Bleaching response of corals and their <Emphasis Type="Italic">Symbiodinium</Emphasis> communities in southern Africa

The high-latitude coral communities of southern Africa suffered minimal impacts during past mass bleaching events. Recent reports indicate an increase in bleaching frequency during the last decade, yet the actual levels of thermal stress and contributing factors in these bleaching events, and the degree of acclimatisation or adaptation on these reefs are poorly understood. During the 2005 warm-water anomaly in the southern Indian Ocean we conducted bleaching surveys and collected samples for genotyping of the algal symbiont communities at 21 sites in southern Mozambique and South Africa. Coral bleaching reached unprecedented levels and was negatively correlated with both latitude and water depths. Stylophora pistillata and Montipora were the most susceptible taxa, whereas three common branching corals had significantly different bleaching responses (Stylophora > Acropora > Pocillopora). Temperature records indicated that localised strong upwelling events coupled with persistent above-average seawater temperatures may result in accumulated thermal stress leading to bleaching. Symbiodinium in 139 scleractinian corals belonged almost exclusively to clade C, with clade D symbionts present in only 3% of the colonies. Two atypical C subclades were present in Stylophora and Pocillopora colonies and these were more abundant in shallow than deeper sites. Taxon-specific differences in bleaching responses were unrelated to different clades of algal symbionts and suggest that Symbiodinium C subtypes with diverse thermal tolerance, coupled with acclimatisation and morphology of the host colony influence the bleaching response. Additionally, the predominance of putatively thermal-sensitive Symbiodinium in southern African corals may reflect a limited experience of bleaching and emphasises the vulnerability of these reefs to moderate levels of thermal stress.     

Variation in colony geometry modulates internal light levels in branching corals, <Emphasis Type="Italic">Acropora humilis</Emphasis> and <Emphasis Type="Italic">Stylophora pistillata</Emphasis>

Colonial photosynthetic marine organisms often exhibit morphological phenotypic plasticity. Where such plasticity leads to an improved balance between rates of photosynthesis and maintenance costs, it is likely to have adaptive significance. To explore whether such phenotypic plasticity leads to more favourable within-colony irradiance for reef-building branching corals, this relationship was investigated for two coral species Acropora humilis and Stylophora pistillata, along a depth gradient representing light habitats ranging from 500 to 25 μmol photons m⁻² s⁻¹, during 2006 at Heron Island, Great Barrier Reef (23.44°S, 151.91°E). In the present study changes in flow-modulated mass transfer co-varied with light as a function of depth. In low-light (deep) habitats, branch spacing (colony openness) in A. humilis and S. pistillata was 40–50% greater than for conspecifics in high-light environments. Also, branches of A. humilis in deep water were 40–60% shorter than in shallow water. Phenotypic changes in these two variables lead to steeper within-colony light attenuation resulting in 38% higher mean internal irradiance (at the tissue surface) in deep colonies compared to shallow colonies. The pattern of branch spacing was similar for S. pistillata, but this species displayed an alternate strategy with respect to branch length: shade adapted deep and cave colonies developed longer and thinner branches, allowing access to higher mass transfer and irradiance. Corals in cave habitats allowed 20% more irradiance compared to colonies found in the deep, and had a 47% greater proportion of irradiance compared to colonies in the shallow high-light environment. Such phenotypic regulation of internal light levels on branch surfaces partly explains the broad light niches of many branching coral species. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Biochemical composition,energy content and chemical antifeedant and antifoulant defenses of the colonial Antarctic ascidian<Emphasis Type="Italic"> Distaplia cylindrica</Emphasis>

The colonial ascidian Distaplia cylindrica occurs both as scattered individual colonies or in gardens of colonies in fine-grained soft substrata below 20 m depths off Anvers Island along the Antarctic Peninsula. Individual colonies, shaped as tall rod-like cylinders and anchored in the sediments by a bulbous base, may measure up to 7 m in height. D. cylindrica represent a considerable source of materials and energy for prospective predators, as well as potential surface area for fouling organisms. Nonetheless, qualitative in situ observations provided no evidence of predation by sympatric predators such as abundant sea stars, nor obvious biofouling of colony surfaces. Mean energy content of whole-colony tissue of D. cylindrica was relatively high for an ascidian (14.7 kJ g^–1 dry wt), with most of this energy attributable to protein (12.7 kJ g^–1 dry wt). The sympatric omnivorous sea star Odontaster validus consistently rejected pieces of D. cylindrica colonies in laboratory feeding assays, while readily ingesting similarly sized alginate food pellets. Feeding deterrence was determined to be attributable to defensive chemistry, as colonies of D. cylindrica are nutritionally attractive and lack physical protection (conspicuous skeletal elements or a tough outer tunic), and O. validus display significant feeding-deterrent responses to alginate food pellets containing tissue-level concentrations of organic extracts. In addition, high acidity measured on outer colony surfaces (pH 1.5) as well as homogenized whole-colony tissues (pH 2.5) are indicative of surface sequestration of inorganic acids. Agar food pellets prepared at tissue levels of acidity resulted in significant feeding deterrence in sea stars. Thus, both inorganic acids and secondary metabolites contribute to chemical feeding defenses. D. cylindrica also possesses potent antifoulant secondary metabolites. Tissue-level concentrations of hydrophilic and lipophilic extracts caused significant mortality in a sympatric pennate diatom. Chemical feeding deterrents and antifoulants are likely to contribute to the abundance of D. cylindrica and, in turn, play a role in regulating energy transfer and community structure in benthic marine environments surrounding Antarctica.Communicated by P.W. Sammarco, Chauvin     

Cytoenzymatic investigation of intracellular digestion in the symbiont-bearing hydrothermal bivalve <Emphasis Type="Italic">Bathymodiolus azoricus</Emphasis>

Invertebrates harbouring endosymbiotic chemoautotrophic bacteria are widely distributed in a variety of reducing marine habitats, including deep-sea hydrothermal vents. Bathymodiolids are dominants of the biomass at geochemically distinct vent sites of the Mid Atlantic Ridge (MAR) and thus are good candidates to study biological processes in response to site-specific conditions. To satisfy their nutritional requirements, these organisms depend to varying extent on two types of chemoautotrophic symbionts and on filterfeeding. The quantitative relationships of the nutritional modes are poorly understood. Using enzyme cytochemistry, electron microscopy and X-ray microanalysis, the structural and functional aspects of the cellular equipment necessary for lysosomal digestion was studied. We provide evidence for the following: (1) the basis of intracellular digestion of symbionts in Bathymodiolus azoricus from two geochemically distinct vent sites was not mainly in the large lysosomal bodies as previously thought (based on the membranous content resembling bacteria); (2) senescent bacteria are autolysed, possibly by bacterial acid phosphatase, that is more likely a cell cycling of the symbionts rather than an active lysosomal digestion by the host; (3) the consistent absence of hydrolases may indicate the improper use of the name “lysosome” for large vesicles at the base of the gill bacteriocytes (4) nutrient transfer in B. azoricus, therefore, may more likely be accomplished through leaking of metabolites from the symbiont to the host, not excluding lysosomal resorption of dead bacteria as an auxiliary strategy for organic molecule transfer; (5) evidence is provided for microvillar transfer of substances from the seawater that may indicate filter-feeding, in non-symbiotic ciliated gill cells of mussels from Lucky Strike; (6) two types of lysosomal vesicles can be distinguished in digestive cells based on their enzymatic content and their elemental composition.     

The origin of the chemical profiles of fungal symbionts and their significance for nestmate recognition in <Emphasis Type="Italic">Acromyrmex</Emphasis> leaf-cutting ants

Cuticular hydrocarbon profiles are essential for nestmate recognition in insect societies, and quantitative variation in these recognition cues is both environmentally and genetically determined. Environmental cues are normally derived from food or nest material, but an exceptional situation may exist in the fungus-growing ants where the symbiotic fungus garden may be an independent source of recognition compounds. To investigate this hypothesis, we quantified the chemical profiles of the fungal symbionts of 18 sympatric colonies of Acromyrmex echinatior and Acromyrmex octospinosus and evaluated the quantitative variation of the 47 compounds in a multivariate analysis. Colony-specific chemical profiles of fungal symbionts were highly distinct and significantly different between the two ant species. We also estimated the relative genetic distances between the fungal symbionts using amplified fragment length polymorphism (AFLP) and correlated these with the overall (Mahalanobis) chemical distances between the colony-specific profiles. Despite the standardized laboratory conditions, the correlations were generally weak, but a statistically significant portion of the total variation in chemical profiles could be explained by genetic differences between the fungal symbionts. However, there was no significant effect of ant species in partial analyses because genetic differences between symbionts tend to coincide with being reared by different ant species. However, compound groups differed significantly with amides, aldehydes, and methyl esters contributing to the correlations, but acetates, alkanes, and formates being unrelated to genetic variation among symbionts. We show experimentally that workers that are previously exposed to and fed with the fungal symbiont of another colony are met with less aggression when they are later introduced into that colony. It appears, therefore, that fungus gardens are an independent and significant source of chemical compounds, potentially contributing a richer and more abundant blend of recognition cues to the colony “gestalt” than the innate chemical profile of the ants alone. Freddie-Jeanne Richard and Michael Poulsen contributed equally to this work.     

Microbial associations in sponges. II. Numerical analysis of sponge and water bacterial populations

Three taxonomically distant sponges Pericharax heteroraphis, Jaspis stellifera and Neofibularia irata contain phenotypically similar bacterial symbionts which differ from bacteria in the ambient water. These symbionts are predominant in the sponges and were detected after computer analysis of 526 heterotrophic bacterial strains tested for 76 characters. These facultative anaerobic symbionts metabolize a wide range of compounds and may be important in removing waste products while the sponges are not circulating water. The bacteria produce sticky-mucoid colonies and thus would contribute to sponge structural rigidity. The fourth sponge Ircinia wistarii contains a mixed aerobic population similar to that in the ambient water. The majority of the bacteria are located around the inhalant canals, facilitating the uptake of dissolved organic matter and oxygen from the incoming water.     

Bacterial sulfate reduction within reduced microniches of oxidized marine sediments

Bacterial sulfate reduction was demonstrated in the oxidized surface layers of a coastal marine sediment using a radiotracer technique. The obligate anaerobic process takes place within reduced sediment pellets of 50 to 200 m diameter. The H₂S produced diffuses out into the interstitial solution and is oxidized before any detectable accumulation takes place. This microniche structure explains the presence of sulfate-reducing (Desulfovibrio spp.) and sulfide oxidizing (Beggiatoa spp.) bacteria and of ferrous sulfide and pyrite in the oxidized sediment. Sulfate reduction was also demonstrated within detrital particles experimentally decomposed in oxic seawater or sediment. The limiting conditions for the maintenance of a reduced microniche within an oxic environment is discussed in terms of a theoretical model.     

Symbiont specificity and bleaching susceptibility among soft corals in the 1998 Great Barrier Reef mass coral bleaching event

Considerable variability in bleaching was observed within and among soft coral taxa in the order Alcyonacea (Octocorallia: Cnidaria) on the central Great Barrier Reef (GBR, latitude 18.2°–19.0°S, longitude 146.4°–147.3°E) during the 1998 mass coral bleaching event. In April 1998, during a period of high sea surface temperatures, tissue samples were taken from bleached and unbleached colonies representative of 17 soft coral genera. The genetic identities of intracellular dinoflagellates (Symbiodinium spp.) in these samples were analyzed using PCR-denaturing gradient gel electrophoresis fingerprinting analysis of the internal transcribed spacer regions 1 and 2. Alcyonaceans from the GBR exhibited a high level of symbiont specificity for Symbiodinium types mostly in clade C. A rare clade D type (D3) was associated only with Clavularia koellikeri, while Nephthea sp. hosted symbionts in clade B (B1n and B36). Homogenous Symbiodinium clade populations were detected in all but one colony. Colonies that appeared bleached possessed symbiont types that were genetically indistinguishable from those in nonbleached conspecifics. These data suggest that parameters other than the resident endosymbionts such as host identity and colony acclimatization are important in determining bleaching susceptibility among soft corals. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The bacterial symbiont from the hydrothermal vent tubewormRiftia pachyptila is a sulfide specialist

We have investigated the metabolic adaptations of the chemolithotrophic bacterial symbionts ofRiftia pachyptila. Specimens of the tubeworm were collected by submersible from depths of 2600 m at 13°N on the East Pacific Rise in 1987, and 2450 m at the Galápagos Rift in 1988. Isolated bacteria utilize sulfide, but not thiosulfate or sulfite, as their sole reduced-sulfur energy source. The bacteria rapidly oxidize a wide range of sulfide concentrations (5µM to 2 mM), with maximal respiration rates at concentrations >1 mM, and unlike many sulfur-oxidizing bacteria, show no inhibition in oxygen consumption at sulfide concentrations up to 2 mM. Incubations of freshly homogenized trophosome tissue or isolated bacteria with sodium [³⁵S] sulfide and subsequent analysis of sulfur products by high-performance liquid chromatography and flow-through scintillation counting showed that sulfide disappeared almost completely within 1 min. Both soluble and insoluble products of sulfide oxidation were produced. The soluble fraction contained sulfate and polysulfides, with no thiosulfate produced. However, the majority of the radioactivity was in the water-insoluble fraction, mostly as elemental sulfur. Whole-worm experiments under pressure showed a rapid removal of³⁵S-sulfide from the incubation water, with sulfide, sulfate, and polysulfides appearing in the blood within 4 h. There was no utilization of thiosulfate by the whole worms, freshly homogenized trophosome tissue, or isolated bacteria.     

Sexual reproduction of the photosymbiotic ascidian <Emphasis Type="Italic">Diplosoma virens</Emphasis> in the Ryukyu Archipelago,Japan: vertical transmission,seasonal change,and possible impact of parasitic copepods

Diplosoma virens is a colonial ascidian hosting prokaryotic algae Prochloron sp. in the common cloacal cavity of the colonies and is sometimes parasitized by notodelphyid copepods. In ascidian–Prochloron symbiosis, it is generally known that the host larvae acquire the algal symbionts from their mother colonies to maintain the symbiosis. A histological study of the sexually mature colonies of D. virens showed that the algal symbionts attach to pre-hatching larvae on the rastrum (plant rake) projected from the postero-dorsal part of the larval trunk, and then the rastrum is packed in the posterior half of the larval trunk that will become a cloacal cavity after metamorphosis. This process is the same as that of D. simile. Monthly sampling of D. virens colonies showed that they have embryos in summer in Ryukyus, situated near the northern-most limit of the coral reefs in the West Pacific. While the frequencies of copepod parasitism were variable among the populations, the colonies from a highly parasitized population had a significantly smaller number of eggs/embryos per zooid than the colonies from the less parasitized populations. The parasites probably have an inhibitory impact on the sexual reproduction of the host colonies.Communicated by T. Ikeda, Hakodate     

Chemical camouflage by myrmecophilous beetles Zyras comes (Coleoptera: Staphylinidae) and Diaritiger fossulatus (Coleoptera: Pselaphidae) to be integrated into the nest of Lasius fuliginosus (Hymenoptera: Formicidae)

Summary. The myrmecophilous beetles, Zyras comes (Staphylinidae) and Diaritiger fossulatus (Pselaphidae) are guests of the black shining ant Lasius fuliginosus. Host worker ants never attacked these beetles, and often gave regurgitant to Z. comes following tactile communication with the beetle. By contrast, the workers from colonies without the myrmecophiles showed hostile responses towards Z. comes before tactile contact, but were not aware of D. fossulatus until contact. In L. fuliginosus, workers within a colony shared profiles, but the profiles differed among colonies. GC analyses showed that both Z. comes and D. fossulatus beetles had the same hydrocarbons as L. fuliginosus, and the profiles were more similar to those of the host colony workers than the foreign workers. Both Z. comes and D. fossulatus appear to imitate the hydrocarbon profile of their host workers, allowing integrating into the host nest. A Y-maze bioassay indicated that Z. comes can follow the trail pheromone of L. fuliginosus. This suggests that Z. comes may detect other chemical signals of L. fuliginosus to keep closer interactions with the workers. Received 22 June 2001; accepted 12 November 2001.     

Response of hydrothermal vent vestimentiferan <Emphasis Type="Italic">Riftia pachyptila</Emphasis> to differences in habitat chemistry

Vestimentiferan tubeworms, which rely on intracellular sulfide-oxidizing autotrophic bacteria for organic carbon, flourish at deep-sea hydrothermal vents despite the erratic nature of their habitat. To assess the degree to which differences in habitat chemistry (sulfide, pH/CO₂) might impact host and symbiont metabolic activity, Riftia pachyptila tubeworms were collected from habitats with low (H₂S < 0.0001 mM) and high (up to 0.7 mM) sulfide concentrations. The elemental sulfur content of the symbiont-containing trophosome organ was lower in specimens collected from the low-sulfide site. Symbiont abundance, RubisCO activity, and trophosome carbon fixation rates were not significantly different for individuals collected from low- versus high-sulfide habitats. Carbonic anhydrase activities were higher in the anterior gas exchange organs of R. pachyptila from the low-sulfide habitat. Despite large differences in habitat chemistry, symbiont abundance and autotrophic potential were consistent, while the host appears to tailor carbonic anhydrase activity to environmental CO₂ availability.     

Lipids and stable carbon isotopes in two species of Hawaiian corals,<Emphasis Type="Italic"> Porites compressa</Emphasis> and<Emphasis Type="Italic"> Montipora verrucosa</Emphasis>, following a bleaching event

Mass coral bleaching events have occurred on a global scale throughout the worlds tropical oceans and can result in large-scale coral mortality and degradation of coral reef communities. Coral bleaching has often been attributed to periods of above normal seawater temperatures and/or calm conditions with high levels of ultraviolet radiation. Unusually high shallow-water temperature (>29°C) in Kaneohe Bay, Hawaii, USA, in late summer (20 August–9 September) and fall (1–7 October) of 1996 produced visible bleaching of two dominant corals, Porites compressa Dana, 1864 and Montipora verrucosa Dana, 1864. The present study examined chlorophyll a (chl a), total lipid concentrations, and lipid class composition in corals of both species in which the entire colony was non-bleached, moderately bleached, or bleached. Skeletal, host tissue, and algal symbiont ¹³C values were also measured in non-bleached and bleached colonies. In additional unevenly bleached colonies, paired samples were collected from bleached upper surfaces and non-bleached sides. Samples were collected on 20 November 1996 during the coral recovery phase, a time when seawater temperatures had been back to normal for over a month. Chl a levels were significantly lower in bleached colonies of both species compared with non-bleached specimens, and in bleached areas of unevenly bleached single colonies. Total lipid concentrations were significantly lower in bleached P. compressa compared with non-bleached colonies, whereas total lipid concentrations were the same in bleached and non-bleached M. verrucosa colonies. The proportion of triacylglycerols and wax esters was lower in bleached colonies of both species. Both bleached and non-bleached M. verrucosa had from ~17% to 35% of their lipids in the form of diacylglycerol, while this class was absent in P. compressa. ¹³C was not significantly different in the host tissue and algal symbiont fractions in non-bleached and bleached samples of either species. This suggests that the ratio of carbon acquired heterotrophically versus photosynthetically was the same regardless of condition. Skeletal ¹³C was significantly lower in bleached than in non-bleached corals. This is consistent with previous findings that lower rates of photosynthesis during bleaching results in lower skeletal ¹³C values. The two species in this study displayed different lipid class compositions and total lipid depletions following bleaching, suggesting that there is a difference in their metabolism of lipid reserves and/or in their temporal responses to bleaching and recovery.Communicated by J.P. Grassle, New Brunswick     

Ecology of<Emphasis Type="Italic"> Leptothorax</Emphasis> ants: impact of food,nest sites,and social parasites

In a long-term field manipulation, we demonstrate strong reactions of Leptothorax longispinosus ant colonies to food- and nest-site supplementation. Demographic and genetic responses varied over small geographic scales, and the two ecological factors interacted with the presence of the social parasite Protomognathus americanus. We conducted a 2×2 experiment in three blocks and found that the blocks, which were less than 100 m apart, reacted very differently to the treatments. Blocks differed in degree of polygyny, intranest relatedness, colony size, productivity, and sexual investment. Furthermore, these differences were associated with the presence of slave-making ants and the local availability of nest sites. Nest-site supplementation had a strong effect only in the site with the highest prevalence of social parasites, influencing there the density and investment patterns of colonies. L. longispinosus ants in the least parasitized area were strongly affected by both food- and nest-site supplementation. There, food supplementation led to a decrease in the number of queens per colony and consequently to an increase in intranest relatedness, while colonies in nest-site-supplemented areas invested fewer resources in males and produced a female-biased allocation ratio. By contrast, in a third block with a very low intracolonial relatedness, food supplementation induced an absolute and relative higher investment in males. We conclude that ecological factors influencing social organization in insect societies cannot be studied in isolation, because the interactions among factors produce far richer responses than any one variable.Communicated by L. Sundström     

Larval feeding by adult bumble bee workers (Hymenoptera: Apidae)

Summary Laboratory studies on colonies of Bombus terrestris (L.) and B. terricola Kby. showed that while the overall rate of larval feeding is highly correlated with total larval biomass, feeding of individual larvae is only weakly regulated. Nevertheless, the temporal distributions of inspections and feedings to larvae by adult nurse bees did provide evidence for a feedback mechanism.The behavior of individual workers engaged in larval feeding is highly contagious through time. A laboratory colony of B. terrestris compensated for the effect of experimental removal of half the worker population by increases in the feeding rate of individual workers. The results appear to be consistent with recent suggestions that overall colony performance is a mass effect resulting from the partially stochastic operations of its individual components.     

Sulfide-oxidizing bacteria in the burrowing echinoid,Echinocardium cordatum (Echinodermata)

Symbiotic filamentous bacteria thrive in the intestinal caecum of the deposit-feeding echinoid Echinocardium cordatum. Specimens of E. cordatum were collected at Wimereux (Nord Pas-de-Calais, France) in 1991. Their symbiotic bacteria build nodules by forming multilayered mats around detrital particles that enter the caecum. The morphological features of the bacteria are those of Thiothrix, a sulfide-oxidizing genus. The filaments, which may form rosettes, are sheathed and made by a succession of hundreds of rod-shaped bacteria which store elemental sulfur in the presence of external sulfide. Live bacteria are restricted to the outer layers of the nodules. Their sulfide-oxidizing activity was investigated, using a Biological Oxygen Monitor, by measuring the O₂-consumption when reduced sulfur compounds are provided. They oxidize thiosulfate and sulfide. Optimal sulfide oxidation occurs at intermediary pO₂ (100 to 160 M O₂l^-1). Spectrophotometry has confirmed that the sulfur content of the filamentous symbiotic sulfideoxidizing bacteria depends on the presence of external sulfide. This is the first report of symbiotic intradigestive Thiothrixspp.-like bacteria; it lengthens the list of symbioses between sulfide-oxidizing bacteria and invertebrates from sulfide-rich habitats.     

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

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

Competitive interactions between related clades: evolutionary implications of overgrowth interactions between encrustin cyclostome and cheilostome bryozoans

Overgrowth interactions between encrusting cyclostome (eight species) and cheilostome (over 50 species) colonies were examined in subtidal communities in the northern Adriatic Sea off Rovinj, Croatia, in May–June 1988 and September–October 1990. Cheilostome colonies occupied ca. 80% of the available substrate space, whereas the cyclostome colonies occupied <5%. Out of 210 recorded interactions, cheilostomes overlapped cyclostomes in 164, while cyclostomes overlapped cheilostomes in only 16; 30 encounters resulted in mutual overlap or at least temporary growth termination along the line of contact. Most of the recorded interactions were for the cyclostomes Diplosolen obelia (Johnston) and Plagioecia patina (Lamarck). Both species elevated their colony margins on contact with a cheilostome in some instances, but D. obelia only overgrew competing cheilostomes in 12/120 encounters, and P. patina never prevailed. I propose that the cheilostomes have a key evolutionary innovation, lacking in cyclostomes, consisting of more rapid ontogenetic development of zooids along colony margins so that encrusting cheilostomes have substantially higher colony margins. This results in an outflow of filtered water along the edge of encrusting cheilostome colonies, while cyclostomes take water in at colony margins. These contrasting feeding currents apparently give a competitive advantage to the cheilostomes where colony margins approach and make contact with those of cyclostomes.     

Size and spatial structure in deep versus shallow populations of the Mediterranean gorgonian <Emphasis Type="Italic">Eunicella singularis</Emphasis> (Cap de Creus,northwestern Mediterranean Sea)

In the Western Mediterranean Sea, the gorgonian Eunicella singularis (Esper, 1794) is found at high densities on sublittoral bottoms at depths from 10 to 70 m. Shallow colonies have symbiotic zooxanthellae that deeper colonies lack. While knowledge of the ecology of the shallow populations has increased during the last decades, there is almost no information on the ecology of the deep sublittoral populations. In October and November 2004 at Cap de Creus (42°19′12″ N; 03°19′34″ E), an analysis of video transects made by a remotely operated vehicle showed that shallow populations (10–25 m depth) were dominated by small, non-reproductive colonies, while deep sublittoral populations (50–67 m depth) were dominated by medium-sized colonies. Average and maximum colony heights were greater in the deeper populations, with these deeper populations also forming larger patch sizes and more extensive regions of continuous substrate coverage. These results suggest that shallow habitats are suitable for E. singularis, as shown by the high recruitment rate, but perturbations may limit or delay the development of these populations into a mature stage. This contrasts with the deep sublittoral habitats where higher environmental stability may allow the development of mature populations dominated by larger, sexually mature colonies.