Ingestion of pico- and nanoplankton by the Mediterranean red coral Corallium rubrum

This study investigates the feeding behaviour of the precious red coral Corallium rubrum on bacterioplankton. The effects of flow rate, prey concentration, and seawater temperature were tested. The results obtained show that C. rubrum was able to prey on both pico- and nanoplankton cells. Flagellates constituted the major bacterioplankton food source in terms of carbon and nitrogen, representing from 43 to 70% of the C and N ingested. Flow speed (2, 6, and 11 cm s⁻¹) had no effect on grazing rates, maybe due to the small size of the ingested particles. Conversely, feeding rates increased with prey concentration and seawater temperature. There was a doubling of the picoplankton ingestion rate for a sixfold increase in its concentration. The ingestion of autotrophic flagellates, however, increased at the same time as their concentration, indicating a preference for this type of food. Considering the range of concentrations typically found in the Ligurian Sea, the ingestion of pico- and nanoplankton brings 148 ng C polyp⁻¹ day⁻¹ and 28 ng N polyp⁻¹ day⁻¹. This type of food represents only ca. 4.5% of the total carbon gained by C. rubrum from the different sources, but might be the most important in terms of nitrogen, phosphorus, and other essential elements.     

Diet and seasonal prey capture rates in the Mediterranean red coral (Corallium rubrum L.)

Gorgonians are passive suspension feeders, contributing significantly to the energy flow of littoral ecosystems. More than in active suspension feeders (such as bivalves, ascidians and sponges) their prey capture is affected by spatial and temporal prey distribution and water movement. Corallium rubrum is a characteristic gorgonian of Mediterranean sublittoral hard bottom communities. This study found a high variability in the annual cycle of prey capture rate, prey size and ingested biomass, compared to other Mediterranean gorgonians. Detrital particulate organic matter (POM) was found throughout the year in the polyp guts and constituted the main proportion of the diet (25–44%). Crustacean fragments and copepods (14–46%) accounted for the second major proportion, while invertebrate eggs (9–15%) and phytoplankton (8–11%) constituted the smallest part of the diet. To verify the importance of detrital POM in the energy input of this precious octocoral species, in situ experiments were carried out during the winter–spring period. The results confirm the importance of detrital POM as the main source of food for C. rubrum [0.13±0.04 μg C polyp⁻¹ h⁻¹ (mean±SD)]. This study also compares the prey capture rates of two colony size classes and two depth strata: Within the same patch, small colonies (<6 cm height) captured significantly more prey per polyp (0.038±0.09 prey polyp⁻¹ h⁻¹) than larger colonies (>10 cm high) (0.026±0.097 prey polyp⁻¹ h⁻¹) and showed a higher proportion of polyps containing prey (17% compared to 10%). Comparing colonies of similar size (<6 cm height) revealed that the colonies situated at 40 m depth captured significantly more prey (0.038±0.09 prey polyp⁻¹ h⁻¹) than the ones at 20 m (0.025±0.11 prey polyp⁻¹ h⁻¹). One pulse of copepods was recorded that constituted 16% of all captured prey during the 15-month period studied in one of the sampled populations. The data suggest that the variability of hydrodynamic processes may have a higher influence on the feeding rate than seasonal changes in the seston composition. The carbon ingestion combined with data on the density of the exploited population results in 0.4–9.6 mg C m⁻² day⁻¹. The grazing impact of current, heavily exploited and small-sized populations is comparable to that of larger Mediterranean gorgonians, suggesting that unexploited red coral populations may have a high impact compared with other passive suspension feeders.     

Demographic parameters of two populations of red coral (Corallium rubrum L. 1758) in the North Western Mediterranean

The demographic and reproductive structure of populations represents the main data set needed for conservation and management plans. Mediterranean red coral Corallium rubrum has been exploited for 2,000 years, but only recently management plans have been solicited by the international community. We examined and compared the demographic features of two red coral shallow populations located in distinct geographic locations: Portofino (Italy) and Cap de Creus (Spain). Adults and juveniles density, growth rates, population size and age structure, fecundity and fertility were examined. Juveniles were the dominant class (33 %) in both populations. The analysis of the gamete content of 653 colonies revealed that the populations have balanced sex ratios and similar fertility and polyp fecundity. The average annual growth rate, determined on 119 colonies by annual growth rings count, was similar in both populations (0.24 mm year^?1), decreasing with colony age. Maximum life span of 99 % of the colonies was 60 and 40 years at Portofino and Cap de Creus, respectively. Minimum harvestable size (7 mm basal diameter) was reached in 30–35 years, and the percentage of colonies above it was 6.7 % at Portofino and 2.1 % at Cap de Creus, where juvenile and adult colony densities were significantly lower and the percentage of commercial-sized colonies reduced by 25 % in just a few years. Notwithstanding similar growth and fecundity, the two populations showed different densities and size/age structures suggesting local factors, together with different fishing pressures, have to be taken into account in the management plans for this species.     

Population structure of an exploited benthic cnidarian: the case study of red coral (Corallium rubrum L.)

Octocorals are an important part of many ecosystems as they add three-dimensional complexity to the benthos and thereby increase biodiversity. The Mediterranean red coral (Corallium rubrum, L. 1758) is a longevous octocoral that is harvested commercially, yet natural and anthropogenic influences on its population size structure are little understood. This study found that some harvested red coral populations had a significantly different size structure when compared to populations at the nearby Marine Protected Area (MPA) of Medas Islands at the Spanish Costa Brava (NW Mediterranean). Eighty-nine percent of the red corals in the harvested Costa Brava area are less than 10 years old and 96% of all colonies have not yet grown more than second-order branches. The size/age distribution of the harvested population is notably skewed towards younger and smaller colonies. Thus, although red coral is still abundant, its population structure is strongly distorted by harvesting. The results confirm that MPAs are useful to distinguish between anthropogenic and natural influences on population structure. However, 14 years of protection appears to be an insufficient recovery time for a longevous octocoral population such as red coral.     

On the potential use of magnesium and strontium concentrations as ecological indicators in the calcite skeleton of the red coral (Corallium rubrum)

 The variability of magnesium, strontium and calcium concentrations in the two skeleton types (sclerites and axis) of the red coral (Corallium rubrum) was assessed by using X-ray fluorescence spectroscopy and microprobe analysis as a prerequisite for their use as ecological indicators. Axis cross sections showed light and dark circular growth bands corresponding to fast and slow growth, respectively. Using microprobe analyses the Mg and Sr concentrations ranged from 2.8 to 3.0% and from 0.21 to 0.29%, respectively, in the sclerites and from 2.4 to 3.0% and from 0.1 to 0.28%, respectively, in the axis. Mg/Ca and Sr/Ca ratios varied within sclerites (average, 12% for both ratios) and within the axis of single colonies (average, 20% for Mg/Ca and 48% for Sr/Ca). Sr/Ca ratios in the axis were lower in fast-growing branch tips than in older, more basal parts of the colonies, whereas the Mg/Ca ratio did not differ significantly between colony regions. Mg/Ca and Sr/Ca ratios in the axis decreased significantly with depth, and we estimated an increase of the Mg/Ca ratio of 0.004–0.006 per degree Celsius. In all depth layers, Sr/Ca ratios showed a significant direct relationship with skeleton density in axis cross sections, whereas there was no significant relationship for Mg/Ca. Overall, our data indicate that temperature promotes the incorporation of Mg in C. rubrum as in other calcite skeletons, whereas Sr concentrations are inversely related to growth rate. This preliminary study suggests that Mg and Sr concentrations in the axis of the red coral have a strong potential as ecological indicator for temperature and growth rate. Received: 3 March 2000 / Accepted: 30 August 2000     

Effects of spatial variability and colony size on the reproductive output and gonadal development cycle of the Mediterranean red coral (Corallium rubrum L.)

Red coral (Corallium rubrum, L. 1758) is an over-exploited Mediterranean gorgonian. The gonadal development cycle of this gorgonian is examined at the Costa Brava (NW Mediterranean) taking into account for the first time colony size, depth and spatial horizontal variability. This study compares the gonad development and fertility in two colony size classes (colonies <6-cm height, and >10-cm height, both at 40–45-m depth), and two populations at different depths (16–18-m depth, and 40–45-m depth, both consisting of <6-cm high colonies) in a 15-month period. The fertility of seven size classes (<2 cm to >12 cm high colonies, in 2 cm intervals) was examined in the deep population, where large colonies were present. Furthermore, reproductive output was compared in 6 populations (distributed along more than 70-km coastline) one month before spawning (June). Red coral was found to be dioecious and gonochoric with a sex ratio of 1:1, which differs from other NW Mediterranean populations. On the other hand, fertility of different size classes indicates that small colonies of 2-cm height already produce gonads, which is in line with previous studies. Female and male polyp fertility and sperm sac size increase significantly with colony size [sperm sac diameter: 476±144 μm (mean±SD) and 305±150 μm in the >10-cm and <6-cm height colonies, respectively), whereas no significant effect on oocyte diameter was found (oocyte diameter: 373.7±18.7 μm). Depth staggered spawning, that is, an earlier release of gonads in the shallow populations, was observed in summer 2003, coinciding with the highest temperature gradient between shallow and deep water during the study period. Colonies of <6-cm height were significantly less fertile than colonies >12 cm, thus the recommendation of this study is that a minimum height should be incorporated into fishing regulations. The six studied populations at the Costa Brava showed a comparable reproductive potential, which demonstrates little variability within the homogenous population structure and range of size classes (due to overharvesting) found at the Costa Brava. The study of reproductive output is an important tool for ecosystem management, and this work recommends basing specific exploitation laws for distinctive populations on colony size, which is found to have a larger effect on reproductive potential than mesoscale variability. An erratum to this article can be found at     

Fecundity of red coral <Emphasis Type="Italic">Corallium rubrum</Emphasis> (L.) populations inhabiting in contrasting environmental conditions in the NW Mediterranean

In this study, we examined the variability and potential patterns of fecundity in the precious Mediterranean red coral Corallium rubrum (L. 1758). A total of 12 populations were selected from the NW Mediterranean Sea. We used a hierarchical sampling design to explore fecundity patterns associated with different environmental conditions found in different cave zones (entrance vs. interior), depths (15–22 vs. 39–42 m), and geographic locations (Côte Bleue vs. Calanques). Overall, 240 apical tips from colonies (10 male + 10 female colonies per population) were analyzed. Fecundity ranged between 1.0 ± 0.7 and 3.2 ± 2.3 mature gonads per polyp in female colonies and between 2.5 ± 1.6 and 6.9 ± 2.5 mature gonads per polyp in male colonies. The fecundity of red coral varied significantly for populations dwelling in different cave zones and geographic areas but not for the examined depths. Our results contribute to the knowledge of red coral fecundity in populations not yet studied in the NW Mediterranean and elucidate significant variability in fecundity within different environmental conditions. The information on coral fecundity can contribute to the development of management and conservation plans for red coral populations.     

Survey of deep-dwelling red coral (<Emphasis Type="Italic">Corallium rubrum</Emphasis>) populations at Cap de Creus (NW Mediterranean)

The distribution and population structure of the eurybathic gorgonian Corallium rubrum were studied off Cap de Creus (Costa Brava, Northwestern Mediterranean Sea). Red coral is endemic to the Mediterranean Sea and the adjacent NE Atlantic coast, where it has been over exploited for centuries. This study presents, the first quantitative data on the spatial distribution and structure of a population extending between 50 (common SCUBA limits) and 230 m depth, and compared it with shallow populations previously studied in the same area. Different remotely operated vehicles (ROV) and two methodological approaches were employed during four cruises between 2002 and 2006: 1-Extensive surveys: sea to coast transects in which red coral density and patch frequency were recorded; 2-Intensive surveys, in which parameters describing colony morphology were recorded. Most of the hard substrate between 50 and 85 m depth was inhabited by red coral colonies, showing a patch frequency of 8.3 ± 7.9 SD patches per 100 m-transect (total transect area: 34 m²), and within-patch colony densities of 16–376 colonies m⁻² (mean of 43 ± 53 colonies m⁻²). Below 120 m depth red coral was less abundant, and rather than forming dense patches as in shallow water, isolated colonies were more common. The population structure differed between sites that are easily accessible to red coral fishermen, and remote ones (both at similar depth, 60–80 m), as colonies in easily accessible locations were smaller in height and diameter, and showed a less developed branching pattern. At shallower locations (10–50 m depth) the population structure was significantly different from those at deeper locations, due to the heavy harvesting pressure they are exposed to in the shallows. Twenty-five to forty-six percentage of the deeper colonies were taller than 6 cm, while only 7–16% of the shallow water colonies exceeded 6 cm colony height. Forty-six to seventy-nine percentage of the colonies in deeper waters were large enough to be legally harvested, while only 9–20% of the shallow water colonies met the 7 mm legal basal diameter to be collected. The branching pattern was also better developed in deeper colonies, as up to 16% of the colonies showed fourth order branches, compared to less than 1% of the shallow water colonies (of which 96% consisted of only one single branch). The results thus confirm that C. rubrum populations above 50 m depth are exposed to a higher harvesting intensity than deeper populations in the same area.     

Determination of canthaxanthin in the red coral (<Emphasis Type="Italic">Corallium rubrum</Emphasis>) from Marseille by HPLC combined with UV and MS detection

Corallium rubrum is the most famous and precious coral due to the intense characteristic red colour of its skeleton. We have determined the presence of carotenoids in natural samples of C. rubrum from Marseille, Riou, and investigated their chemical composition. Analysis was performed both on soft tissues and hard tissues including spicules and skeleton. Since hard tissues are made of a mineral fraction and an organic fraction obtained after demineralization, extraction was performed both with and without demineralization by EDTA. The extracts were analyzed by complementary methods of TLC, HPLC/DAD and HPLC/MS. The components were separated by RP-18 chromatography column using acetonitrile/methanol for HPLC/DAD and acetonitrile/water gradient for HPLC/MS analysis. Our results give the first evidence of the presence of canthaxanthin, 4,4′-diketo-β-carotene, as the major carotenoid in all samples. Spicules exhibited higher content in carotenoids than skeleton. Demineralization treatment improved the efficiency of carotenoid extraction by an average factor of 5 and showed that organic matrix contains canthaxanthin.     

Sexual reproduction, larval development and benthic planulae of the solitary coral Monomyces rubrum (Scleractinia: Anthozoa)

The reproductive biology of the solitary ahermatypic coral Monomyces rubrum was studied in northeastern New Zealand between January 1996 and January 1998. The period of oogenesis lasted around 11 months, from late January to December, while spermatogenesis was more rapid, starting in late August and culminating in a spawning period in early December. Reproduction commenced at a polyp size of around 1,000 mm³ (5-6 years old) and the maximum estimated fecundity of the largest corals (7,000 mm³) was no more than 200 eggs. Oocytes were probably fertilized while within the mesentery and were shed into the coelenteron where they developed, via a solid blastula stage, for approximately 1 month. Planulae were relatively large, 3-4 mm in length and 1-2 mm diameter at the time of release, and crawled or swam immediately to the substratum. Peaks of planula shedding were semi-lunar in January 1997, but only one peak was observed in January 1998. The production of a few large rapidly settling larvae by this member of the family Flabellidae is consistent with the trend for solitary short-lived corals from other families to brood larvae rather than spawn gametes.     

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

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

Shifts in coral community structures following cyclone and red tide disturbances within the Gulf of Oman (United Arab Emirates)

This study documents the effects of two consecutive disturbances on coral community structures in the Gulf of Oman (United Arab Emirates); Cyclone Gonu in June 2007 and the Cochlodinium polykrikoides harmful algal bloom (HAB) that persisted from August 2008 until May 2009. Coral cover, colony densities, size class frequency distributions, and geometric growth rates derived from size class transition probability matrices were used to assess the post-Gonu and post-HAB recovery trajectories at four sites. The net effects of these disturbances were fourfold: (i) storm damage caused >50% losses of live branching and tabular coral cover by fragmentation and dislodgment of pocilloporid and acroporid colonies; (ii) Pocillopora damicornis colonies that survived the cyclone experienced mass mortality during the first 3?months of the HAB, resulting in localized extirpation of this species; (iii) variable Acropora mortality during the HAB indicated individual colony, rather than taxa-wide, susceptibility; and (iv) massive colony coral taxa were resistant to both disturbances.     

Cyst formation of the noxious red tide flagellate Chattonella marina (Raphidophyceae) in culture

Cyst formation of the noxious red tide flagellate Chattonella marina (Raphidophyceae), cultured from a cyst in sediments of Suo-nada, Japan in 1986, was investigated by culture experiments. Nitrogen limitation was effective in inducing cyst formation. After incubation in N-limited medium at 25°C with 10000 lux on a 14 h light, 10 h dark photo-cycle, cultures were subjected to six different light intensities (0 10000 lux) in order to determine effects of light intensity on cyst formation. Cysts were formed under light intensities of 1000 lux or below, but not at 10000 lux. Cysts formed in culture displayed morphological characteristics quite similar to those natural cysts observed in sediments collected from the Inland Sea of Japan. Observations on nucleus, using diaminidinophenylindole (DAPI)-staining and epifluorescence microscopy, indicated that the majority of cysts formed in culture were uninucleate, a few were binucleate. Germination of cysts was observed under adequate conditions for germination (22°C, 3500 lux with a 14 h light, 10 h dark photo-cycle) even after storage at 11°C in the dark for more than 4 mo.     

Growth and form in the reef-building coral Montastrea annularis

The in-situ growth rate of the reef coral Montastrea annularis on a reef at Jamaica, W. Indies was determined for a 1-year period using Alizarin Red-S staining techniques. Growth rate is correlated with water depth and growth form. The flattened growth form of M. annularis allows for continued rapid increase in colony surface area at low light intensities. The geomorphology of Jamaican reefs may in part be controlled by the population ecology of M. annularis.     

Ecological versatility and the decline of coral feeding fishes following climate driven coral mortality

Coral reefs are under threat due to climate-mediated coral mortality, which affects some reef coral genera more severely than others. The impact this has on coral reef fish is receiving increasing attention, with one focal area assessing impacts on fish that feed directly on live coral. It appears that the more specialised a species of corallivore, the more susceptible it is to coral declines. However data are sparse for the Indian Ocean, and little is known about why some corals are preferentially fed upon over others. Here I assess feeding specialisation in three species of coral feeding butterflyfish in the Chagos Archipelago, central Indian Ocean, assess the food quality of the coral genera they target and document patterns of decline in the Seychelles following a severe coral mortality event. Cheatodon trifascialis was the most specialised coral feeder, preferentially selecting for Acropora corals, however, when Acropora was scarce, individuals showed considerable feeding plasticity, particularly for the dominant Pocillopora corals. C. trifasciatus also preferentially fed on Acropora corals, but fed on a much more diverse suite of corals and also displayed some selectivity for Porites. C. auriga is a facultative corallivore and consumed ∼55% live coral, which lies within the wide range of coral dependence reported for this species. C:N ratio analysis indicated Lobophyllia and Acropora have the highest food quality, with Pocillopora having the lowest, which conforms with diet selection of corallivores and helps explain preferential feeding. Obligate specialist feeders displayed the greatest declines through coral mortality in the Seychelles with obligate generalists also declining substantially, but facultative feeders showing little change. Clearly a greater understanding of the species most vulnerable to disturbance, their habitat requirements and the functional roles they play will greatly assist biodiversity conservation in a changing climate.     

Production and role of hyaline hairs in Ceramium rubrum

Light intensity within the range of 40 to 210 E · m^-2 · s^-1 had relatively little effect on the abundance, length or distribution of Ceramium rubrum unicellular, hyaline hairs. External NH ₄ ⁺ concentrations less than 0.5 M stimulated hair initiation and growth in apical regions of the thalli. Ammonium concentrations in excess of 20 M inhibited hair formation on all regions of the thalli. Ammonium uptake velocities of plants with hairs were approximately twice those of plants without hairs. These hairs may be adaptive to take advantage of intermittent bursts of nutrients by increasing the plant's surface area and, hence, the number of nutrient uptake sites.Woods Hole Oceanographic Institution Contribution No. 5526     

Communication and coexistence in a coral community

A large number of biotic relationships among dissimilar organisms is characteristic of tropical coral reefs. Such relationships involve at least four of the 16 species associated with the coral Pocillopora damicornis (L.) at Heron Island on the Great Barrier Reef. Behavioural interactions among the most conspicuous resident species (2 fish, 1 crab and 1 shrimp) were studied in the field under natural and experimental conditions. The inhabitants have the potential to harass, capture and eat, or exclude one another from the coral habitat but a system of signals among the residents facilitates coexistence and maintains the integrity of the community. The signals include contact and shivering movements by the fish which apparently inform the crustaceans of the fish's resident status.