Marginal tentacles of the corallimorpharian Rhodactis rhodostoma. 2. Induced development and long-term effects on coral competitors

Polyps of the corallimorpharian Rhodactis rhodostoma (Ehrenberg, 1934) form aggregations that monopolise patches of space on the shallow reef flats of some Red Sea coral reefs. Some of these polyps bear specialised bulbous marginal tentacles (BMTs) where they contact cnidarian competitors. BMTs differ from the normally filiform marginal tentacles (FMTs) of R. rhodostoma, and appear to develop from them. However, their morphogenesis and long-term impacts on spatial competition with reef corals are unknown. We experimentally induced contacts between R. rhodostoma polyps and colonies of the branching stony coral Acropora eurystoma on a shallow coral reef at Eilat, northern Red Sea. During the first 24 d of contact, the A. eurystoma colonies extruded mesenterial filaments that damaged the tissues of the corallimorpharian polyps. After 18 d,>90% of R. rhodostoma individuals had developed BMTs, which resulted in a reversal in the direction of competitive damage. During the subsequent 1.5 years of observation, the corallimorpharians maintained well-developed BMTs, unilaterally damaged the tissues of A. eurystoma, and in some cases moved onto the stony coral skeletons and partially overgrew them. BMTs developed from FMTs in a series of four distinct stages, accompanied by significant changes in their morphology, cnidom, and density of nematocysts. Isolated control polyps did not develop BMTs or show any signs of damage. In contrast, corallimorpharian polyps transplanted into contact with colonies of the massive stony coral Platygyra daedalea began to develop sporadic BMTs, but were unilaterally and severely damaged by the corals, and started to disappear within 21 d, after the corals developed sweeper tentacles. We conclude that long-term outcomes of competition between R. rhodostoma and reef-building corals depend largely on the relative aggressive reach of the competitive mechanisms developed by each species. As a consequence, this corallimorpharian is an intermediate competitor in the aggressive hierarchy among Indo-Pacific reef corals. This study confirms that R. rhodostoma polyps may actively damage and overgrow some stony corals, leading to the formation of an almost continuous blanket of polyps in large patches of some shallow reef flats. Received: 15 July 1998 / Accepted: 24 March 1999     

Adaptation strategies of the corallimorpharian <Emphasis Type="Italic">Rhodactis rhodostoma</Emphasis> to irradiance and temperature

Corallimorpharians may dominate some habitats on coral reefs and compete with stony corals for access to light, yet little is known concerning their photosynthetic traits. At Eilat in the northern Red Sea, we observed that the abundance of individuals of the corallimorpharian Rhodactis rhodostoma decreased significantly with depth on the reef slope. Field and laboratory experiments revealed that they employ several mechanisms of photoadaptation to high irradiance on the shallow reef flat. Their endosymbiotic microalgae (zooxanthellae) varied significantly in both abundance and chlorophyll content with level of irradiance. Use of a diving pulse amplitude modulated fluorometer revealed that the zooxanthellae of R. rhodostoma effectively disperse excess light energy by expressing significantly higher values of non-photochemical quenching and maximum excitation pressure on photosystem II when experimentally exposed to high light (HL) versus low light (LL). Host corallimorpharian tissues mediated this response by shielding the algal symbionts from high irradiance. The endoderm of host tentacles thickened significantly and microalgal cells were located further from the mesoglea in HL than in LL. The clades of zooxanthellae hosted by the corallimorpharians also varied with depth. In shallow water, all sampled individuals hosted clade C zooxanthellae, while in deep water the majority hosted clade D. The photosynthetic output of individuals of R. rhodostoma was less affected by HL than was that of a stony coral examined. When exposed to both high temperature (HT) and HL, individuals of R. rhodostoma reduced their maximum quantum yield, but not when exposed to HL at low temperature (LT). In contrast, colonies of the scleractinian coral Favia favus reduced their photosynthetic output when exposed to HL in both temperature regimes. After 2 weeks of HT stress, R. rhodostoma polyps appeared to bleach completely but re-established their zooxanthella populations upon return to ambient temperature. We conclude that mechanisms of photoadaptation to high irradiance employed by both the endosymbiotic zooxanthellae and host corallimorpharians may explain in part the abundance of R. rhodostoma on some shallow reef flats. The ability to survive for weeks at HT while bleached also may allow corallimorpharians to repopulate shallow reef areas where scleractinians have been killed by thermal stress. B. Kuguru and G. Winters contributed equally to this work.     

The reef environment and competitive success in the Corallimorpharia

Competitive success within coral reef communities is controlled by various factors. In addition to competitive abilities in direct interactions with a contestant, external influences such as disturbance caused by nutrient input may determine the outcome of antagonistic interactions. We examined the competitive success of corallimorpharians on coral reefs by investigating their distribution patterns within reefs and how well they perform in interference competition with staghorn corals in different environments. Substrate composition and corallimorpharian growth were examined on three reefs in Tanzania under different disturbance regimes using the line-intercept transect and point techniques. A transplant experiment was conducted in which staghorn corals (Acropora formosa) were exposed to the polyps of Rhodactis rhodostoma to establish how competition between corals and corallimorpharians affects their respective distributions. Within reefs corallimorpharians seemed to be more competitive in shallow waters. This could be due to both environmental factors as well as varied competitive abilities depending on surrounding benthos that changed with depth. Reef environment also seemed to influence corallimorpharian growth among reefs as they had the highest densities in the areas with the highest nutrient loads. The transplant experiment revealed that the corallimorpharians had a competitive advantage over the corals, and in comparisons of reefs influenced by different degrees of disturbance, corallimorpharians were most competitive in the area with the highest nutrient content. Hence, stress on coral reefs in the form of raised nutrient loads may favour the competitive success of corallimorpharians.Communicated by P.W. Sammarco, Chauvin     

Migration,habitat use,and competition among mobile corals (Scleractinia: Fungiidae) in the Gulf of Eilat,Red Sea

We examined the impact of seven species of mobile mushroom corals (Fungiidae) on the community structure of sheltered reef slopes in terms of their patterns of migration, habitat use and competition with other benthic organisms. On fringing reefs at Eilat, Red Sea, polyps detached at 1 to 6 cm length, and grew to 11–55 cm length. Attached mushroom corals were oriented vertically in reef cavities. Detached corals migrated downward on the reef slope and onto rubble or soft substratum at the reef base, at 29 to 71 cmyr^–1. Mobility decreased with corallum size and extent of undersurface ornamentation. In aquaria, small corals righted themselves and migrated up to 6 cm d^–1 by nocturnally inflating and pushing their tissues against the substratum. Autonomous coral behavior and storm-generated water motion appeared to account for most fungiid mobility at Eilat. Mushroom corals did not damage each other upon contact, even in multi-species aggregations, but unilaterally damaged non-fungiid scleractinian corals. Their dominance during contact interactions retards overgrowth by larger attached scleractinians, and mobility allows them to colonize soft substrata not accessible to most other reef corals.     

Analysis of ribosomal ITS1 sequences indicates a deep divergence between Rhodactis (Cnidaria: Anthozoa: Corallimorpharia) species from the Caribbean and the Indo-Pacific/Red Sea

This paper explores the implications of ribosomal ITS1-sequence analysis for evolutionary relationships between four species of corallimorphs which are morphologically very similar but have distinct distributions. The analyses highlight the unreliability of morphological criteria in establishing systematic relationships amongst tropical corallimorphs. Both parsimony and distance methods of phylogenetic analysis strongly support a deep divergence between the Caribbean species Rhodactics sanctithoma and the three morphologically-related species R. howesii and R. indosinensis from the Indo-Pacific and R. rhodostoma from the Red Sea. The analyses also strongly support a closer relationship between the Red Sea species, R. rhodostoma, and one of the Indo-Pacific species, R. howesii, than between these two and the second Indo-Pacific species, R. indosinensis. The first of these results is presumably a consequence of the closure of the Central American Seaway, the event which led to the separate development of Atlantic and Indo-Pacific coral faunas. However, the lack of detailed distribution data represents a major barrier to reconstructing the evolutionary history of the genus.     

Predators selectively graze reproductive structures in a clonal marine organism

Although the fitness consequences of herbivory on terrestrial plants have been extensively studied, considerably less is known about how partial predation impacts the fitness of clonal marine organisms. The trophic role of Caribbean parrotfish on coral reefs is complex: while these fish are important herbivores, as corallivores (consumers of live coral tissue), they selectively graze specific species and colonies of reef-building corals. Though the benefits of parrotfish herbivory for reef resilience and conservation are well documented, the negative consequences of parrotfish grazing for coral reproductive fitness have not been previously determined. We examined recently grazed colonies of Montastraea annularis corals to determine whether grazing was positively associated with coral reproductive effort. We measured gonad number, egg number and size, and proportional reproductive allocation for grazed and intact coral colonies 2–5 days prior to their annual spawning time. We found that parrotfish selectively grazed coral polyps with high total reproductive effort (number of gonads), providing the first evidence that parrotfish selectively target specific tissue areas within a single coral colony. The removal of polyps with high reproductive effort has direct adverse affects on coral fitness, with additional indirect implications for colony growth and survival. We conclude that chronic grazing by parrotfishes has negative fitness consequences for reef-building corals, and by extension, reef ecosystems.     

Transplantation of juvenile corals: a new approach for enhancing colonization of artificial reefs

 Coral reefs in the northern Gulf of Eilat are exposed to continuous man-made disturbances, resulting in decreased coral coverage and reduced recruitment at the Nature Reserve of Eilat. The construction of artificial reefs on sandy bottoms is a possible option to decrease diving pressure on natural reefs. In the present study we tested this hypothesis by submerging an experimental artificial reef anchored to the bottom at 18 m depth and floated vertically 3 m below water surface. The reef was composed of PVC plates, attached both vertically and horizontally along a wire. Propagules of two coral species, the stony coral Stylophora pistillata and the soft coral Dendronephthya hemprichi, were transplanted to this artificial reef. Planulae of S. pistillata were obtained during the breeding season, seeded in petri dishes in the laboratory and after 2 wk the dishes were transferred to the experimental artificial reef. Automized fragments of D. hemprichi which had previously settled on 10 × 10 cm PVC plates were transplanted onto the experimental artificial reef. The survivorship of the transplanted D. hemprichi colonies was significantly higher on the lower sides of shallower plates. Survivorship of S. pistillata colonies increased with depth when located on the vertical plates, or on the upper sides of the horizontal plates. The highest survivorship of this coral was on the vertical plates and on the upper sides of the horizontal plates, while very low survivorship was recorded on the lower sides. The results indicate that vertical artificial surfaces offer the optimal biotic and abiotic conditions for the survival of the two examined corals. The vertical plates are characterized by low sed imentation rates, low coverage of turf-algae, minimal grazing by sea urchins and absence of the competitor tunicate Didemnum sp. In addition, the vertical orientation of the experimental plates reduces shading and offers the required light intensity for zooxanthellate corals such as S. pistillata. Only a few studies to date have tried to implement artificial reefs in a coral reef environment. The results of the present study indicate the potential of enhancing recruitment of corals by transplantation of juvenile recruits onto appropriate artificial structures. Maximal survivorship of these recruits is dependent upon the structural features of the artificial reef, which should offer optimal conditions. Received: 25 May 1996 / Accepted: 15 July 1996     

Maintenance of living space by sweeper tentacles of Montastrea cavernosa,a caribbean reef coral

The reef-building coral Montastrea cavernosa Linnaeus possesses sweeper tentacles which have enlarged nematocyst batteries. Sweeper tentacles appear to be used in defense of the coral's living space and may successfully deter mesenterial filament attacks from the more aggressive coral M. annularis. M. cavernosa therefore possesses a specialized defensive strategy that has not been taken into account by present models describing spatial competition in coral reef ecosystems.     

Allorathkea ankeli n. gen. n. sp. und Thamnostoma eilatensis n. sp. —zwei neue Hydromedusen (Anthomedusae: Hydrozoa) aus dem Roten Meer

Two new hydromedusae collected in 1970 in the Eilat Bay (Red Sea) are here described as Allorathkea ankeli n. gen. n. sp. and Thamnostoma eilatensis n. sp. A. ankeli (Family Rathkeidae) is characterized by 8 radial canals, dichotomously branched oral arms, and solid marginal tentacles arranged in 8 groups. T. eilatensis (Family Bougainvilliidae) possesses 4 solitary marginal tentacles, 4 interradial gonads and dichotomously branched oral tentacles inserted above mouth-rim.     

Annual density banding in massive coral skeletons: result of growth strategies to inhabit reefs with high microborers’ activity?

Porites and Montastraea are the major reef-building massive coral genera in the Indo-Pacific and Atlantic oceans, respectively. They are also the most commonly used genera in sclerochronological studies. Despite the marked differences in the way these genera use calcareous material to construct their skeletons (growth strategies) and in their skeletal architectural structure, they form annual high and low density bands in their skeletons, that result from the positive relationship of coral calcification rate with sea surface temperature and seasonal changes of the latter. Evidence in the literature suggests that the different growth strategies allow these organisms to construct denser skeletons far from terrigenous inputs, on reefs where microborers’ activity is high. It seems quite probable that this has consequences for the evolution, diversity, distribution and abundance of reef corals.     

Antimicrobial activity of Red Sea corals

Scleractinian corals and alcyonacean soft corals are the two most dominant groups of benthic marine organisms inhabiting the coral reefs of the Gulf of Eilat, northern Red Sea. Antimicrobial assays performed with extracts of six dominant Red Sea stony corals and six dominant soft corals against marine bacteria isolated from the seawater surrounding the corals revealed considerable variability in antimicrobial activity. The results demonstrated that, while the majority (83%) of Red Sea alcyonacean soft corals exhibited appreciable antimicrobial activity against marine bacteria isolated from the seawater surrounding the corals, the stony corals had little or no antimicrobial activity. From the active soft coral species examined, Xenia macrospiculata exhibited the highest and most potent antimicrobial activity. Bioassay-directed fractionation indicated that the antimicrobial activity was due to the presence of a range of compounds of different polarities. One of these antibiotic compounds was isolated and identified as desoxyhavannahine, with a minimum inhibitory concentration (MIC) of 48 μg ml⁻¹ against a marine bacterium. The results of the current study suggest that soft and hard corals have developed different means to combat potential microbial infections. Alcyonacean soft corals use chemical defense through the production of antibiotic compounds to combat microbial attack, whereas stony corals seem to rely on other means.     

Coral recruitment: a spatio-temporal analysis along the coastline of Eilat,northern Red Sea

Recruitment rates of stony corals to artificial substrates were monitored for 2 years at 20 sites along the coast of Eilat, northern Red Sea, to compare with those recorded at other coral reef locations and to assess variation in recruitment at several spatial scales. Coral recruitment was low compared to that observed on the Great Barrier Reef in Australia, but was similar to levels reported from other high-latitude reef locations. Pocilloporids were the most abundant coral recruits in all seasons. Recruitment was twofold higher during the first year than during the second year of study. There was considerable spatial variability, with the largest proportion of variance, apart from the error term, attributable to differences between sites, at a scale of 10² m. Spearmans ranked correlation showed consistency in spatial patterns of recruitment of pocilloporid corals between years, but not of acroporid corals. During spring, when only the brooding pocilloporid coral Stylophora pistillata reproduces at this locality, most coral recruitment occurred at central and southern sites adjacent to well-developed coral reefs. During summer, recruitment patterns varied significantly between years, with wide variation in the recruitment of broadcasting acroporid corals at northern sites located distant from coral reefs. Settlement was low at all sites during autumn and winter. This work is the first detailed analysis of coral recruitment patterns in the Red Sea, and contributes to the understanding of the spatial and temporal scales of variation in this important reef process.Communicated by O. Kinne, Oldendorf/Luhe     

Management of nematocysts in the alimentary tract and in cnidosacs of the aeolid nudibranch gastropod<Emphasis Type="Italic"> Cratena peregrina</Emphasis>

The nudibranch gastropod Cratena peregrina (Opisthobranchia: Aeolidaceae), when feeding on polyps of the hydrozoan Eudendrium racemosum (Cnidaria: Anthoathecata), devours masses of small microbasic eurytele and holotrichous isorhiza nematocysts. Large proportions of these nematocysts were found undischarged in the alimentary tracts of the snails. Feeding experiments in this study tracked the fate of nematocysts as they passed through the alimentary canal to the digestive gland in the dorsal appendages, the cerata, to the cnidosacs, and finally in the faeces. In digestive cells, many structurally intact nematocysts were present in large phagosomes that remained unaffected even after 2 days fasting, and phagosomes containing nematocysts were found in the faeces. Thus, it is inferred that fusion of nematocyst-containing phagosomes with lysosomes and subsequent digestion of nematocysts is blocked. Masses of exposed, undigested and structurally intact nematocysts were discarded in the faeces. In the tips of the cerata, other nematocysts were phagocytosed by cnidophages and stored in the cnidosacs. After release, in contact with seawater, cnidosac nematocysts were able to discharge. When cnidophore tentacles of E. racemosum with only holotrichous isorhizas were fed to the snails, the isorhizas arrived in the cnidosacs about 2 h after feeding and mixed with existing small euryteles. Some cnidosacs also contained very large microbasic eurytele or large macrobasic eurytele nematocysts, possibly from Eudendrium ramosum and Eudendrium glomeratum, respectively. This indicates that the various types of nematocysts from food were all incorporated into the cnidosacs. Evidence for a selection process or digestion of a certain nematocyst type in the cnidophages was not obtained. It is concluded that a large proportion of the nematocysts ingested with the food are not digested, but are eliminated, structurally and functionally intact, via the alimentary canal and the tips of the cerata.     

Three party symbiosis: acoelomorph worms,corals and unicellular algal symbionts in Eilat (Red Sea)

Epizoic worms were found to occur on certain coral colonies from reefs off the coast of Eilat (Red Sea). We identified 14 coral species infested by acoelomorph worms at a depth range of 2–50 m. The host corals were all zooxanthellate and included both massive and branching stony corals and a soft coral. Worms from all hosts were identified as belonging to the genus Waminoa and contained two distinct algal symbionts differing in size. The smaller one was identified as Symbiodinium sp. and the larger one is presumed to belong to the genus Amphidinium. Worm-infested colonies of the soft coral, Stereonephthya cundabiluensis, lacked a mucus layer and exhibited distinct cell microvilli, a phenotype not present in colonies lacking Waminoa sp. In most cases, both cnidarian and Acoelomorph hosts displayed high specificity for genetically distinctive Symbiodinium spp. These observations show that the epizoic worms do not acquire their symbionts from the “host” coral.     

Temporal patterns of larval settlement and survivorship of two broadcast-spawning acroporid corals

Acroporid corals are the main reef-building corals that provide three-dimensional habitats for other reef organisms, but are decreasing on many reefs worldwide due to natural and anthropogenic disturbances. In this study, temporal patterns of larval settlement and survivorship of two broadcast-spawning acroporid coral species, Acropora muricata and A. valida, were examined through laboratory rearing experiments to better understand the potential for larval dispersal of this important coral group. Many larvae were attached (but not metamorphosed) to settlement tiles on the first examination 3–4 days after spawning (AS). The first permanent larval settlement (i.e. metamorphosed and permanently settled juvenile polyps) occurred at 5–6 days AS, and most larval settlement (85–97% of total) occurred within 9–10 days AS. Larval survivorship decreased substantially to around 50% by the first week of the experiment and to approximately 10% by the second to third week. The rates of larval attachment, settlement, and the initial drop in survivorship of larvae suggest that effective dispersal of some acroporid species may largely be completed within the first few weeks AS.     

Tetracycline reduces sedimentation damage to corals

Sediment deposition on coral reefs occurs naturally and is also caused by man-made disturbances such as dredging; it can result in the death of scleractinian corals by an unknown mechanism. Sedimentation experiments with corals were carried out in El Nido, Northern Palawan, Philippines, in 1986, and in Honolulu, Oahu, Hawaii in 1988. Four species of Indo-Pacific reef corals (Oxypora glabra, Montipora verrucosa, Porites lobata, Pocillopora meandrina) were subjected to sedimentation tests with and without the antibiotic tetracycline to investigate the possible role of microorganisms in the process of sedimentation damage to corals.O. glabra, Porites lobata andPocillopora meandrina were rapidly damaged andO. glabra was always killed by sedimentation.Montipora verrucosa was not injured and may be physiologically resistant to sedimentation damage. Tetracyclinetreated seawater reduced the rate of tissue necrosis and prevented colony mortality, suggesting that tetracycline-sensitive bacteria are involved in the process of tissue necrosis and may be partially responsible for coral mortality following sediment deposition.     

Ecological and biological phenomena influencing coral-species composition on the reef tables at Eilat (Gulf of Aqaba,Red Sea)

Observations performed on coral colonies damaged during a very low tide on the shallow-water reef tables at Eilat (Red Sea), showed that most colonies are able to regenerate if parts of the living tissue remain intact on the skeletons. Brain-like corals, such asFavia favus andPlatygyra lamellina, were found to be more resistant to dessication than the delicate bush-like forms. It was also observed that the exposed skeletons of corals were immediately covered by a successive group of algae with a rich invertebrate epifauna among them. It is postulated that unpredicted extreme low water, occurring once in several years, could be the factor that prevents crowding on the reef tables, thus keeping high the level of coralspecies diversity.Supported, in part, by a grant from the Smithsonian Institution, USA. SFC-70074.This paper was presented at the Intecol Symposium held in Dubrovnik in March, 1972.     

Light harvesting in the green alga Ostreobium sp., a coral symbiont adapted to extreme shade

Ostreobium sp. (Chlorophyta: Siphonales) can be found as green bands within the skeletal material of a number of stony corals in the Indo-Pacific and Caribbean regions. Many of these corals also contain symbiotic dinoflagellates in the overlaying coral polyps that effectively screen out all the typical photosynthetically active radiation from the algae in the green bands below. Ostreobium sp., nevertheless, grows photosynthetically. Its action spectrum and absorption spectrum have been shown to extend much further into the near infra-red compared to other green algae. In the present study, carried out in 1987, fluorescence excitation and emission spectra were measured in Ostreobium sp. and compared to spectra obtained from the green alga Ulva sp. and the brown alga Endarachne sp. Xanthophylls, probably siphonein and an unidentified xanthophyll probably related to siphonaxanthin, are photosynthetically active in Ostreobium sp., and can sensitize Photosystem II fluorescence at 688 nm and Photosystem I (PS I) fluorescence at 718 nm. The fluorescence emission spectra of Ostreobium sp. measured at 25° C and 77 K were not remarkably different from those of the green alga Ulva sp. Absorbance changes induced by light were measured in Ostreobium sp. from 670 to 750 nm and were like those normally seen in green plants except that, in addition to the minimum expected for the reaction-center chlorophyll of PS I (P700) at 703 nm, another minimum was seen at 730 nm. It is possible that this spectrumreflects the functioning of a reaction center of Photosystem I that has adapted to function in light highly enriched in far-red wavelengths.CIW-DPB Publication No. 1021     

Assessing the antipredatory defensive strategies of Caribbean non-scleractinian zoantharians (Cnidaria): is the sting the only thing?

The relative importance of chemical, nematocyst, and nutritional defenses was examined for 18 species of Caribbean sea anemones (actinarians), zoanthids, and mushroom polyps (corallimorpharians) from the Florida Keys and the Bahamas Islands, 2008–2010. Feeding assays were performed using the fish Thalassoma bifasciatum with artificial foods containing crude organic extracts of cnidarian tissues. A novel behavioral assay using brine shrimp nauplii was used to assess nematocyst defenses. The nutritional quality of cnidarian tissues was examined using bomb calorimetry and soluble protein assays. In general, actinarians invested in nematocyst defenses, zoanthids in either nematocyst or chemical defenses, and corallimorpharians lacked both, except for 1 of 3 species that was chemically defended. Relative to other coral reef invertebrates, cnidarian tissues had similar caloric values but lower soluble protein concentrations. Trade-offs between chemical and nematocyst defenses were observed for 65% of species, while habitat and behavior provided a likely explanation for undefended species.     

Ingestion of a medusa (Aegina citrea) by the nematocyst-containing ctenophore Haeckelia rubra (formerly Euchlora rubra): phylogenetic implications

The rare ctenophore Haeckelia rubra (formerly Euchlora rubra) has long been known to have nematocysts rather than colloblasts in its tentacles. Five specimens were collected in the San Juan Archipelago, Washington State, USA in 1980 and 1981, and their feeding behavior was observed in the laboratory. We found that H. rubra readily eats the tentacles of a medusa, Aegina citrea, whose nematocysts (apotrichous isorhizas) are nearly identical in morphology to the nematocysts of the ctenophore. When H. rubra was offered 16 other species of hydromedusae and 1 siphonophore in the laboratory, the ctenophores showed little or no tendency to ingest these potential prey items. In addition to its routinely positive response to A. citrea, the ctenophore could be induced by manipulation and starvation to accept and ingest bits of the bodies of 4 additional species of hydromedusae and 1 siphonophore. These results, combined with the histological and rearing experiments of other investigators, leave little doubt that the nematocysts in H. rubra are not endogenous, but are kleptocnidae similar to those nematocysts retained and subsequently used by some species of nudibranchs that feed on Cnidaria. A close phylogenetic link between the Cnidaria and the Ctenophora is most unlikely.