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.     

Net negative growth detected in a population of <Emphasis Type="Italic">Leptogorgia sarmentosa</Emphasis>: quantifying the biomass loss in a benthic soft bottom-gravel gorgonian

Gorgonian species may contribute to the three-dimensional seascape in soft bottom-gravel areas, but the information on their biology and ecology is very scarce. The biometry and secondary production of the Mediterranean soft bottom-gravel passive suspension feeder Leptogorgia sarmentosa (Cnidaria: Octocorallia) was studied using photographic monitoring of the primary branches from May 1998 to September 2000. The primary branches observed had a high density of polyps (2.2 ± 0.2 SD polyps mm⁻¹) and a high organic matter content (63.2 ± 9.1 SD %). During the two-year sampling period, there was a net negative growth in 90% of the gorgonian population. The mean loss during the 27-month period was −2.9 ± 0.9 SD cm per branch (5.7 mg C branch⁻¹). However, considering only the initial and final diameters and maximum height in the 27 months elapsed time, the gorgonians showed positive growth, which meant that photographic sampling of single branches was a more appropriate method for gorgonian secondary production monitoring. A water mass anomaly detected in 1999 in the north-western Mediterranean Sea may have been the cause of the net negative growth in L. sarmentosa in the studied area. Partial mortality due to different factors, such as strong currents, predation, disease, etc., could be a common strategy in sessile colonial benthic populations that would facilitate their maintenance even during very stressful circumstances.     

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.     

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     

Life cycle of the jellyfish <Emphasis Type="Italic">Rhizostoma pulmo</Emphasis> (Scyphozoa: Rhizostomeae) and its distribution,seasonality and inter-annual variability along the Catalan coast and the Mar Menor (Spain,NW Mediterranean)

Rhizostoma pulmo is one of the most abundant scyphomedusae along the Mediterranean coasts. To understand changes in the population densities of the medusa stage and its relationship with the benthic stage, we describe all developmental stages from the life cycle of R. pulmo, from the scyphistoma stage to young medusae reared in the laboratory. Mature scyphistomae showed a mean total body length of 1.7 mm and the average tentacle number was 16. Asexual reproduction occurred by lateral budding, podocyst and pedalocyst production. Each strobila released up to 8 ephyrae that had a mean total body diameter of 3.5 mm. Moreover, we also present data on the temporal and spatial distribution of the species in the Catalan Sea and the coastal lagoon on Mar Menor (NW Mediterranean) during the years 2007–2009. In the Catalan Sea, the presence of adult R. pulmo was recorded as extended along the whole coast, but most of the observation was concentrated in the central area; the highest abundances were recorded during the months of July and August. The highest number of observations was detected in 2008 in coincidence with the inter-annual variation observed in the Mar Menor lagoon.     

Spatial variability in reproductive cycle of the gorgonians <Emphasis Type="Italic">Paramuricea clavata</Emphasis> and <Emphasis Type="Italic">Eunicella singularis</Emphasis> (Anthozoa,Octocorallia) in the Western Mediterranean Sea

Paramuricea clavata (Risso, 1826) and Eunicella singularis (Esper, 1794) are the most representative gorgonian species in hard bottoms sublittoral communities in the Western Mediterranean Sea. Reproductive cycles of two populations of both species were studied in two distinct locations approximately 600 km apart (Medes Islands and Cape of Palos), in order to assess interpopulation variability on a relevant geographic scale. Seasonal variation of lipid concentration levels in the gorgonian tissue was used as a tool to quantify energy storage by each studied population in order to explain possible interpopulation differences in gonadal output. Sex ratio in Medes Islands populations of both species was 1:1, while in Cape of Palos sex ratio was significantly male biased (1:7) in P. clavata, and female biased (1.7:1) in E. singularis populations. Spawning timing occurred in all cases coinciding with a marked increase in sea-water temperature in spring, and after the most successful feeding season, but comparing localities there was a clear temporal shift in the time of gametes release, appearing well linked to the shift in sea-water temperature rising in spring in both sites at the depth where populations are placed. Therefore, in this study the temperature appears as the main synchronizing factor of gonadal development within these populations. Significant differences in gonadal volume per polyp were found in both species owing mainly to differences in the number of gonads per polyp between populations, with Cape of Palos populations displaying higher values in both studied species, suggesting that the exposition to different local conditions may be reverted in a different gonadal output. But the observed patterns in lipid concentrations levels in gorgonians disable us to conclude that lipid concentration levels explain the observed differences in gonadal output found in this study.     

Temporal variability in zooplankton prey capture rate of the passive suspension feeder<Emphasis Type="Italic"> Leptogorgia sarmentosa</Emphasis> (Cnidaria: Octocorallia), a case study

There is increasing evidence that suspension feeders play a significant role in plankton–benthos coupling. However, to date, active suspension feeders have been the main focus of research, while passive suspension feeders have received less attention. To increase our understanding of energy fluxes in temperate marine ecosystems, we have examined the temporal variability in zooplankton prey capture of the ubiquitous Mediterranean gorgonian Leptogorgia sarmentosa. Prey capture was assessed on the basis of gut content from colonies collected every 2 weeks over a year. The digestion time of zooplankton prey was examined over the temperature range of the species at the study site. The main prey items captured were small (80–200 µm), low-motile zooplankton (i.e. eggs and invertebrate larvae). The digestion time of zooplankton prey increased when temperature decreased (about 150% from 21°C to 13°C; 15 h at 13°C, 9 h at 17°C, and 6 h at 21°C), a pattern which has not previously been documented in anthozoans. Zooplankton capture rate (prey polyp^–1 h^–1) varied among seasons, with the greatest rates observed in spring (0.16±0.02 prey polyp^–1 h^–1). Ingestion rate in terms of biomass (g C polyp^–1 h^–1) showed a similar trend, but the differences among the seasons were attenuated by seasonal differences in prey size. Therefore, ingestion rate did not significantly vary over the annual cycle and averaged 0.019±0.002 g C polyp^–1 h^–1. At the estimated ingestion rates, the population of L. sarmentosa removed between 2.3 and 16.8 mg C m^–2 day^–1 from the adjacent water column. This observation indicates that predation by macroinvertebrates on seston should be considered in energy transfer processes in littoral areas, since even species with a low abundance may have a detectable impact.Communicated by S.A. Poulet, Roscoff     

Distribution of a population of annelid polychaetes in the “trottoir” of the midlittoral zone on the coast of North-East Spain,Western Mediterranean

The discharge and mode of action of the microbasic mastigophores (b-Rhabdoids) of Anemonia sulcata Pennant (Anthozoa) tentacles were examined using light and electron microscopical techniques. The statistical analysis of different morphological parameters of 100 nematocysts before and after their complete discharge revealed four different phases in the process of discharge. The first phase consists of the opening of the three opercular flaps. The second phase is characterized by the extrusion of the shaft. Osmotic pressure generated by the nematocyst propagates the extremely fast shaft discharge. The third phase, the evagination of the long tubule, is propagated by intrinsic mechanical forces stored in the tubule's twists and coils. During the process of evagination the tubule length increases by 95%, its circumference by 17%. Its forward motion is produced by the dilatation of its circumference and undoing of its twists. This motion is supported by the pressure which is generated by the hydration of the capsular matrix. The fourth phase consists in the extrusion of soluble capsular material at the tip of the evaginated tubule. During discharge the net volume of the nematocyst increases by 182%. This expansion is only possible if a high osmotic pressure is generated within the cyst. The extrusion of the shaft is the crucial phase in the discharge of the microbasic mastigophores, because without the puncture produced by the shaft the tubule is not able to overcome even surfaces of low viscosity media. The function of these tentacular nematocysts is the mechanical capture of prey organisms and, most probably, the introduction of unknown toxins into the latter.     

A hierarchical,multi-scale,management-responsive model of Mediterranean vegetation dynamics

Ecosystems of the Mediterranean basin are characterized by a heterogeneous and dynamic landscape mosaic of vegetation formations. This landscape has been shaped over millenia by disturbances associated with agropastoral land use: clearing, grazing and burning, and by regeneration processes of the natural woody vegetation. The ability to predict the effects of management decisions on the structure and composition of the vegetation is essential for present and future land management. To improve this ability we developed a hierarchical multi-scale, management-responsive model of vegetation dynamics.