Defensive strategies of soft corals (Coelenterata: Octocorallia) of the Great Barrier Reef

Summary The relationship between ichthyotoxicity and predation-related defensive functional morphology was examined in alcyonacean soft corals of the central and northern regions of the Great Barrier Reef (GBR), Australia. Approximately 170 specimens were assessed encompassing a number of genera within three families: 1) the Alcyoniidae (Lobophytum, Sarcophytum, Sinularia, Cladiella, Parerythropodium, and Alcyonium); 2) Neptheidae (Lemnalia, Paralemnalia, Capnella, Lithophyton, Nephthea, Dendronephthya, Scleronephthya, and Stereonephthya), and 3) Xeniidae (Anthelia, Efflatounaria, Cespitularia, Heteroxenia, and Xenia). Ichthyotoxicity data were derived from earlier studies which used Gambusia affinis Baird and Girard (Vertebrata, Pisces) as a test organism. These data were compared to morphological data collected from specimens in the field and laboratory. Three sets of statistical analyses were performed, each considering a progressively narrower group of taxa. The first included 68 specimens and considered 16 morphological characters in each, falling into the general categories of gross colony form, colony texture, presence of mucus, colony color, polyp retractility, and sclerite morphology and distribution. These were tested for independence against ichthyotoxicity data. The second set of analyses involved a more restricted morphological data set derived from 28 species of Sinularia in combination with 28 species within the Nephtheidae, comparing them to their respective toxicity ranks. The third analysis considered the previous two taxonomic groups separately in relation to their toxicity levels.The attempt to consider many morphological characters in a taxonomically diverse collection did not reveal any general association in the Alcyonacea between defensive morphology and toxicity, and those associations which did emerge were clearly erroneous. The second analysis, considering only Sinularia spp. and nephtheids, demonstrated a negative association between ichthyotoxicity and the morphological characters of a) polypary armament, b) microarmament of the individual polyp, and c) strong mineralization of the coenenchyme. The third analysis revealed that the negative association found between toxicity and the first two characters was derived entirely from the nephtheids while the association detected between toxicity and the third character was restricted to Sinularia. It is concluded that a relationship between toxicity and morphology can be demonstrated, but it is heavily dependent upon which specific morphological characters are being considered and at what level of taxonomic resolution the analysis is being performed. An approach utilizing many characters over many taxa is unlikely to yield significant, reliable, or meaningful results.Australian Institute of Marine Science Contribution Number 383     

Competitive strategies of soft corals (Coelenterata: Octocorallia). II. Variable defensive responses and susceptibility to scleractinian corals

Interactions involving competition for space between several species of alcyonacean and scleractinian corals were assessed experimentally on Britomart Reef, central region of the Great Barrier Reef, Australia. Colonies of three soft coral species, Sarcophyton ehrenbergi Marenzeller, Nephthea brassica Kukenthal, and Capnella lacertiliensis Macfayden Forskal (Coelenterata:Alcyonacea) were relocated within stands of two scleractinian corals, Parités andrewsi Vaughan (= P. cylindrica Dana) and Pavona cactus Förskal (Coelenterata:Scleractinia). Undisturbed scleractinian and relocated alcyonacean controls were also monitored.Alcyonacean corals induced necrosis of tissue in scleractinian corals. Necrosis was significantly more pronounced when colonies were in contact but was also observed in the absence of contact, implicating the presence of active allelopathic agents. Scleractinian coral species varied in their susceptibility to the ill effects of alcyonaceans, with Pontes andrewsi being more susceptible than Pavona cactus. Of the soft corals, Nephthea caused the highest degree of mortality in the two scleractinian corals examined and Sarcophyton the least. Some soft corals appear to retain their toxins while others release them, implying a combination of anti-predatory and anti-competitor roles for the secondary metabolites. Scleractinian corals were often overgrown by soft corals.Both species of scleractinian corals were found to cause approximately equal amounts of tissue necrosis in alcyonaceans. These effects were more pronounced when colonies were in direct contact. The necrotic effects among alcyonacean corals were species-specific. Alcyonaceans also overgrew scleractinian corals and secreted a protective polysaccharide layer in areas proximal to scleractinians. Secretion of this layer was stimulated differentially by the two scleractinian species and also varied in frequency of occurrence among the alcyonaceans.High levels of tissue necrosis were observed in both groups of organisms within 3 wk of initiation of the experiment. Necrosis increased with time in the scleractinian corals and decreased in the alcyonaceans. The development of a protective polysaccharide layer in the alcyonaceans increased with time.     

Feeding behaviour and feeding ecology of the Octocorallia (Coelenterata: Anthozoa)

The feeding behaviour of some 30 species of Octocorallia was examined in the laboratory and in the field. All of the species from the Orders–Alcyonacea, Gorgonacea, Stolonifera and Telestacea, appear to have a common, basic feeding strategy. Fine particulate matter and zooplankton were captured in a raptorial manner by the tentacles and pinnules. Upon capture of food particles, the tentacles were flexed rapidly inwards and closed or wiped across the mouth. At the same time the mouth opened and ingestion was accomplished by directional ciliary currents in the mouth and the pharynx. Food capture by means of mesenterial filaments, mucus strands or ciliary currents was not observed. The examination of gut contents of Alcyonacea showed that they feed upon zooplankton. Epibenthic copepods from the demersal coral reef zooplankton were an important element of the diet of alcyonaceans.     

A new family of soft corals (Anthozoa,Octocorallia, Alcyonacea) from the aphotic tropical eastern Pacific waters revealed by integrative taxonomy

The new family Aquaumbridae is described based on distinct morphological characters and supported by a molecular phylogenetic analysis. Aquaumbridae is similar to Nidaliidae, Nepththeidae and Alcyoniidae in having arborescent colonies and lacking axis structure or stolons, but differs from them in having very different sclerite composition and having conspicuous transparent jelly-like lobes. Phylogenetic analysis of two mitochondrial genes, ND2 and mtMutS, strongly supports its placement in a separated clade. Herein we describe Aquaumbra klapferi sp. nov., gen. nov. in the new family. The organisms were obtained from the seamounts, ridges and canyons out of the insular shelf of Isla del Coco, Costa Rica, down to 400 m depth. The new species represents the first discovery of a soft coral in an eastern Pacific oceanic island, and provides hints of the biodiversity of the largely unexplored deep waters of the tropical eastern Pacific.     

Chemical studies of marine invertebrates. XXIII. A novel polyhydroxylated sterol from the soft coral Litophyton viridis (Coelenterata, Octocorallia, Alcyonacea).

The structure of 24-methylenecholest-5-en-3beta,7beta,19-triol (II) isolated from the soft coral Litophyton viridis, has been established by X-ray diffraction analysis. The compound is accompanied by its 7-monoacetate derivative.     

A new genus of soft coral (Cnidaria,Octocorallia) from the Republic of Congo (Pointe-Noire Region)

A new genus of soft coral from the Republic of Congo is described, Complexum gen. n. Nine West African octocoral species previously described in the genus Alcyonium by Tixier-Durivault (1955) are referred to this new genus, and a new species is described and figured, Complexum pusillum sp. n. The new species is characterized by having encrusting growth form and abundant spiny clubs in the surface of the polyparium. It colonizes shallow calcareous rocky banks (5 to 20 m depth) existing in coastal water of the region of Pointe-Noire. Based on molecular phylogeny this new genus is well separated from Alcyonium species.     

In situ sampling of interstitial water from lake sediments

A sampler with a relatively high resolution has been developed, which allows interstitial water to be obtained from lake sediments at well defined depths, without serious disturbance of sediment structure. Oxidation effects are excluded. Sampling time is in the order of a day. Installation requires little additional equipment. The instrument has been developed for use in shallow lakes.     

Morphological and genetic analyses of xeniid soft coral diversity (Octocorallia; Alcyonacea)

Studies on the biodiversity and evolution of octocorals are hindered by the incomplete knowledge of their taxonomy, which is due to few reliable morphological characters. Therefore, assessment of true species diversity within abundant and ecologically important families such as Xeniidae is difficult. Mitochondrial genes provide a reliable solution to this problem for a wide range of taxa. However, low mutation rates of the mitochondrial DNA in octocorals result in insufficient variability for species discrimination. We compared the variation of a fragment of the Signal Recognition Particle 54 gene (SRP54, proposed for octocorals) and the mitochondrial ND6/ND3 marker among members of the xeniid genera Ovabunda, Xenia, Heteroxenia and Bayerxenia. The mean uncorrected pairwise sequence divergence was 39 % for SRP54 compared to 2 % for ND6/ND3. Morphological assignments were not always supported by genetics: Species diversity was underestimated (one case) or overestimated, probably reflecting intraspecific polymorphisms or hinting at recent speciations. ND6/ND3 is informative for some species-level assignments, whereas SRP54 shows the variability needed for species delimitations within this understudied taxon. Our results on both genes show their potential for evolutionary and biodiversity studies in Xeniidae.     

Colony induction and growth inhibition in Desmodesmus quadrispina (Chlorococcales) by allelochemicals released from the filamentous alga Uronema confervicolum (Ulotrichales)

In biofilms, the competition between microorganisms for light, nutrients and space is extreme. Moreover, planktonic algae can be considered as competitors insofar as they decrease the available light for the benthic algae. One of the strategies employed by microorganisms to eliminate competitors is the release of inhibiting compounds, a process known as allelopathy. Here we demonstrate that a benthic/epiphytic alga, Uronema confervicolum, produces allelopathic compounds that induce oxidative stress and growth inhibition in the planktonic Desmodesmus quadrispina. Some of these compounds can also trigger the formation of colony in D. quadrispina. As colonies have higher sedimentation rates than unicells, their induction by U. confervicolum might decrease shading. This study is the first report of colony induction in the context of alga-alga interaction. Our results also suggest the implication of mitogen-activated protein (MAP) kinases in the transduction of the signal leading to the formation of reactive oxygen species in the cells. A comparison with allelochemicals from another planktonic green alga, Monoraphidium aff. dybowski, emphasizes the specificity of colony induction by U. confervicolum, in contrast with oxidative stress which is induced by several compounds. The reciprocal production of inhibiting compounds by D. quadrispina makes this interaction an interesting example of co-evolution between two microorganisms belonging to different compartments of the ecosystem.     

A new deep-sea pennatulacean (Anthozoa: Octocorallia: Chunellidae) from the Porcupine Abyssal Plain (NE Atlantic)

During the BENGAL cruises, an important collection of deep-sea benthic organisms was sampled. Among the pennatulacean colonies, a previously undescribed species of chunellid was collected. That material is here described as the type species of a new genus, Porcupinella gen. nov. The new genus and species are described based on material collected in the Porcupine Abyssal Plain (NE Atlantic), 4,839–4,847 m in depth. This is the first time that a chunellid is reported from the Atlantic Ocean. The new genus is compared with the other genera in the family, and some phylogenetic remarks about the families Chunellidae and Umbellulidae are also provided.     

Two new dimorphic soft-coral species (Anthozoa: Octocorallia) from Antarctica

Sphaeralcyon shackletonisp. nov. and Sphaeralcyon scottisp. nov. are described and illustrated from material collected at the Scotia Sea and Weddell Sea on the Polarstern cruises ANT XIX/3 (ANDEEP-I), ANT XIX/5 (LAMPOS), and ANT XXI/2 (BENDEX). With the discovery of Sphaeralcyon shackletoni and S. scotti, three species are now known in the genus Sphaeralcyon, all them reported from the Southern Ocean. The diagnosis of the genus has been slightly modified to accommodate some of the characters of the new species.     

Exploring the utility of an indel-rich, mitochondrial intergenic region as a molecular barcode for bamboo corals (Octocorallia: Isididae)

The DNA barcoding initiative has advocated the use of the 5'-end (～658bp) of mitochondrial (mt) cytochrome c oxidase subunit 1 (cox1) to genetically distinguish species. However, this has proven difficult within the subclass Octocorallia due to extraordinarily low substitution rates within mt protein-coding genes. Intergenic regions (IGRs), which have been little examined among octocorals, may be subject to high mutation rates and have proven useful target regions at both the interspecific and population levels of metazoans. Herein we examine a mt IGR (igr4) between the cytochrome b (cob) and NADH dehydrogenase subunit 6 (nad6) genes among species of the bamboo coral subfamily Keratoisidinae to evaluate its utility for barcoding and phylogenetic studies. Among 77 keratoisidin specimens, we found igr4 to vary in length between either 42bp (Acanella Gray, 1870 and Orstomisis Bayer, 1990) or 302-605bp (Isidella Gray, 1857, Lepidisis Verrill, 1883, Keratoisis Wright, 1869, and two undescribed genera). We interpreted the short igr4 sequence of Acanella eburnea (Pourtalès, 1868) as potentially indicative of additional mt genome-related novelties and thus sequenced its entire mt genome; gene content and gene order were the same as in a previously-sequenced bamboo coral mt genome. Alignment of the longer igr4 sequences included 108 parsimony-informative characters, as well as numerous indels ranging from 2-262bp in length. Uncorrected pairwise 'p' distances indicated sequence variation of 0-27.2%, as compared to 0-4.8% among the same specimens for the MutS homolog (msh1), currently the most widely sequenced octocorallian mt gene, and <0.4% for cox1 for a subset of the taxa. Despite the greater levels of variation, fewer unique haplotypes were observed at igr4 compared to msh1; however, in combination, the two gene regions revealed increased mt haplotype diversity relative to either gene region on their own.     

Two new gorgonian genera (Octocorallia: Primnoidae) from Southern Ocean waters

Tauroprimnoa austasensis gen. nov., sp. nov. and Digitogorgia kuekenthali gen. nov., sp. nov. are described and illustrated from Southern Ocean waters. The most distinctive characters of the newly proposed genera are, in Tauroprimnoa, the existence of four marginal scales, two abaxials with a strong thorn, and the presence of a single abaxial longitudinal row of body scales. In the case of Digitogorgia, the colony branching pattern, the structure of the opercular scales, and the presence of a complete cycle of accessory opercular scales are the distinct features to distinguish it from previously known genera. Tauroprimnoa austasensis sp. nov. is reported from the Eastern Weddell Sea, Antarctica, while Digitogorgia kuekenthali sp. nov. has been found in the SubAntarctic waters off Burdwood Bank and, in the south east of Isla Nueva in Chilean Patagonia.     

Electrophysiology of a myoid epithelium inChelophyes (Coelenterata: Siphonophora)

Summary The passive electrical properties, and the ionic basis of the action potential have been examined in the subumbrella myoid epithelium of the siphonophoreChelophyes. The myoepithelial cells are electrically coupled, and are 20 m wide, some 1 mm long, and only 5 m thick. Membrane constants determined by a 2-electrode study were: = 280 m; R_m = 0.11 kOhm/cm²; R_i = 24 Ohm/cm. Mean resting potential was – 85 mV. The first action potential of a series (whether evoked by repetitive stimulation, or occurring in a natural unstimulated swimming burst) shows a rapid rise and fall with no afterpotential. The overshoot is small, but successive action potentials show a remarkable facilitation, overshooting by as much as 70 mV. They also show a plateau phase after the initial rapid rise, which is terminated by a rapid fall. Conduction velocity was 27 cm/s.Changes in the external milieu, and the effects of Ca²⁺ blocking agents indicated that the action potentials are complex events. Although insensitive to TTX, the action potential is dependent on external sodium concentration, and is not abolished by Ca²⁺ blocking agents: in this respect it resembles the sodium-dependent action potentials of other siphonophore tissues.The ionic basis of the facilitated action potentials is not yet clear, but it seems probable that a fast potassium conductance terminating the unfacilitated action potential is progressively inactivated during repetitive activity, and that the plateau phase of the facilitated action potential is maintained by a sodium conductance mechanism, to be terminated by a calcium-activated potassium conductance.Abbreviations EGTA 1,2-bis-[2-di(carboxymethyl)-amino-ethoxyl]-ethane - TEA tetraethyl ammonium chloride - TTX tetrodotoxin This work was carried out during a visit to the Station Zoologique, Villefranche-sur-Mer in the spring of 1979; it is a pleasure to express our thanks to Prof. P. Bougis and his staff for their kind hospitality. Q.B. and P.A.V.A. were supported by a grant from the British Council which is gratefully acknowledged.     

Chemical studies of marine inverterbrates. XXIX 4 alpha-Methyl-3 beta 8-beta-dihydroxy-5 alpha-ergost-24(28)-en-23-one, a novel polyoxygenated sterol from the soft coral Litophyton viridis. (Coelenterata, Octocorallia, Alcyonacea).

A novel sterol, 4α-methyl-3β,8β-dihydroxy-5α-ergost-24(28)-en-23-one (I), has been isolated from the soft coral Litophyton viridis. Its structure and relative configuration has been established by X-Ray diffraction analysis.     

A second, cryptic species of the soft coral genus Incrustatus (Anthozoa: Octocorallia: Clavulariidae) from Tierra del Fuego, Argentina, revealed by DNA barcoding

The encrusting soft coral Incrustatus comauensis is a common denizen of hard substrates in the shallow sub-tidal zone from the central Chilean fjords to the Cape Horn region of southern South America. DNA barcoding of specimens collected from the Beagle Channel, Tierra del Fuego, Argentina, revealed the presence of a second, cryptic species of Incrustatus that is syntopic with I. comauensis. We describe Incrustatus niarchosi, a new species that can be distinguished morphologically from I. comauensis by differences in the microscopic ornamentation of the coenenchymal sclerites. To date, I. niarchosi n. sp. is known only from the Beagle Channel. A population of I. comauensis discovered in the intertidal zone in eastern Tierra del Fuego represents a new record of the species for that habitat and geographic region. Although the intertidal population is also distinct genetically, it is morphologically indistinguishable from sub-tidal Chilean populations of I. comauensis, and at present, there is insufficient evidence to support its status as a separate species.     

A new gorgonian genus from deep-sea Antarctic waters (Octocorallia, Alcyonacea, Plexauridae)

Mesogligorgia scotiae gen. nov., sp. nov. is described and illustrated from a colony collected in the Scotia Sea, 2,201–2,213 m in depth, on the ANDEEP-I cruise. The new taxon is placed in the family Plexauridae because of: 1) the presence of a horny axis with a cross-chambered central core and numerous loculi, 2) retractile polyps in calyces with distinct spicular components, and 3) armed polyps with large sclerites with a poorly- developed collaret and eight well-developed points. The irregularly distributed sclerites running along the axis, into a thick mesogloeal coenenchyme, and the elongated spindles with irregular ends are the most distinctive characters of the newly proposed genus.