Stefano Goffredo

Growth and population dynamic model of the reef coral Fungia granulosa Klunzinger, 1879 at Eilat, northern Red Sea.

The lack of population dynamic information for most species of stony corals is due in part to their complicated life histories that may include fission, fusion and partial mortality of colonies, leading to an uncoupling of coral age and size. However, some reef-building corals may produce compact upright or free-living individuals in which the above processes rarely occur, or are clearly detectable. In some of these corals, individual age may be determined from size, and standard growth and population dynamic models may be applied to gain an accurate picture of their life history. We measured long-term growth rates (up to 2.5 years) of individuals of the free-living mushroom coral Fungia granulosa Klunzinger, 1879 at Eilat, northern Red Sea, and determined the size structure of a population on the shallow reef slope. We then applied growth and population models to the data to obtain estimates of coral age, mortality rate, and life expectancy in members of this species. In the field, few F. granulosa polyps suffered partial mortality of >10% of their tissues. Thus, the majority of polyps grew isometrically and determinately, virtually ceasing growth by about 30-40 years of age. Coral ages as revealed by skeletal growth rings were similar to those estimated from a growth curve based on field data. The frequency of individuals in each age class on the reef slope decreased exponentially with coral age, indicating high mortality rates when corals were young. The maximum coral age observed in the field population (31 years) was similar to that estimated by application of a population dynamic model (30 years). Calculated rates of growth, mortality and life expectancy for F. granulosa were within the range of those known for other stony corals. Our results reveal a young, dynamic population of this species on Eilat reefs, with high turnover rates and short lifespans. Such information is important for understanding recovery of coral reefs from disturbances, and for application to the management of commercially exploited coral populations.

Unite research with what citizens do for fun: “recreational monitoring” of marine biodiversity

Institutes often lack funds and manpower to perform large-scale biodiversity monitoring. Citizens can be involved, contributing to the collection of data, thus decreasing costs. Underwater research requires specialist skills and SCUBA certification, and it can be difficult to involve volunteers. The aim of this study was to involve large numbers of recreational divers in marine biodiversity monitoring for increasing the environmental education of the public and collecting data on the status of marine biodiversity. Here we show that thousands of recreational divers can be enrolled in a short time. Using specially formulated questionnaires, nonspecialist volunteers reported the presence of 61 marine taxa encountered during recreational dives, performed as regular sport dives. Validation trials were carried out to assess the accuracy and consistency of volunteer-recorded data, and these were compared to reference data collected by an experienced researcher. In the majority of trials (76%) volunteers performed with an accuracy and consistency of 50-80%, comparable to the performance of conservation volunteer divers on precise transects in other projects. The recruitment of recreational divers involved the main diving and tour operators in Italy, a popular scientific magazine, and mass media. During the four-year study, 3825 divers completed 18757 questionnaires, corresponding to 13539 diving hours. The volunteer-sightings-based index showed that in the monitored area the biodiversity status did not change significantly within the project time scale, but there was a significant negative correlation with latitude, suggesting improved quality in the southernmost areas. This trend could be related to the presence of stressors in the northern areas and has been supported by investigations performed by the Italian Ministry of the Environment. The greatest limitation with using volunteers to collect data was the uneven spatial distribution of samples. The benefits were the considerable amounts of data collected over short time periods and at low costs. The successful development of citizen-based monitoring programs requires open-mindedness in the academic community; advantages of citizen involvement in research are not only adding large data sets to the ecological knowledge base but also aiding in the environmental education of the public.

Growth and population dynamics model of the Mediterranean solitary coral Balanophyllia europaea (Scleractinia, Dendrophylliidae)

Complex life history processes of corals, such as fission, fusion, and partial mortality of colonies, that decouple coral age from size, are rare or clearly detectable in corals that produce distinct colonial or solitary forms. In some of these corals, individual age may be determined from size, and standard age-based growth and population dynamics models may be applied. We determined population size and structure and measured growth rates of Balanophyllia europaea individuals at Calafuria in the eastern Ligurian Sea. We then applied demographic models to these data. Growth rate decreased with increasing coral size. The age–size curve derived from field measurements of growth rates fits that obtained from the computerized tomography analysis of skeletal growth bands. The frequency of individuals in each age class decreased exponentially with age, indicating a population in a steady state. The survival curve showed a turnover time of 3.6 years and a maximum life span of 20 years. This is nearly three times the turnover time and maximum life span recorded for Balanophyllia elegans living off the western coasts of North America, the only congeneric species whose population dynamics has been studied. The Beverton and Holt population model may be useful for comparative analyses of demographic traits and for resource management of solitary or compact, upright growth forms that rarely fragment. This paper completes the description of the main life-strategy characteristics of the Mediterranean endemic coral B. europaea, together with our previous studies on the reproductive biology of this species. This constitutes a major advance in the understanding of the biology and ecology of Mediterranean scleractinian corals, and represents the most complete description of a coral from this geographic area to date that we are aware of.

The Skeletal Organic Matrix from Mediterranean Coral Balanophyllia europaea Influences Calcium Carbonate Precipitation

Scleractinian coral skeletons are made mainly of calcium carbonate in the form of aragonite. The mineral deposition occurs in a biological confined environment, but it is still a theme of discussion to what extent the calcification occurs under biological or environmental control. Hence, the shape, size and organization of skeletal crystals from the cellular level through the colony architecture, were attributed to factors as diverse as mineral supersaturation levels and organic mediation of crystal growth. The skeleton contains an intra-skeletal organic matrix (OM) of which only the water soluble component was chemically and physically characterized. In this work that OM from the skeleton of the Balanophyllia europaea, a solitary scleractinian coral endemic to the Mediterranean Sea, is studied in vitro with the aim of understanding its role in the mineralization of calcium carbonate. Mineralization of calcium carbonate was conducted by overgrowth experiments on coral skeleton and in calcium chloride solutions containing different ratios of water soluble and/or insoluble OM and of magnesium ions. The precipitates were characterized by diffractometric, spectroscopic and microscopic techniques. The results showed that both soluble and insoluble OM components influence calcium carbonate precipitation and that the effect is enhanced by their co-presence. The role of magnesium ions is also affected by the presence of the OM components. Thus, in vitro, OM influences calcium carbonate crystal morphology, aggregation and polymorphism as a function of its composition and of the content of magnesium ions in the precipitation media. This research, although does not resolve the controversy between environmental or biological control on the deposition of calcium carbonate in corals, sheds a light on the role of OM, which appears mediated by the presence of magnesium ions.

Environmental implications of skeletal micro-density and porosity variation in two scleractinian corals

The correlations between skeletal parameters (bulk density, micro-density and porosity), coral age and sea surface temperature were assessed along a latitudinal gradient in the zooxanthellate coral Balanophyllia europaea and in the azooxanthellate coral Leptopsammia pruvoti. In both coral species, the variation of bulk density was more influenced by the variation of porosity than of micro-density. With increasing polyp age, B. europaea formed denser and less porous skeletons while L. pruvoti showed the opposite trend, becoming less dense and more porous. B. europaea skeletons were generally less porous (more dense) than those of L. pruvoti, probably as a consequence of the different habitats colonized by the two species. Increasing temperature had a negative impact on the zooxanthellate species, leading to an increase of porosity. In contrast, micro-density increased with temperature in the azooxanthellate species. It is hypothesized that the increase in porosity with increasing temperatures observed in B. europaea could depend on an attenuation of calcification due to an inhibition of the photosynthetic process at elevated temperatures, while the azooxanthellate species appears more resistant to variations of temperature, highlighting possible differences in the sensitivity/tolerance of these two coral species to temperature changes in face of global climate change.

Growth and Demography of the Solitary Scleractinian Coral Leptopsammia pruvoti along a Sea Surface Temperature Gradient in the Mediterranean Sea

The demographic traits of the solitary azooxanthellate scleractinian Leptopsammia pruvoti were determined in six populations on a sea surface temperature (SST) gradient along the western Italian coasts. This is the first investigation of the growth and demography characteristics of an azooxanthellate scleractinian along a natural SST gradient. Growth rate was homogeneous across all populations, which spanned 7 degrees of latitude. Population age structures differed between populations, but none of the considered demographic parameters correlated with SST, indicating possible effects of local environmental conditions. Compared to another Mediterranean solitary scleractinian, Balanophyllia europaea, zooxanthellate and whose growth, demography and calcification have been studied in the same sites, L. pruvoti seems more tolerant to temperature increase. The higher tolerance of L. pruvoti, relative to B. europaea, may rely on the absence of symbionts, and thus the lack of an inhibition of host physiological processes by the heat-stressed zooxanthellae. However, the comparison between the two species must be taken cautiously, due to the likely temperature differences between the two sampling depths. Increasing research effort on determining the effects of temperature on the poorly studied azooxanthellate scleractinians may shed light on the possible species assemblage shifts that are likely to occur during the current century as a consequence of global climatic change.

Ultrastructural observations of the spermatogenesis of the hermaphroditic solitary coral Balanophyllia europaea (Anthozoa, Scleractinia)

Abstract Spermatogenesis ultrastructure was studied in a simultaneous hermaphrodite population of the solitary coral Balanophyllia europaea. In this species, spermatogenesis takes place in spermatocysts located within gametogenetic mesenteries surrounded by a bilayered boundary. Spermatogonia and spermatocytes are large flagellate cells, densely packed at the outermost edges of the spermatocyst. Spermatids and sperm are loosely distributed near the centre of the spermatocyst. The cytoplasm of spermatogonia and primary spermatocytes often contains short lengths of free axonemes, probably derived from the reabsorption of a primitive flagellum. Maturing spermatids either contain long intracytoplasmic axonemes, that may be stages of the tail synthesis, or have a flagellum. The morphological features of the sperm of this hermaphroditic scleractinian, very similar to those observed in the sperm of gonochoric taxa, support the hypothesis that the hermaphroditism of this population is an adaptive condition.

Genetic assessment of population structure and connectivity in the threatened Mediterranean coral Astroides calycularis (Scleractinia, Dendrophylliidae) at different spatial scales

Understanding dispersal patterns, population structure and connectivity among populations is helpful in the management and conservation of threatened species. Molecular markers are useful tools as indirect estimators of these characteristics. In this study, we assess the population genetic structure of the orange coral Astroides calycularis in the Alboran Sea at local and regional scale, and at three localities outside of this basin. Bayesian clustering methods, traditional F‐statistics and Dest statistics were used to determine the patterns of genetic structure. Likelihood and coalescence approaches were used to infer migration patterns and effective population sizes. The results obtained reveal a high level of connectivity among localities separated by as much as 1 km and moderate levels of genetic differentiation among more distant localities, somewhat corresponding with a stepping‐stone model of gene flow and connectivity. These data suggest that connectivity among populations of this coral is mainly driven by the biology of the species, with low dispersal abilities; in addition, hydrodynamic processes, oceanographic fronts and the distribution of rocky substrate along the coastline may influence larval dispersal.

An adaptive management approach to an octocoral fishery based on the Beverton-Holt model

Coral reef species are frequently the focus of bio-prospecting, and when promising bioactive compounds are identified there is often a need for the development of responsible harvesting based on relatively limited data. The Caribbean gorgonian Pseudopterogorgia elisabethae has been harvested in the Bahamas for over a decade. Data on population age structure and growth rates in conjunction with harvest data provide an opportunity to compare fishery practices and outcomes to those suggested by a Beverton-Holt fishery model. The model suggests a minimum colony size limit of 7–9 years of age (21–28 cm height), which would allow each colony 2–4 years of reproduction prior to harvesting. The Beverton-Holt model assumes that colonies at or above the minimum size limit are completely removed. In the P. elisabethae fishery, colonies are partially clipped and can be repeatedly harvested. Linear growth of surviving colonies was up to 3 times that predicted for colonies that were not harvested and biomass increase was up to 9 times greater than that predicted for undisturbed colonies. The survival of harvested colonies and compensatory growth increases yield, and yields at sites that had previously been harvested were generally greater than predicted by the Beverton-Holt model. The model also assumes recruitment is independent of fishing intensity, but lower numbers of young colonies in the fished populations, compared to unfished populations, suggest possible negative effects of the harvest on reproduction. This suggests the need for longer intervals between harvests. Because it can be developed from data that can be collected at a single time, the Beverton-Holt model provides a rational starting point for regulating new fisheries where long-term characterizations of population dynamics are rarely available. However, an adaptive approach to the fishery requires the incorporation of reproductive data.

Gains and losses of coral skeletal porosity changes with ocean acidification acclimation

Ocean acidification is predicted to impact ecosystems reliant on calcifying organisms, potentially reducing the socioeconomic benefits these habitats provide. Here we investigate the acclimation potential of stony corals living along a pH gradient caused by a Mediterranean CO2 vent that serves as a natural long-term experimental setting. We show that in response to reduced skeletal mineralization at lower pH, corals increase their skeletal macroporosity (features >10 μm) in order to maintain constant linear extension rate, an important criterion for reproductive output. At the nanoscale, the coral skeletons structural features are not altered. However, higher skeletal porosity, and reduced bulk density and stiffness may contribute to reduce population density and increase damage susceptibility under low pH conditions. Based on these observations, the almost universally employed measure of coral biomineralization, the rate of linear extension, might not be a reliable metric for assessing coral health and resilience in a warming and acidifying ocean.