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     

Chemical aspects of mass spawning in corals. I. Sperm-attractant molecules in the eggs of the scleractinian coral Montipora digitata

This paper provides the first evidence for sperm chemotaxis in the Scleractinia. Montipora digitata Dana, 1845 (Scleractinia: Coelenterata) is a hermaphroditic coral which reproduces bi-annually, releasing egg-sperm bundles during the mass spawning at Magnetic Island (19°10S; 146°52E) in late spring-early summer, and autumn each year. The buoyant egg-sperm bundles float to the surface where they break apart, releasing eggs and sperm into the ocean. Fertilisation occurs after 30 min. Unfertilized eggs were collected, washed free of sperm, and freeze-dried. The eggs were extracted with dichloromethane, fractionated by chromatography on silica gel, and the fractions assayed for their ability to attract M. digitata sperm. The active fraction was further fractionated by high-performance liquid chromatography, resulting in the isolation of three highly unsaturated fatty alcohols: (1) dodeca-2,4-diynol; (2) tetradec-13-ene-2,4-diynol; (3) (14Z)-heptadeca-14,16-diene-2,4-diynol. Of these three compounds, only Compound 1 attracted sperm of M. digitata. Synthetic Compound 1, produced from simple precursors by known reactions, possessed sperm-attracting activity comparable to the naturally derived attractant. Preliminary experiments suggest that the natural mixture of Compounds 1, 2 and 3 in the ratio 1:4:9 is more effective in attracting sperm from M. digitata than sperm from other Montipora species. Sperm attractants may act to reduce the incidence of hybridisation between different species of Montipora.     

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.     

Effect of nutrient enrichment on the complementary (secondary) metabolite composition of the soft coral Sarcophyton ehrenbergi (Cnidaria: Octocorallia: Alcyonaceae) of the Great Barrier Reef

 A long-term study of the effects of nutrient enrichment on coral reefs (ENCORE Experiment) was carried out at One Tree Island (23°30′S; 152°96′E), Great Barrier Reef, between 1992 and 1996. The experiment involved the addition of water-soluble nutrients to 12 microatolls which contained a range of organisms and were situated within the larger lagoon. Three replicates of each of three nutrient treatments (nitrogen, phosphorus, nitrogen plus phosphorus) and an untreated set of three control atolls were prepared using the 12 selected microatolls. As part of the larger ENCORE experiment, changes in the chemical composition of the alcyonacean soft coral Sarcophyton ehrenbergi Marenzeller (Octocorallia: Alcyonacea), placed in the treatment microatolls, were monitored for a 1 yr period in an attempt to detect any responses attributable to nutrient enrichment. Multivariate analyses were performed to determine whether there were any patterns of response in the different nutrient treatments. At the level of individual metabolites, there were no clear treatment effects. However, the ratio of bioactive or stress metabolites (terpenes) to energy storage metabolites (lipids), referred to as the “physiological-change indicator”, revealed effects of nutrient enrichment. Nitrogen enrichment resulted in a trend towards higher physiological-change indicators than control or phosphorus treatments in the majority of cases, while phosphorus enrichment significantly decreased the ratio relative to controls. In most cases, the physiological-change indicator increased in soft soft corals relocated into contact with the scleractinian Pocillopora damicornis. The potential of soft corals to serve as indicators of a changed nutrient regime is discussed. Received: 23 November 1998 / Accepted: 23 September 1999     

Transformation of soft coral (Coelenterata: Octocorallia) terpenes by Ovula ovum (Mollusca: Prosobranchia)

The faecal pellets from specimens of the prosobranch mollusc Ovula ovum found feeding on the soft coral Sarcophyton sp. at Eclipse Island, Palm Island Group (18°46S; 146°33E) in November 1980 were analysed. The only terpene present in the faeces, 7,8-deoxysarcophytoxide, differed from the major constituent of the soft coral, sarcophytoxide, suggesting that the latter had been transformed into the former within the cowrie. This transformation is not trivial, and could not be produced simply by acid catalysis. Subsequent analysis of tissues dissected from different regions of O. ovum indicates that the transformation is probably effected by enzymes in the digestive diverticula stomach region of the prosobranch. The transformed compound is significantly less toxic to the mosquito fish Gambusia affinis Baird and Girard than the ingested compound.     

Seasonal cycle of sexual reproduction of the Mediterranean soft coral Alcyonium acaule (Anthozoa, Octocorallia)

Alcyonium acaule (Cnidaria, Octocorallia) is a common, hard-bottom soft coral in the northwestern Mediterranean Sea. This study describes sexual reproduction and the gamete development cycle of this soft coral. A population at 15–18 m depth in the Marine Protected Area of the Medes Islands (42º02′55″ N, 3º13′30″ E) was sampled from July 1994–August 1995. A. acaule is gonochoristic and a surface brooder, spawning once a year in early summer. The mean diameter of ripe spermatic sacs was 400 ± 91 (SD) μm, and the mean diameter of mature oocytes was 473 ± 37 (SD) μm. There were 30 spermatic sacs polyp^?1 in males and 14 oocytes polyp^?1 in females. Different phases of gametogenesis in female and male colonies were examined separately with respect to seasonal changes in bottom temperature and solar irradiance. The data suggest that the relatively constant temperatures in January–April are probably not related to oocyte maturation, but that rising temperatures in May could affect sperm maturation. Rapidly increasing solar irradiance in March may be the trigger for vitellogenesis and oocyte maturation, although the mechanism for this in anthozoans is not understood.     

Effects of diterpenes derived from the soft coral Sinularia flexibilis on the eggs,sperm and embryos of the scleractinian corals Montipora digitata and Acropora tenuis

Sinularia flexibilis Quoy and Gaimard, 1833 is a dominant soft coral on many Indo-Pacific coral reefs, and has been found to release toxic compounds (diterpenes), which cause tissue necrosis and death in nearby scleractinian corals. This study investigates how S. flexibilis-derived diterpenes inhibit the development of Acropora tenuis Dana, 1846 and Montipora digitata Dana, 1846 eggs and larvae in vitro. Collection and experimental sites at Magnetic (146°49'E; 19°8'S) and Orpheus (146°28'E; 18°32'S) Islands, Queensland, Australia, were utilized during the spawning seasons of 1989–1992. Freshly spawned coral eggs were placed in solutions of three different terpenes, flexibilide, dihydroflexibilide and sinulariolide, at 5 and 10 ppm, before, during, and after fertilisation. The majority of eggs which were fertilised in the presence of the diterpenes lost their cellular integrity and burst just a few hours after treatment. Terpenes were not toxic to unfertilised eggs, nor to 24 h-old embryos, although sperm ceased swimming activity after 1 h of treatment. The terpenes were not fatal to the sperm because fertilisation still occurred in their presence. The ability of the soft coral-derived diterpenes to inhibit cell division suggests that they may have potential applications in cancer chemotherapy.     

Movement in soft corals: An interaction between Nephthea brassica (Coelenterata: Octocorallia) and Acropora hyacinthus (Coelenterata: Scleractinia)

An interaction between the alcyonarian octocoral Nephthea brassica Kukenthal, 1904 and the scleractinian Acropora hyacinthus Dana was photographed and the assemblage collected at Rib Reef (18°15S; 146°45E), Central Region of the Great Barrier Reef, in September 1980. The interaction, which did not involve allelopathy, was characterised by extensive deposition of undifferentiated aragonite by the scleractinian in the vicinity of the soft coral, which moved across the surface of the acroporid. The basal attachment region of the nephtheid showed structural features which permitted movement. The net effect of this interaction was a series of divided trails across the surface of the acroporid plate evidencing the movement through growth and asexual division of the parent colony. These trails were susceptible to secondary colonisation by epifauna and algae which may reduce the fitness of the acroporid. This movement appears to afford N. brassica a distinct advantage in competing for space with acroporid corals.     

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.     

Chemical consequences of relocation of the soft coral Lobophytum compactum and its placement in contact with the red alga Plocamium hamatum

Seven colonies of Lobophytum compactum Tixier-Durivault, 1956, which produce isolobophytolide as the major secondary metabolite, were selected from a fringing reef in the Pelorus Channel, Palm Island Group (18°34S; 146°29E), North Queensland, Australia. In September 1991, they were sectioned to afford two portions which were relocated to a grid, and a significant part of the parent colony which was left in place. The aim of the experiment was to determine the effect of relocation and contact with a toxic alga on the secondary metabolite content of a soft coral. A significant increase in the concentration of isolobophytolide was observed for all relocated colonies (n=14, p=0.001) compared to the non-relocated control colonies. This decreased after 2 mo, and was not significantly different from that of the non-relocated control colonies (n=14, p=0.881). After 1 mo, Plocamium hamatum J. Agardh plants were placed in direct contact with 50% of the relocated colonies. All soft-coral colonies in contact with the alga (n=7), showed tissue necrosis on the parts in direct contact with the alga after a further 2 wk. Tissues of the relocated control colonies (n=7), and those portions of treated colonies which were not in direct contact with the alga, were not affected. The parts of the colonies in contact with the alga showed a significant decrease in lipid content over time (n=7, p=0.001) and also a decrease in the concentration of the diterpene isolobophytolide (n=7, p=0.001). The effects of P. hamatum on the soft coral were essentially restricted to contract necrosis; chemical variations in the affected tissue were the outcomes of this necrosis. These results indicate that stress due to relocation is a more important factor in the variation of isolobophytolide levels in the soft coral L. compactum than is contact with the alga P. hamatum.     

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     

Multiple spawning events and sexual reproduction in the octocoral Sarcophyton elegans (Cnidaria: Alcyonacea) on Lizard Island, Great Barrier Reef

Sarcophyton elegans is a common symbiotic (zooxanthellate) octocoral species in the shallow waters of the Great Barrier Reef (GBR). Study of a population at Lizard Island (14°40′S, 145°28′E) on the GBR from October 1991 to January 1994 revealed that, as is typical of tropical alcyonarian corals, S. elegans is a gonochoric broadcast spawner with a 1:1 sex ratio. Sexual reproduction was closely correlated with colony size, with first reproduction at 13-cm basal stalk circumference for females and 12 cm for males. Oogenesis took 19–24 months, with a new cycle commencing every year, and spermatogenesis took 10–12 months. The majority of gametes were released during the annual austral mass coral spawning event after the full moon in November, but gametes were also released after the full moon in each month between August and February. All autozooid polyps participated in reproduction, but those at the outer edge of a colony released their gametes first. During subsequent months, the polyps closer to the center of the colony released their gametes. This is a novel strategy of gamete release, reported here for the first time, which accommodates the demands of feeding and reproduction in a different way than other corals where individual polyps have separate feeding or reproductive roles. Colonies upstream in the prevailing current spawned up to 1 month earlier than those downstream and ceased 1 month earlier. The mechanism controlling this spatial differentiation in spawning time, repeatedly observed over three seasons, is unknown. Sarcophyton elegans appears to have a dual strategy of providing protection for its gametes by releasing most of them concurrently with the single, annual mass spawning of a large number of cnidarians, while also hedging its bets by individual colonies spawning a fraction of their gametes over an extended period of 6 months.     

Behavioral aspects of sperm competition in cuttlefish, Sepia officinalis (Sepioidea: Cephalopoda)

Sexual selection studies in cephalopods indicate that sperm competition is a central feature of their mating systems, yet this has not been studied experimentally in any detail. In 1998 we staged 20 matings of the cuttlefish, Sepia officinalis L., in the laboratory. Males rapidly initiated mating in the “head-to-head” position, with no apparent courtship. Mating lasted an average of 10 min (range 7 to 14 min). For the first 6 min (on average 63% of the mating duration), the male flushed strong jets of water directly at the female's buccal membrane, which sometimes resulted in the expulsion of parts of spermatangia placed there in recent matings. Then, in a single discrete movement that lasted an average of only 14 s, the male's modified fourth left arm – the hectocotylus – wrapped around a single large bundle of spermatophores (ca. 150 to 300) and transferred them to the female's buccal membrane. For the remainder of the mating (average 3 min, range 1.5 to 5.0 min), the hectocotylus repeatedly broke the spermatophores open, and manipulated them, so that sperm were released and many spermatangia were attached along the ventral buccal membrane, near the paired seminal receptacles. Approximately 140 spermatangia were attached in rows 3 to 5 deep around the ventral buccal membrane in a single mating; the rest were usually discarded during mating. Histology revealed that each of the seminal receptacles consists of a series of sperm storage bulbs connected by a central duct, which leads to a single pore at the surface of the buccal membrane. Baseline data on sperm motility were obtained, but the mechanism by which sperm enter the seminal receptacle remains unknown. Females seemed to initiate termination of mating, then males guarded their mates temporarily. These results, combined with other recent laboratory experiments, provide evidence that sperm competition may be a major feature of the mating system of S. officinalis. Received: 4 January 1999 / Accepted: 18 May 1999     

Comparative aspects of sexual reproduction in the Caribbean coral genus Diploria (Scleractinia: Faviidae)

The increasing decline of Caribbean reef-building species underlines the importance of more information on their reproductive biology and ecology. This study compares the reproductive biology of the three species of Diploria, an important Caribbean reef-building genus affected by bleaching and disease, by: (a) characterizing their gametogenetic cycles simultaneously, (b) exploring the spatial and temporal variability in reproductive traits within and across species, (c) assessing fecundity and the minimum size of sexual reproduction, and (d) comparing information with other studies in the Caribbean. Starting in 1999, one tissue core was collected every month (for 17 months) from each of at least five large, tagged colonies of each species in La Parguera, Puerto Rico. Three more temporally spaced samplings were done during summer of 2000 and winter–spring of 2001 and 2002. Tissues were fixed in Helly’s solution, rinsed in fresh water, decalcified with 10% HCl, and preserved in 70% ethanol, embedded in paraplast, cut, and thin slides (7 μm) prepared and stained with Heidenhain’s Aniline-Blue method. Maximum egg size, number of eggs and spermaries were measured and gametogenetic cycles assessed. Microscopic observations confirmed that all three species were simultaneous hermaphrodites with a single, annual gametogenetic cycle. Spermatocytes and oocytes developed within the same mesentery but were not intermingled. Female gametes developed several months earlier than that of males, but both reached maturation simultaneously. Minor differences in the onset of gametogenesis were found for D. strigosa and D. clivosa, with both spawning after 10 p.m. between 8 and 10 days after the August and/or September full moons. D. labyrinthiformis on the other hand, began oogenesis in early July and spawned after 11 p.m. between 7 and 10 days after the April and/or May full moons. Spatial and temporal variability in mean number of spermaries and eggs/mesentery among colonies within and among species were found. D. labyrinthiformis had consistently significant higher mesenterial and polyp fecundity compared to the other two species over the 3 years. Average egg diameter varied between 240 and 246 μm and was similar among the three species. D. strigosa and D. clivosa sexual reproductive characteristics were similar to those of other large broadcast spawning Faviidae (i.e. Montastraea spp) in the region. In contrast, D. labyrinthiformis was a spring spawner (similar to Colpophyllia natans in Puerto Rico), which together with its significantly different micromorphology suggests that it might be phylogenetically more distant to the other taxa.     

Reproduction in Anthelia glauca (Octocorallia: Xeniidae). II. Transmission of algal symbionts during planular brooding

The soft coral Anthelia glauca Lamarck, 1816, of the family Xeniidae, is found on the reefs of KwaZulu-Natal, South Africa. Its gastrodermal cells contain numerous endosymbiotic unicellular algae (zooxanthellae). A. glauca is a gonochoric species that simultaneously broods its planulae within the pharyngeal cavity of the polyps. Symbiotic algae appear with zygote formation within the pharynx, embedded in amorphous material. The algal cells adhere to the ciliated ectodermal surface of immature planulae and are most probably endocytosed by them. Zooxanthellae are translocated towards the basal part of the ectoderm. Gaps are subsequently opened in the mesoglea into which symbionts surrounded by ectodermally derived material, including plasma membrane, pass. The basal membrane of endodermal cells disintegrates, and the algae bulge into spaces formed in the underlying endoderm. Throughout the process, each zooxanthella resides within a vacuolar membrane in the detached ectodermal cytoplasm. The acquisition process is essentially one in which zooxanthellae are translocated from the pharyngeal cavity into the ectoderm and then through the mesoglea into the endoderm, culminating in the final symbiotic state. The direct transmission of symbiotic algae to the eggs or larvae probably provides the most efficient means whereby zooxanthellae are acquired by the host progeny. Received: 15 July 1997 / Accepted: 25 February 1998     

Distribution,abundance and survival of juvenile hermatypic corals (Scleractinia) and the importance of life history strategies in the parent coral community

The distribution and abundance of juvenile corals were examined at depths from 3 to 37 m on the reefs of Curaçao and Bonaire (Netherlands Antilles). Juveniles of Agaricia agaricites were most abundant (60.6%), followed by Helioseris cucullata (8.3%). The large massive corals such as Montastrea annularis, M. cavernosa and branched species such as Madracis mirabilis and Acropora palmata had few juveniles. This, combined with species characteristics, shows that these species employ very different life history strategies. In some species the abundance of juveniles over the reef paralleled that of larger colonies, but not for example in Agaricia agaricites. The composition of the coral community was apparently no direct function of juvenile abundance. A change in angle of settlement of A. agaricites juveniles with increasing depth, from vertical to horizontal surfaces, seems to reflect the preferred light intensity. We studied the survival of juvenile corals during a half-year period. One-third remained unharmed, one-third died or disappeared, and one-third was limited in growth by factors such as spatial competition. This was the same for all depths, but factors influencing survival varied with depth.     

Effects of the 1996 and 1998 positive sea-surface temperature anomalies on corals,coral diseases and fish in the Arabian Gulf (Dubai,UAE)

Two positive sea-surface temperature anomalies occurred in the Arabian Gulf in short sequence. Between May and August 1996 and 1998, sea-surface temperatures in the southern Arabian Gulf were elevated by 2°C above average. The consequences for coral fauna, coral diseases and coral regeneration were studied in Dubai (United Arab Emirates) between Jebel Ali and Ras Hasyan. In 1996, coral death was widespread, affecting primarily the genus Acropora. In Acropora-dominated areas, live coral cover was reduced from 90% to about 26% in 1996, while in 1998 only a reduction from 26% to 22% of the remaining coral cover occurred. In the study area, all six Acropora species suffered total mortality in 1996, thus the coral fauna was reduced from 34 species to 27. The nearest areas with surviving Acropora were 30 km to the east (Deira) and 20 km to the west (Al Jazira). Massive coral species suffered negligible mortality, and slowly increased in space cover. The Acropora overkill turned 7.9 km² (19.7% of total coral-covered area) of previously lush coral gardens into a dead framework that was increasingly bioeroded. Acropora recruitment only started in 1998, average recruit size in 1999 was 7Dž cm, and recruits were rare. Prior to the mass mortality event, coral diseases were common and seasonal (14LJ% of corals, mainly Acropora, affected in summer, in winter 7Lj%, mainly massives), after the mortality event seasonality was lost and infection remained below winter levels (6LJ%, only massives infected). In fish, overall species richness decreased from 95 to 64 species in point counts, but frequency only decreased in one species (Pseudochromis persicus). Guild structure changed inasmuch as herbivores and planktivores increased, and invertivores decreased, although differences were not statistically significant. The most abundant family, both prior to and after the coral mass mortality, was Lutjanidae. It appears that even though much of the coral was dead, the maintenance of structural complexity allowed the fish assemblage to avoid a similar catastrophic change to that experienced by the coral assemblage.     

Chemical defense in the eggs and embryos of antarctic sea stars (Echinodermata)

Fifteen species of reproductively mature echinoderms (11 sea stars, 3 sea urchins, 1 sea cucumber) were collected from McMurdo Sound, Antarctica, during austral spring and summer of 1985 and 1986; eggs and embryos were obtained, and were tested for ichthyonoxicity using the common marine killifishFundulus grandis as a model predator. Chemical deterrents occurred in the large, yolky eggs of the pelagic lecithotrophic sea starPerknaster fuscus and the planktotrophic sea starPorania antarctica. Brooded embryos of the sea starsDiplasterias brucei andNotasterias armata were also noxious. Significant ichthyonoxicity was not detected in the remaining 7 species of sea stars, 3 sea urchins, and 1 sea cucumber. Chemical deterrents were generally effective at concentrations below a single egg or embryo per agar test-pellet. Although chemicals found in these eggs and embryos are noxious to an allopatric fish, they may not be noxious to sympatric fish.     

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.