Black coral forests enhance taxonomic and functional distinctiveness of mesophotic fishes in an oceanic island: implications for biodiversity conservation.

The degradation of shallow ecosystems has called for efforts to understand the biodiversity and functioning of Mesophotic Ecosystems. However, most empirical studies have been restricted to tropical regions and have majorly focused on taxonomic entities (i.e., species), neglecting important dimensions of biodiversity that influence community assembly and ecosystem functioning. Here, using a subtropical oceanic island in the eastern Atlantic Ocean (Lanzarote, Canary Islands), we investigated variation in (a) alpha and (b) beta functional (i.e., trait) diversity across a depth gradient (0-70 m), as a function of the presence of black coral forests (BCFs, order Antipatharian) in the mesophotic strata, a vulnerable but often overlooked 'ecosystem engineer' in regional biodiversity. Despite occupying a similar volume of the functional space (i.e., functional richness) than shallow (< 30 m) reefs, mesophotic fish assemblages inhabiting BCFs differed in their functional structure when accounting for species abundances, with lower evenness and divergence. Similarly, although mesophotic BCFs shared, on average, 90% of the functional entities with shallow reefs, the identity of common and dominant taxonomic and functional entities shifted. Our results suggest BCFs promoted the specialization of reef fishes, likely linked to convergence towards optimal traits to maximize the use of resources and space. Regional biodiversity planning should thus focus on developing specific management and conservation strategies for preserving the unique biodiversity and functionality of mesophotic BCFs.

Stochastic diffusion characterises early colony formation in Mediterranean coral Corallium rubrum.

The colony formation in Mediterranean coral Corallium rubrum is initiated by a larva that metamorphoses into the first polyp of the emerging colony approximately two weeks after settlement. The primary polyp then sets up a slow process that eventually, at least after a few years, gives rise to a tree-like rigid colony structure on which other polyps flourish. For a mature colony, this axial skeleton provides support for new polyps. However, the first emergence of the characteristic axial skeleton can take two years or more from the larva stage. The early colony morphology, instead, is shaped exclusively by the polyps' abundant deposition of sclerites, a magnesian calcite biomineral that has a different granularity from the distinctive red-coloured skeleton. With the appearance of the first polyp, a growing sclerite heap in a mesoglea layer provides a base for the emerging colony. In this paper, to elucidate the mechanical processes of early skeleton development in C. rubrum colonies, we present a computational model whereby the mesoglea layer provides a diffusion medium for the sclerites that the polyps deposit. We show that our stochastic model with three parameters captures the dynamic variability observed in measurements on living colonies. Our simulation results provide evidence for a diffusion process whereby the interplay between polyp budding and sclerite deposition are the main determinants of structure in early colony formation. Our model demonstrates that the frequency of budding events in an early colony can be described as a function of the available mesoglea surface whereas the number of polyps on the colony plays a secondary role in determining this frequency. We show that these model predictions are confirmed by direct observations on the colonies in our sample. Moreover, our results indicate that diffusion is a prevalent mechanism of colony development also at later stages of a colony's life span.

Deep-Dwelling Populations of Mediterranean Corallium rubrum and Eunicella cavolini: Distribution, Demography, and Co-Occurrence.

Corallium rubrum and Eunicella cavolini are two octocorals, reported as co-occurring species in the deep rocky habitats of the Mediterranean Sea with a high hydrodynamic and moderate eutrophication. Their spatial distribution and demography in the deep sea are mainly affected by temperature and direct and indirect anthropogenic activities; however, knowledge of the factors that potentially influence their co-existence is scarce. This paper provides novel data on the distribution and demography of these two species, at depths between 50 and 290 m in the Western Mediterranean Sea, providing insights on their co-occurrence. Both species exhibited the highest population density at deeper sites (>150 m), showing an inverse size-density relation. Density values ranged from 0.03 colonies m-2 to 32 and 80 col. m-2 for yellow gorgonian and red coral, respectively. The two species co-occurred in 13% of the total frames examined, mostly dwelling between 120 and 160 m depth. Distance-based linear modeling (DistLM) emphasized that when co-occurring the variability of the two species' densities were significantly driven by the density-rather than the morphology (i.e., height)-of the other species. We stress the need for further studies to elucidate the possible mutual effects of suspension feeders and to test the role of different environmental factors potentially influencing inter-specific relationships.

Colonization of plastic debris by the long-lived precious red coral Corallium rubrum: New insights on the "plastic benefits" paradox.

Seafloor macrolitter is ubiquitous in world's oceans; still, huge knowledge gaps exist on its interactions with benthic biota. We report here the colonization of plastic substrates by the Mediterranean red coral Corallium rubrum (L. 1758), occurring both in controlled conditions and in the wild at ca. 85 m depth in the Western Mediterranean Sea. Juveniles settled on seafloor macro-litter, with either arborescent or encrusting morphology, ranging from 0.6 to 3.5 mm in basal diameter and 0.2-7.1 years of age, also including a fraction (20%) of potentially sexually mature individuals. In controlled conditions, larvae settled and survived on plastic substrates for >60 days. Our insights show that marine plastic debris can provide favourable substrate for C. rubrum settlement either in controlled conditions or in the wild, suggesting their possible use in restoration activities. However, we pinpoint here that this potential benefit could result in adverse effects on population dynamics.

The rise of octocoral forests on Caribbean reefs.

Coral reefs throughout the tropics have experienced large declines in the abundance of scleractinian corals over the last few decades, and some reefs are becoming functionally dominated by animal taxa other than scleractinians. This phenomenon is striking on many shallow reefs in the tropical western Atlantic, where arborescent octocorals now are numerically and functionally dominant. Octocorals are one of several taxa that have been overlooked for decades in analyses of coral reef community dynamics, and our understanding of why octocorals are favoured (whereas scleractinians are not) on some modern reefs, and how they will affect the function of future reef communities, is not commensurate with the task of scientifically responding to the coral reef crisis. We summarize the biological and ecological features predisposing octocorals for success under contemporary conditions, and focus on those features that could have generated resistance and resilience of octocoral populations to environmental change on modern reefs. There is a rich set of opportunities for rapid advancement in understanding the factors driving the success of octocorals on modern reefs, but we underscore three lines of inquiry: (1) the functional implications of strongly mixotrophic, polytrophic, and plastic nutrition, (2) the capacity to recruit at high densities and maintain rapid initial rates of vertical growth, and (3) the emergent properties associated with dense animal forests at high colony densities.

Density-dependence mediates coral assemblage structure.

Density dependence (DD) controls community recovery following widespread mortality, yet this principle rarely has been applied to coral assemblages. The reefs of Mo'orea, French Polynesia, provide the opportunity to study DD of coral population growth, because coral assemblages in this location responded to declines in abundance with high recruitment and an increase in cover during which recruitment of pocilloporid corals was inversely associated with density. This study tests for DD in this system, first, by describing the context within which it operates: coral cover changed from 46% in 2005, to <1% in 2010 following an outbreak of a corallivorous sea star and a cyclone, and then increased to 74% by 2017, in large part through inverse density-associated pocilloporid recruitment. Second, a test for DD of recruitment was conducted by decreasing Pocillopora spp. cover from 33% to 19%: one year later, the density of Pocillopora spp. recruits was 1.65-fold higher in the low vs. high cover treatment. Finally, the effects of DD were investigated by comparing simulated and empirical distributions of pocilloporid colonies: as predicted by DD, small colonies were randomly distributed, while large colonies were uniformly distributed. Together these results demonstrate DD of population regulation for Pocillopora spp. corals, thus revealing the potential importance of this ecological principle in determining the resilience of coral assemblages.

Limited Genetic Connectivity between Gorgonian Morphotypes along a Depth Gradient.

Gorgonian species show a high morphological variability in relation to the environment in which they live. In coastal areas, parameters such as temperature, light, currents, and food availability vary significantly with depth, potentially affecting morphology of the colonies and the structure of the populations, as well as their connectivity patterns. In tropical seas, the existence of connectivity between shallow and deep populations supported the hypothesis that the deep coral reefs could potentially act as (reproductive) refugia fostering re-colonization of shallow areas after mortality events. Moreover, this hypothesis is not so clear accepted in temperate seas. Eunicella singularis is one of the most common gorgonian species in Northwestern Mediterranean Sea, playing an important role as ecosystem engineer by providing biomass and complexity to the coralligenous habitats. It has a wide bathymetric distribution ranging from about 10 m to 100 m. Two depth-related morphotypes have been identified, differing in colony morphology, sclerite size and shape, and occurrence of symbiotic algae, but not in mitochondrial DNA haplotypes. In the present study the genetic structure of E. singularis populations along a horizontal and bathymetric gradient was assessed using microsatellites and ITS1 sequences. Restricted gene flow was found at 30-40 m depth between the two Eunicella morphotypes. Conversely, no genetic structuring has been found among shallow water populations within a spatial scale of ten kilometers. The break in gene flow between shallow and deep populations contributes to explain the morphological variability observed at different depths. Moreover, the limited vertical connectivity hinted that the refugia hypothesis does not apply to E. singularis. Re-colonization of shallow water populations, occasionally affected by mass mortality events, should then be mainly fueled by larvae from other shallow water populations.

Habitat constraints and self-thinning shape Mediterranean red coral deep population structure: implications for conservation practice.

The Mediterranean red coral, Corallium rubrum, is one of the most precious corals worldwide. Below 50 m depth, C. rubrum populations are generally characterised by large and sparse colonies, whereas shallow populations (above 50 m depth) show high densities of small colonies. We show here instead that populations dwelling between 80 and 170 m depth exhibited a continuous range of population density (from 2 to 75 colonies per 0.25 m(2)), with less than 1% of variance explained by water depth. An inverse relationship between maximum population density and mean colony height was found, suggesting that self-thinning processes may shape population structure. Moreover, demographically young populations composed of small and dense colonies dominated along rocky vertical walls, whereas mature populations characterised by large and sparsely distributed colonies were found only in horizontal beds not covered by sediment. We hypothesise that, in the long term, shallow protected populations should resemble to present deep populations, with sparsely distributed large colonies. Since the density of red coral colonies can decay as a result of self-thinning mechanisms, we advise that future protection strategies should be based also on a measure of red coral spatial coverage instead of population density.

Modelling ecological complexity for marine species conservation: the effect of variable connectivity on species spatial distribution and age-structure.

Connectivity is currently emphasized as a key factor in conservation for its role in enhancing biodiversity of an area and giving benefit to the adjacent areas. For most marine species, connectivity is synonomous of larval dispersal. We applied a spatially explicit meta-population model to test the hypothesis that larval dispersal can affect local demographical features, consequently misleading conservation practice in the marine environment. Simulations were carried out in the Gulf of Lions where coastal circulation displays highly variable temporal and spatial submeso-scale structures. Two different benthic invertebrate species were considered: a soft bottom short lived species and a hard bottom long lived one. In the first case, simulations showed that highest densities at equilibrium do not inform on self-persistent populations location. In the second case, simulations showed that connectivity effects may result in out-of-equilibria demographical structure. We emphasized the caveats in the parameterization of demographical models when local demography is controlled by connectivity.

The role of light in mediating the effects of ocean acidification on coral calcification.

We tested the effect of light and PCO2 on the calcification and survival of Pocillopora damicornis recruits settled from larvae released in southern Taiwan. In March 2011, recruits were incubated at 31, 41, 70, 122 and 226 µmol photons m(-2) s(-1) under ambient (493 µatm) and high PCO2 (878 µatm). After 5 days, calcification was measured gravimetrically and survivorship estimated as the number of living recruits. Calcification was affected by the interaction of PCO2 with light, and at 493 µatm PCO2 the response to light intensity resembled a positive parabola. At 878 µatm PCO2, the effect of light on calcification differed from that observed at 493 µatm PCO2, with the result that there were large differences in calcification between 493 µatm and 878 µatm PCO2 at intermediate light intensities (ca. 70 µmol photons m(-2) s(-1)), but similar rates of calcification at the highest and lowest light intensities. Survivorship was affected by light and PCO2, and was highest at 122 µmol photons m(-2) s(-1) in both PCO2 treatments, but was unrelated to calcification. In June 2012 the experiment was repeated, and again the results suggested that exposure to high PCO2 decreased calcification of P. damicornis recruits at intermediate light intensities, but not at lower or higher intensities. Together, our findings demonstrate that the effect of PCO2 on coral recruits can be light dependent, with inhibitory effects of high PCO2 on calcification at intermediate light intensities that disappear at both higher and lower light intensities.

Applying cathodically polarised substrata to the restoration of a high value coral.

Larval settlement of the high value red coral, Corallium rubrum, was studied on three different CaCO(3) substrata, viz. lithogenic (marble), electro-accreted calcium carbonate in the presence and in the absence of cathodic polarisation. The last two substrata consisted of stainless steel plates galvanically coupled with Zn anodes. The electrochemical characterization of the settlement device was studied in order to investigate correlations between cathodic parameters (polarisation potential, current density, calcareous deposit composition) and larval settlement. The results obtained in the natural habitat (at 35 m depth) showed that settlement was five times lower on the electro-accreted aragonite in the presence of low cathodic current densities (i≤1 µA cm(-2)) compared to both marble tiles and electro-accreted aragonite in the absence of polarisation. These last two substrata showed similar settlement values. The implications of these findings on restoration strategies for C. rubrum are discussed.

Population dynamics and conservation biology of the over-exploited Mediterranean red coral.

The main goal of ecologists is nowadays to foster habitat and species conservation. Life-history tables and Leslie-Lewis transition matrices of population growth can be powerful tools suitable for the study of age-structured over harvested and/or endangered species dynamics. Red coral (Corallium rubrum L 1758) is a modular anthozoan endemic to the Mediterranean Sea. This slow growing, long lived species has been harvested since ancient times. In the last decades harvesting pressure increased and the overall Mediterranean yield reduced by 23. Moreover, mass mortality (putatively-linked to global warming) recently affected some coastal populations of this species. Red coral populations are discrete genetic units, gonochoric, composed by several overlapping generations and provided of a discrete (annual) reproduction. A population of this precious octocoral was studied in detail and its static life table was compiled. In order to simulate the trends overtime of the population under different environmental conditions and fishing pressures, a discrete, non-linear model, based on Leslie-Lewis transition matrix, was applied to the demographic data. In this model a bell-shaped curve, based on experimental data, representing the dependence of recruitment on adult colonies density was included. On these bases the stability of the population under different density, reproduction and mortality figures was analysed and simulations of the population trends overtime were set out. Some simulations were also carried out applying to the studied population the mortality values measured during the anomalous mass mortality event which really affected some red coral populations in 1999. The population under study showed high stability and a strong resilience capability, surviving to a 61% reduction of density, to a 27.7% reduction of reproduction rate and to an unselective harvesting affecting 95% of the reproductive colonies.

Ecology through time, an overview.

This brief review is an attempt to condense the major events in the history of Ecology into short 10 sections. In little more than two generations, a new science has emerged and developed into a fundamental part of our lives, spurred on by increasing interest in "natural systems" and concerns over the environmental changes we are witnessing. Ecology, rather neglected until the early decades of last century, flourished and established itself as mature science during the mid-1900s, producing many theories, models, hypothesis and trends of thought. Ecology deals with interacting natural systems and eclectically applies tools drawn from several different sciences (Biology, Mathematics, Statistics, Chemistry, Geology, Physics and so on). The beauty of the subject and its multidisciplinary approach makes Ecology extremely intriguing for researchers. One of the main goals of Ecology is to forecast population, community and ecosystem trends over time. Ecological systems are complex: they are composed of such a large number of different, interacting components that their overall behavior can only be understood in terms of emerging properties. Therefore, some interactions and effects are difficult to predict. However, some insights into population and community trends can be inferred from exhaustive data sets and sufficiently long-term, time-series data and dynamic models. Unfortunately, due to the limited funding of environmental data collection, only a few exhaustive, long-term samplings have been carried out, and systematic record keeping for the purposes of ecological research has only recently become widespread. However, it is still possible to garner some insights from historical reports (proxies), which clearly show how the population structures of most of today's communities have been affected by human activities. This short review is based on a talk given at Arcidosso (Grosseto, Italy) during the September, 2005 workshop: "Scientific research and society during the last fifty years", organized by the Interdisciplinary Center for the Study of Complex Systems of the University of Pisa (CISSC), in cooperation with the Domus Galilaeana. The talk was addressed to scientists in various fields with the threefold aim of presenting Ecology as a true science, stimulating curiosity and laying the bases for further insights. We hope that this review, as brief as it is, may at least partially attain such goals.