Chemically mediated interactions between the red algaPlocamium hamatum (Rhodophyta) and the octocoralSinularia cruciata (Alcyonacea)

Interactions between the red algaPlocamium hamatum J. Agardh (Rhodophyta) and other benthic organisms including the alcyonacean soft coralSinularia cruciata (Tixier-Durivault) were investigated on an inshore fringing reef environment in whichP. hamatum was the dominant large fleshy alga. Field observations of sessile reef organisms including octocorals and sponges living in close proximity toP. hamatum revealed that varying degrees of tissue necrosis were suffered by the invertebrates when in physical contact with the alga. In order to establish whether the chemical constituents of the alga, especially chloromertensene, played a role in this necrosis, manipulative field experiments were carried out in the Pelorus Channel, Palm Island group (18°34S; 146°29E), North Queensland, Australia, in November and December 1988. The first experiment involved the relocation of healthy plants and soft corals into contact and non-contact situations on a mesh grid. In all cases of contact betweenP. hamatum andS. cruciata, the soft coral suffered tissue necrosis (n=6,p=0.0022). The second experiment had the same design, but involved the use of artificial plants both uncoated and coated with natural levels of chloromertensene, in contact withS. cruciata. In all cases of contact with coated treatments, necrosis was observed inS. cruciata (n=4,p=0.025). In cases where uncoated artificial fronds were placed in contact with soft corals,S. cruciata showed minor abrasion effects, but no appreciable necrosis. Coated treatments were not fouled by epiphytes during the experiment and were not consumed by predators. Uncoated treatments were rapidly reduced in size by predation and any remaining material was biofouled. These experiments thus demonstrated that the deleterious effects observed in soft corals in the field were caused by contact with the algaP. hamatum, that these effects were indeed chemically mediated by chloromertensene, and that physical contact without chemical intervention caused no such deleterious effects. This is the first experimental evidence which conclusively demonstrates allelopathy between an alga and other marine organisms and identifies the compound responsible for the observed allelopathic effects.     

The effects of different strains of zooxanthellae on the secondary-metabolite chemistry and development of the soft-coral host Lobophytum compactum

We have assessed the secondary-metabolite chemistry of freshly metamorphosed coral polyps, with and without zooxanthellae, using extremely sensitive electro-spray and Fourier-transform mass spectrometry. Coral larvae of the soft coral Lobophytum compactum of the same genetic background were reared, then inoculated with zooxanthellar strains of different taxonomic and geographic origin, and their terpenoid chemistry analysed. The identification of isolobophytolide in individuals of all treatment groups, including aposymbiotic control polyps, demonstrates that control of terpene production lies with the host coral and not their symbiotic algae. Received: 29 November 1999 / Accepted: 22 November 2000     

Population structure, growth and recruitment of the euryalinid brittle-star Astrobrachion constrictum (Echinodermata: Ophiuroidea) in Doubtful Sound, Fiordland, New Zealand

The euryalinid brittle-star (snake star) Astrobrachion constrictum (Farquhar) lives coiled around the branches of black coral (Antipathes fiordensis) colonies. Twenty-two vertical transects, 10 m wide by 30 m deep, were swum in Doubtful Sound over a 2.5 yr period from 1993 to 1995. Numbers, disc diameters and colour morphotypes of brittle-stars inhabiting coral colonies were recorded. 36.3% of the coral colonies >200 mm tall (n = 292) hosted ≥1 Astrobrachion constrictum (range 0 to 12). Overall, the population was patchily distributed on the available coral habitat. The dark red colour morphotype of A. constrictum was most common (87%, n = 279) followed by the yellow, striped, and then spotted varieties. The population was comprised mainly of large (≥10 mm disc diam) individuals, and juveniles were rarely encountered, indicating low rates of recruitment or a high mortality of recruits. Disc-diameter data gathered from this and previous studies indicated that growth in A. constrictum is initially rapid, with individuals reaching a disc diameter of 15 mm in ≃2.5 yr; growth decreases with age, as in other deep-sea ophiuroids. Growth rate within years, however, was not constant, with faster growth in the spring/summer. Maximum size for A. constrictum is reached in ≃8 yr at ˜23 mm disc diam. Anecdotal evidence indicates that A. constrictum may not be confined solely to black coral colonies. Received: 25 September 1996 / Accepted: 16 October 1996     

Seasonal and sex-specific variations in levels of photo-protecting mycosporine-like amino acids (MAAs) in soft corals

This study focusses on the nature and extent of variation in mycosporine-like amino acids (MAAs) in relation to annual cycles in solar radiation, seawater temperature, and reproduction in reef-flat populations of two soft coral species. The results show MAA tissue concentrations in shallow water colonies of Lobophytum compactum and Sinularia flexibilis to be significantly correlated to annual cycles in solar radiation (P<0.0006 and P<0.0005, respectively) and seawater temperature (P<0.0006 and P<0.0004, respectively). Evidence of seasonal cycles in MAA levels in the tissues of shallow-reef invertebrates positively correlating with annual cycles in solar radiation and temperature suggests that they are an integral component of the soft corals' biochemical defence system against high irradiance and/or temperature stress and thus bleaching. This is further corroborated by the higher production of MAAs in females than males prior to spawning (up to 67% and 56% for L. compactum and S. flexibilis, respectively), presumably to provide a high level of protection against irradiance stress for progeny.     

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     

Stimulation of fat-body production in the polyps of the coralPocillopora damicornis by the presence of mutualistic crabs of the genusTrapezia

A mutualism exists between the xanthid crabs of the genusTrapezia and their host corals,Pocillopora damicornis. It has previously been established that these obligate coral residents benefit the coral hosts by defending them against echinoderm predators and by increasing the survival of polyps located deep between the coral branches. In turn, the corals apparently benefit the crabs by producing lipid-filled structures on which the trapezid crabs feed; these fat bodies may contain some of the lipid which in previous studies of coral metabolism has been termed excess. It was determined by experiments conducted at the Hawaii Institute of Marine Biology that the presence of crabs in colonies ofP. damicornis stimulates the polyps to produce the lipid-filled fat bodies; removal of crabs causes corals to cease producing fat bodies. A structure very similar to the fat bodies ofP. damicornis has been reported inAcropora durvillei. Both of these coral genera ordinarily possess xanthid-crab mutualists. This association between branching corals and crustaceans may have evolved because corals of these genera provide shelter among their branches and because these shallow-water corals are evidently capable of releasing lipid which is excess to the corals' metabolic needs, but which can be utilized by the crabs.     

Algal overgrowth of alcyonacean soft corals

Colonies of the soft coral Lobophytum pauciflorum (Ehrenberg, 1834) (Coelenterata: Octocorallia: Alcyonacea: Alcyoniidae), some of which were heavily overgrown by the algae Ceranium flaccidum (Kuetzing) Ardissone and Enteromorpha sp., and other minor epizoites, were collected at Taylor Reef (17°50S; 146°35E) in the Great Barrier Reef. Overgrown colonies contained the diterpene 2-epi-sarcophytoxide as the major secondary metabolite, while conspecific colonies with clean polyparies contained two diterpenes in approximately equal amounts: 14-hydroxycembra-1,3,7,11-tetraene and 15-hydroxycembra-1,3,7,11-tetraene. By contrast, twenty conspecific pairs of overgrown and clean colonies of other alcyoniid soft corals collected from Pelorus Channel, Palm Island Group (18°34S; 146°29E), showed no chemical differences within in the pairs. Cultures of a common species of Ceramium [C. codii (Richards) Mazoyer] were incubated with different concentrations of nine soft-coral-derived diterpenes and significant algal growth inhibition was observed in many cases. It appears that terpenoids from soft coral may contribute to the lack of epizoic organisms on soft-coral polyparies.     

Population genetic and phylogeographic structure of wahoo,<Emphasis Type="Italic"> Acanthocybium solandri</Emphasis>, from the western central Atlantic and central Pacific Oceans

The wahoo, Acanthocybium solandri (Cuvier, 1832), is a pelagic, highly migratory, scombroid fish, distributed worldwide throughout tropical and warm temperate seas. To evaluate population genetic and phylogeographic structure against a null hypothesis of panmixia, the entire mitochondrial DNA control-region (~890 base pairs) was sequenced for 231 wahoo. Samples were collected from 1997 to 2001 from seven sites: North Carolina (NC; n=23), east central Florida (CF; n=30), Bimini, Bahamas (BB; n=40), southern tip of Florida (SF; n=21), Cayman Islands (CI; n=23), northern Gulf of Mexico (NG; n=54), and Hawaii (HI; n=40). Inter-annual samples were obtained from four of these locations (NC, BB, SF, NG). Seventeen haplotypes were shared by individuals within and among samples; 187 singleton haplotypes were observed. Within-sample haplotype diversities ranged from 0.995 to 1.000 (overall h=0.999) and within-sample nucleotide diversities ranged from 0.049 to 0.055 (overall =0.053). A neighbor-joining tree based on inter-haplotypic distances revealed two monophyletic lineages differing by 13.6% nucleotide divergence. Nested within each major lineage were several, well-supported subclades. There was no evidence of temporal heterogeneity in haplotype distributions. Partitioning mtDNA variation, 99.75% of the variance was within samples and 0.25% (P=0.307) between samples; the fixation index (_ST=0.0025) was not significant. Likewise, pairwise _ST values were low or negative, and none were significant on a table-wide basis. Exact tests for sample differentiation in haplotypes were also non-significant. All population analyses were consistent with the null hypothesis of panmixia. However, analytical power was limited by sample size. Mismatch distributions were inconsistent with expected distributions based on sudden-expansion and static-growth models. Wahoo exhibit concurrently high haplotype and nucleotide diversities, presumably a consequence of secondary contact between historical subpopulations rather than a long, stable evolutionary history. Given the level of geographic and individual sampling, wahoo thus far represent the sole example of a scombroid or xiphioid fish exhibiting coarse-grain genetic homogeneity across a broad, inter-oceanic range despite a deeply coalescing genealogical structure. Accordingly, cooperative fishery management on a broad, inter-ocean scale may be warranted.Communicated by J.P. Grassle, New Brunswick     

Reproduction and reproductive variability in the coral Pocillopora verrucosa from the Republic of Maldives

The sexuality, reproductive mode, and timing of reproduction of Pocillopora verrucosa from the Republic of Maldives, Indian Ocean, were assessed using serial histological sections. These showed that P. verrucosa is an annual simultaneous hermaphrodite, with gonads arranged in two opposing arcs of alternating testes and ovaries, six gonads in each arc. No planulae were observed in any dissection or histological analysis carried out, therefore making brooding unlikely. Broadcast spawning is inferred from the disappearance of mature gametes from samples collected between late March and April 1991. Mean oocyte size at spawning was 53.5 m and mean potential fecundity was 7300 ooctyes cm^-2yr^-1. The reproductive pattern of P. verrucosa in the Maldives is compared to that of the same species in different locations, the short breeding season in March to April occurring earlier in the Maldives than in Red Sea populations. The maximum mean oocyte diameter found in Maldivian specimens was much smaller than elsewhere. The year-to-year variation in numbers of oocytes/polyp within single colonies, and the variation between colonies within the population was significant (p<0.01). Therefore, it is possible that fecundity is not a good variable to use when monitoring stress on coral reefs unless larger numbers of estimates can be routinely made. Some colonies contained immature oocytes at spawning that were not released and continued to grow to approximately twice the size of the spawned oocytes that were presumed to be mature. These unspawned oocytes were oosorbing, and were characterised by the presence of zooxanthellae and large numbers of vacuoles in their cytoplasm. This gradual expansion and oosorption of unspawned oocytes has not been documented previously for corals.     

Variation in UVB sensitivity of planula larvae of the coral Agaricia agaricites along a depth gradient

The effects of natural intensities of ultraviolet A (UVA, 320 to 400 nm) and B (UVB, 280 to 320 nm) radiation on planktonic planula larvae of the reef-building coral Agaricia agaricites (Linnaeus) were investigated through field experiments. Survival, chlorophyll concentrations, and solubilized protein concentrations were determined for larvae spawned from colonies at 3 and 24 m depth and subjected to one of three light regimes at 3, 10, or 24 m depth for 72 h: PAR (photosynthetically active radiation, 40- to 700 nm) only, PAR+UVA, or PAR+UVA+UVB. At 3 m depth, larvae in the PAR+UVA+UVB treatment showed lower survivorship than larvae exposed to either PAR alone or PAR+UVA. Within the PAR+UVA+UVB treatment at 3 m depth, larvae from colonies at 24 m depth suffered higher mortality than those from 3 m. Differences in survivorship between larvae originating from 3 and 24 m depth corresponded with tissue concentrations of UVB-protective mycosporine-like amino acids: larvae from 3 m had higher concentrations of mycosporine-glycine (_max=310nm) and palythine (_max=320nm) than those from 24 m depth. Chlorophyll concentrations were inversely correlated with PAR intensities, but variation in this parameter did not appear to be detrimental. These results show that sensitivity to high intensities of UVB radiation may affect survival of A. agaricites larvae in shallow reef-waters.     

Depth-dependent photoadaption by zooxanthellae of the reef coral Montastrea annularis

Zooxanthellae living in colonies of the Caribbean reef coral Montastrea annularis photoadapt to depth-dependent attenuation of submarine light. Studies carried out at Discovery Bay, Jamaica, show that in shallow-living coral colonies, the zooxanthellae appear photoadapted to function at high light intensities, and do poorly if transplanted to low light intensities; in contrast, zooxanthellae in deeper-living coral colonies can be damaged by high light intensities. The adaptation to decreasing light intensity and changing spectral quality appears to be accomplished by increasing the size of the photosynthetic unit (PSU), as opposed to increasing the number of PSU's per cell. Whole cell absorption increases with depth, partially offsetting the loss of light energy due to depth-dependent attenuation. Calculations of photosynthetically usable radiation, the light an alga is capable of absorbing in its own submarine habitat, suggest that the algae at different depths are optimizing rather than maximizing their ability to harvest submarine light energy.     

Background levels of soil beryllium in several countries

Beryllium and aluminium contents in uncontaminated soils from six countries are reported. The means and ranges of beryllium in the surface soils were as follows: 1.43(0.20–5.50)g g^–1 in Thailand (n=28), 0.7 (0.31–1.03) g g^–1 in Indonesia (n=12), 0.99(0.82–1.32) g g^–1 in New Zealand (n=3), 0.58(0.08-1.68)g g^–1 in Brazil (n=16), 3.52(2.49–4.97)g g^–1 in the former Yugoslavia (n=10), and 1.56(1.01–2.73) g g^–1 in the former USSR (n=8). The mean and range of beryllium contents of the surface soils in Japan (1.17(0.27–1.95)g g^–1 n=27) are situated within the values of the soils from these countries except for the Yugoslav soils derived from limestones. The mean of the mean beryllium contents of the surface soils in all these countries is 1.42 g g^–1 which will be used as a tentative average content of beryllium in uncontaminated surface soils, except for the soils derived from parent materials high in beryllium content. The beryllium contents of the subsoils were higher than those of the surface soils in New Zealand and Yugoslavia as is the case with Japan. The correlation coefficient between the contents of beryllium and aluminium in all the soil samples (n=113) including surface soils and subsoils was 0.505 (p < 0.001).     

Influences of habitat and natural disturbances on contributions of massive Porites corals to reef communities

We compared densities, distributions and size frequencies of massive corals in the genus Porites on five relatively exposed, mid-shelf reefs (50 km offshore) in the central Great Barrier Reef with those on a sheltered inshore reef (10 km offshore). Data included various transect and mapping studies between 1984 and 1990, estimates of size-dependent damage from the crown-of-thorns starfish Acanthaster planci, estimated densities of herbivorous sea urchins (potential predators of juveniles), and observations of size-specific effects of tropical cyclones. Assemblages of Porites spp. on mid-shelf reefs were dominated by small colonies (2 to 10 cm diam) established either from planula larvae or from small tissue remnants that had survived A. planci predation in the early to mid-1980s. Large colonies (up to 10 m diam) were scarce, except for localized aggregations on terraces at the base of reef slopes (6 to 12 m deep). Extensive space suitable for settlement by coral larvae can be attributed to recurrent cyclones and A. planci outbreaks. Despite low sea urchin predation, the slowly growing Porites juveniles are likely to die from overgrowth by numerous, much faster growing corals. On the sheltered inshore reef, the coral community was dominated by very large (>5 m diam) Porites colonies, several centuries old; recruitment was mainly by fragmentation of large colonies; there was little space available for settlement, and probabilities of juvenile mortality from grazing urchins were high. Differences in settlement and early survival of Porites spp. are exacerbated by different regimes of storm damage. A model is proposed that links wave climate with the size and age reached by corals before dislodgement by storm waves, and which is consistent with observed densities and size-frequency distributions of Porites in sheltered and exposed areas.     

Effects of feeding frequency and symbiosis with zooxanthellae on nitrogen metabolism and respiration of the coral Astrangia danae

Colonies of the temperate coral Astrangia danae occur naturally with and without zooxanthellae. Basal nitrogen excretion rates of nonsymbiotic colonies increased with increasing feeding frequency [average excretion rate was 635 ng-at N (mg-at tissue-N)^-1 h^-1]. Reduced excretion rates of symbiotic colonies were attributed to N uptake by the zooxanthellae. Nitrogen uptake rates of the zooxanthellae averaged 8 ng-at N (10⁶ cells)^-1 h^-1 in the dark and 21 ng-at N (10⁶ cells)^-1 h^-1 at 200 Ein m^-2 s^-1. At these rates the zooxanthellae could provide 54% of the daily basal N requirement of the coral if all of the recycled N was translocated. Basal respiration rates were 172 nmol O₂ cm^-2 h^-1 for starved colonies and 447 nmol O₂ cm^-2 h^-1 for colonies fed three times per week. There were no significant differences between respiration rates of symbiotic and nonsymbiotic colonies. N excretion and respiration rates of fed (symbiotic and nonsymbiotic) colonies increased greatly soon after feeding. N absorption efficiencies decreased with increasing feeding frequency. A N mass balance, constructed for hypothetical situations of nonsymbiotic and symbiotic (3×10⁶ zooxanthellae cm^-2) colonies, starved and fed 15 g-at N cm^-2wk^-1, showed that the presence of symbionts could double the N growth rate of feeding colonies, and reduce the turnover-time of starved ones, but could not provide all of the N requirements of starved colonies. Rates of secondary production, estimated from rates of photosynthesis and respiration were similar to those estimated for reef corals.     

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.     

Photosynthetic characteristics of Prochloron sp./ascidian symbioses

The prokaryotic green alga Prochloron sp. (Prochlorophyta) is found in symbiotic association with colonial didemnid ascidians that inhabit warm tropical waters in a broad range of light environments. We sought to determine the light-adaptation features of this alga in relation to the natural light environments in which the symbioses are found, and to characterize the temperature sensitivity of photosynthesis and respiration of Prochloron sp. in order to assess its physiological role in the productivity and distribution of the symbiosis. Colonies of the host ascidian Lissoclinum patella were collected from exposed and shaded habitats in a shallow lagoon in Palau, West Caroline Islands, during February and March, 1983. Some colonies from the two light habitats were maintained under conditions of high light (2 200 E m^–2 s^–1) and low light (400 E m^–2 s^–1) in running seawater tanks. The environments were characterized in terms of daily light quantum fluxes, daily periods of light-saturated photosynthesis (H_sat), and photon flux density levels. Prochloron sp. cells were isolated from the hosts and examined for their photosynthesis vs irradiance relationships, respiration, pigment content and photosynthetic unit features. In addition, daily P:R ratios, photosynthetic quotients, carbon balances and photosynthetic carbon release were also characterized. It was found that Prochloron sp. cells from low-light colonies possessed lower chlorophyll a/b ratios, larger photosynthetic units sizes based on both reaction I and reaction II, similar numbers of reaction center I and reaction center II per cell, lower respiration levels, and lower P_max values than cells from high-light colonies. Cells isolated from low-light colonies showed photoinhibition of P_max at photon flux densities above 800 E m^–2 s^–1. However, because the host tissue attenuates about 60 to 80% of the incident irradiance, it is unlikely that these cells are normally photoinhibited in hospite. Collectively, the light-adaptation features of Prochloron sp. were more similar to those of eukaryotic algae and vascular plant chloroplasts than to those of cyanobacteria, and the responses were more sensitive to the daily flux of photosynthetic quantum than to photon flux density per se. Calculation of daily minimum carbon balances indicated that, though high-light cells had daily P:R ratios of 1.0 compared to 4.6 for low-light cells, the cells from the two different light environments showed nearly identical daily carbon gains. Cells isolated from high-light colonies released between 15 and 20% of their photosynthetically-fixed carbon, levels sufficient to be important in the nutrition of the host. Q₁₀ responses of photosynthesis and respiration in Prochloron sp. cells exposed briefly (15–45 min) to temperatures between 15° and 45°C revealed a discontinuity in the photosynthetic response at the ambient growth temperatures. The photosynthetic rates were found to be more than twice as sensitive to temperatures below ambient (Q₁₀=3.47) than to temperatures above ambient (Q₁₀=1.47). The Q₁₀ for respiration was constant (Q₁₀=1.66) over the temperature range examined. It appears that the photosynthetic temperature sensitivity of Prochloron sp. may restrict its distribution to warmer tropical waters. The ecological implications of these findings are discussed in relation to published data on other symbiotic systems and free-living algae.     

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     

Nestmate recognition and the ontogeny of acceptability in the ant,Leptothorax curvispinosus

Summary In social insects, there is often a brief period following eclosion when workers are highly acceptable in alien nests of their own or other species. This study tested for such an acceptance period in the facultatively polygynous ant, Leptothorax curvispinosus, and compared the duration and effectiveness of this period for conspecific and heterospecific introductions. Workers that eclosed and aged for 1–70 h or 30 days in isolation were introduced into either their parental nests (n=24), alien conspecific nests (n=265), or nests of the closely related and biologically similar species, L. longispinosus (n=341). In alien conspecific nests, acceptance was maximal for workers aged 1–12 h at introduction (67.7% not attacked, 75.8% adopted) and gradually decreased until the level of nonaggression (after 60 h) and adoption (after 36 h) were not significantly different from 30-day-old workers (5.9% not attacked, 17.6% adopted). In heterospecific nests, acceptance was maximal for workers aged 1–4 h at introduction (34.8% not attacked, 37.0% adopted) but thereafter was not significantly different from 30-day-old workers (5.6% not attacked, 8.3% adopted). In their parental nests, workers were generally accepted regardless of age (4–56 h posteclosion, 95.8% not attacked, 100% adopted); a result that is consistent with previous research on older workers (38–157 days posteclosion). This study demonstrates an acceptance period that is more effective and of longer duration within than between these species but that, under uniform laboratory conditions, is often not necessary for the integration of workers into their parental colonies. Within colonies, acceptance periods might only be important during relatively brief periods in a colony's life history when eclosing workers produce genetically based nestmate recongition cues that are not already represented in the colony and must be learned by colony members (e.g., during early colony growth or following adoption of queens), or when young workers must acquire environmentally based nestmate recognition cues to achieve and maintain acceptability.     

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     

Influence of phosphate depletion on the growth and colony formation of Phaeocystis pouchetii

The haptophycean alga Phaeocystis pouchetii (Hariot) Lagerheim, a bloom-forming species in the North Sea, formed massive colonies in batch cultures only when the phosphate concentration in the medium was below 1 mol l^-1. In general, colony cells and single cells showed a similar response to phosphate depletion: decreased cellular phosphate (up to a factor of 32), decreased chlorophyll-a concentration (up to a factor of 4.2) and high activities of alkaline phosphatase (APA). However, the growth rate of colony cells was reduced at low phosphate concentrations (<1 mol l^-1) in contrast to that of single cells. Colony cells tended to have a slightly higher phosphate concentration than single cells, while the increase in APA was delayed. The differences between single cells and colonies in phosphate utilization, cellular phosphate content, and growth rate cause, at low phosphate levels, a shift towards the formation of colonies. Similar changes seem to occur in nature during transition of nutrient-sufficient to nutrientlimited conditions.This investigation was supported by the Foundation for Fundamental Biological Research (BION), which is subsidized by the Dutch Organization for the Advancement of Pure Research (ZWO). Publication No. 8 of the project Ecological Research of the North Sea (EON)