cerianthus (Anthozoa: Cnidaria)

Direct-current electrical stimulation was used as a means of discharging cnidom components from the marginal tentacles of a species of Cerianthus from Southern Queensland, Australia, in order to facilitate the study of cnidom components by light microscopy. The cnidom consists of b-rhabdoids (3 types), spirocysts, and ptychocysts. Threshold pulses of between 1 and 2 V with a frequency of 1 pulse per second for 2 sec discharge all types of cnidae. However, large numbers of certain cnida types discharge after a longer period of stimulation.