Ecological Shifts in Mediterranean Coralligenous Assemblages Related to Gorgonian Forest Loss

Mediterranean gorgonian forests are threatened by several human activities and are affected by climatic anomalies that have led to mass mortality events in recent decades. The ecological role of these habitats and the possible consequence of their loss are poorly understood. Effects of gorgonians on the recruitment of epibenthic organisms were investigated by manipulating presence of gorgonians on experimental panels at 24 m depth, for Eunicella cavolinii, and at 40 m depth, for Paramuricea clavata, at two sites: Tavolara Island (Tyrrhenian Sea) and Portofino Promontory (Ligurian Sea). After 4 months, the most abundant taxa on the panels were encrusting green algae, erect red algae and crustose coralline algae at 24 m depth and encrusting brown algae and erect red algae at 40 m depth. Assemblages on the panels were significantly affected by the presence of the gorgonians, although effects varied across sites and between gorgonian species. Species diversity and evenness were lower on panels with gorgonian branches. Growth of erect algae and recruitment of serpulid polychaetes were also affected by the presence of the gorgonians, primarily at Tavolara. Crustose coralline algae and erect sponges were more abundant on E. cavolinii panels at 24 m depth, while encrusting bryozoans were more abundant on P. clavata panels at 40 m depth. Effects of gorgonians on recruited assemblages could be due to microscale modification of hydrodynamics and sediment deposition rate, or by a shading effect reducing light intensity. Gorgonians may also intercept settling propagules, compete for food with the filter-feeders and/or for space by producing allelochemicals. Presence of gorgonians mainly limits the growth of erect algae and enhances the abundance of encrusting algae and sessile invertebrates. Therefore, the gorgonian disappearances may cause a shift from assemblages characterised by crustose coralline algae to filamentous algae assemblages, decreasing complexity and resilience of coralligenous bioconstructions.     

Morphological features of the gorgonian Paramuricea macrospina on the continental shelf and shelf edge (Menorca Channel,Western Mediterranean Sea)

Morphological variability in gorgonians is frequent and commonly associated with habitat variability, often resulting in segregated morphotypes. Paramuricea macrospina is an endemic Mediterranean gorgonian species found on rocky bottoms between 40 and 160?m depth and has recently been reported as one of the most abundant species in continental shelf and shelf edge environments. Three different chromatic forms of P. macrospina were observed in the Menorca Channel: a yellow form and a light purple form occurring on maërl beds of the continental shelf, and a dark purple form occurring on rocky substrates of the shelf edge. The objective of the present work is to verify if these P. macrospina forms may represent distinct taxonomic units by analysing differences in colony morphology and sclerite size and shape of the three chromatic forms. No significant differences were found in colony shape, suggesting that environmental variability between the continental shelf and the shelf edge is not influential enough to significantly alter colony morphology. Significant differences in sclerite size and shape were found amongst all forms, suggesting that sclerites may be influenced by environmental conditions. However, the co-occurrence of the yellow and light purple forms side by side on the continental shelf may indicate a certain degree of genetic differentiation.     

Estimating surface area of sponges and gorgonians as indicators of habitat availability on Caribbean coral reefs

A rapid method to estimate the three-dimensional (3D) surface area (SA) of marine gorgonians and sponges from field measurements of colony height, diameter, and morphology was developed as an indicator of habitat availability for fish and invertebrates. Colony characteristics for sponges and gorgonians were compiled from field measurements, expert judgment, and taxonomic literature, and employed to generate 3D images using computer-aided design software. The images were used to test various statistical models and geometric surrogates that best estimated SA using only height and diameter measurements. A morphological classification system was devised using shapes and relative proportions of sponges and gorgonians which are commonly found in shallow waters (<25 m depth) of the Central Western Atlantic Ocean. Regression models (linear, quadratic, or cubic) were found to be more robust than geometric surrogates, exhibiting greater accuracy at range extremes. Statistical models explained over 90% of the variation in SA and forecast errors of less than 20%. The best models for estimating SA are presented for eight sponge and nine gorgonian morphologies. Application of these methods with existing estimators for stony corals SA can be used as an indicator of structural habitat availability, which is an important ecosystem service of coral reefs.     

Cycle of gonadal development in Eunicella singularis (Cnidaria: Octocorallia): trends in sexual reproduction in gorgonians

Abstract. Eunicella singularis is a gorgonian species whose members are abundant in hard-bottom sublittoral communities of the Mediterranean Sea. The reproductive biology in this species has been examined to better understand the ability in this species to recover from recent mass mortality events. Eunicella singularis is a stable, gonochoric, iteroparous species that reproduces annually and exhibits a seasonal pattern of gametogenesis characterized by a single annual maturation of the gametes. The sex ratio did not significantly differ from 1:1. Oogenesis lasted 13–17 months, beginning between February and June, and ending with the release of 0.7 planula larvae per polyp between late May and July of the following year. The diameter of mature oocytes ranged 450–860 μm. Spermatogenesis was much shorter than oogenesis and occurred over 5–6 months. Gonadal production of both sexes increased in spring and culminated with the spawning of male colonies in late May–June. Fertilization of oocytes and development of the planula larvae occurred within the polyps of female colonies. Planula release was observed in June and July. The patterns emerging from this and previous studies on sexual reproduction of Mediterranean gorgonians suggest that investment in gonad development appears to be related to resource availability.     

Distribution and biodiversity of Singapore gorgonians (sub-class Octocorallia) — a preliminary survey

Preliminary surveys of the gorgonian fauna of Singapore show the existence of 6 families, comprising at least 11 genera and 21 species. The distribution of this fauna is closely related to the availability of a suitable substrata for settlement and growth. In addition, the greatest number of gorgonians, in terms of abundance and species richness, is found on the lower slopes and bottoms of the reefs in Singapore. The intermediate disturbance hypothesis is applied to the data obtained and shown to be useful in making inferences on the state of succession of gorgonian communities.     

Evaluation of the utility of COI and ITS markers as tools for population genetic studies of temperate gorgonians

The aim of this study was to analyse the polymorphism of two molecular markers, mitochondrial COI and nuclear ITS 2, to determine the genetic structure of Mediterranean gorgonian populations. Four gorgonians, Eunicella cavolinii, Eunicella singularis, Paramuricea clavata and Corallium rubrum were analysed for COI, while only E. cavolinii was analysed for ITS. These species have been recently affected by mass mortality events probably linked to climate change. In the context of these strong disturbances, reliable data on the genetic structure of affected populations could be critical in order to propose management and conservation strategies for these species. A hierarchical sampling plan encompassing different spatial scales (from tenths to hundreds of kilometres) was designed to test these markers. Both markers failed to make evident any genetic structure of these populations. The evolutionary features and utility of these markers are critically discussed and future research directions are proposed.     

Grenadier abundance examined at varying spatial scales in deep waters off Newfoundland, Canada, with special focus on the influence of corals

There is a growing body of research examining the effects of corals on fish communities, species abundances, and biodiversity. Yet, few studies have quantitatively examined what factors are influencing the distribution of individual fish species. In general, many researchers believe they know what influences the distribution of grenadiers on large spatial scales, but numerous studies have shown the distributions of organisms are often determined by various factors that change in relative importance when viewed at differing scales. Our study used video collected from three deep canyons off Newfoundland, Canada (North west Atlantic) to examine how the factors apparently influencing the distribution of four grenadiers (Macrouridae: Coryphaenoides rupestris, Coryphaenoides carapinus, Nezumia bairdii, and Macrourus berglax) change when assessed at varying spatial scales. We paid special attention to the influence of deep-water corals found in the study area (large gorgonians/antipatharians, small gorgonians, sea pens, soft corals, and cup corals). The factors that influenced grenadier presence and/or abundance (and the magnitude of this effect) varied as different sampling resolutions were examined. We found C. rupestris abundance was positively related to cup coral abundance in transects longer than 10 m, likely as a result of similar habitat preferences between both taxa. When significant relationships between depth and C. rupestris presence and/or abundance were found, they were always negative. Depth was a significant predictor of C. carapinus abundance in transects longer than 10 m. Very few predictors of M. berglax abundance or presence could be found. Depth and the number of small gorgonians were consistent predictors of N. bairdii abundance.     

Distribution of cichlid fishes in the littoral of Lake Bacalar, Yucatan Peninsula

The distribution of cichlids in Lake Bacalar was assessed from January to September 1996, in eight shallow-water (< 2 m) stations, using a 20 m long seine net. Temperature, salinity/conductivity and dissolved oxygen were sensor-obtained; depth, submerged vegetation and bottom type were determined by visual inspection. Affinities between stations, based on fish abundance and environmental parameters, were analyzed by multivariate methods. Littoral of Bacalar town was the site with highest richness, diversity and abundance, probably promoted by the additional urban waste of organic matter. PCA defined two station-groups, one associated with muddy substrates, the other one with rocky/sandy bottoms. Cichlids are more abundant over muddy bottoms with abundant submerged vegetation, probably because of the abundance of prey and shelter.     

Variability at different spatial scales of a coralligenous assemblage at Giannutri Island (Tuscan Archipelago,northwest Mediterranean)

This study was carried out on the rocky cliffs of Giannutri Island (Tyrrhenian Sea, Italy) to test the hypothesis that coralligenous assemblages are consistent within the bathymetric range considered (25–35 m depth) over three different spatial scales (1000 m, 100 m and 10 m). A multi-factorial sampling design was used to assess patterns of vertical distribution in the studied area. Data on the percent cover of algae and invertebrates were collected at three depths (25, 30 and 35 m) using a photographic method, and percentage cover was obtained using a visual method. Analysis of the results using ANOVA indicated that the distribution and abundance of algae and some invertebrates of the sublittoral assemblages are clearly heterogeneous. Algae, sponges and bryozoans showed significant variability in distribution and abundance at different depths, but this variability was not consistent amongst transects. Ascidians did not show any variability, while the abundance of anthozoans differed significantly amongst transects. We concluded that heterogeneity in the distribution and abundance of the taxa analysed is related to the smallest spatial scale investigated (10's of m). Some of the possible causes of the observed variability are discussed.     

Diversity and relative abundance of corals, octocorals and sponges at Jaragua National Park, Dominican Republic

The Jaragua National Park is located in a remote area to the SW coast of the Dominican Republic. Fishing and mining are the major human activities. The main reef formations of the Park include: (a) long bank reefs (spur and groove) growing as bands over the platform and running in a SW-NW direction at 12-25 m depth, (b) well developed, deep, fringing reefs at the platform edge (drop-off) areas which could extend from 10 to 45 m depth, and (c) small patch reefs and poorly developed coral-octocoral-sponge-algal communities in shallow platforms near shore, rocky bottoms, and over the submerged walls of the uplifted reef. Nine reef localities were surveyed between Cabo Beata and Bahia Honda using Scuba diving to inventory the diversity and relative abundance of scleractinian corals, octocorals and sponges. Fringing reefs were surveyed starting at the bottom (30 m) and swimming in a zig-zag pattern (50 m on each side) to shallower areas. Bank reefs were surveyed by swimming in zig-zag across the spur-groove formation along 500 m. Sponges were the most diverse group with 83 species in 50 genera followed by the scleractinian corals with 56 species in 26 genera and the octocorals with 47 species in 15 genera. New records included eight coral species, 29 octocoral species and 59 sponges. The diversity, species composition and abundance of particular groups varied across the different localities. Northern reefs within the park and the Los Frailes Island offshore had the highest live cover, relative abundance and diversity for the three groups. In general, the Jaragua National Park had the highest diversity of corals, octocorals and sponges reported for the Dominican Republic and rank amongst the highest reported for the northern Caribbean. It is recommended that the area be protected and that fishing activities be regulated or eliminated altogether.     

Seaweed succession on artificial reefs on different bottom substrata

Artificial rest reefs were set on sandy and rocky bottoms at 5–10 m depth along the coast of southern Japan. Mature thalli ofSargassum, Gelidium and other seaweds were transported from other coastal areas, packed in mesh bags and attached to the reefs to start the beds. After one year, the seaweed flora on the reef on a sandy bottom consisted of more than 20 species, includingSargassum spp. andGelidium amansii, which are important animal food species. Coralline algae were the dominants on the rocky bottom reefs. The lower biomass on reefs on the rocky bottom was due to grazing by urchins. The same number of species was present in the first and second years on reefs on sandy bottoms, but there were moreSargassum thalli the second year.Maximum algal biomass of the artificial reef in May of the second year was 9998 g wet wt m^–2 in sandy areas, 441 g wet wt m^–2 in boulder areas and 228 g wet wt m^–2 in rocky areas. Reefs on rocky bottoms continued to be covered by coralline algae and several species ofCodium andDictyota.     

Gorgonian distribution patterns in coral reef environments of the Gulf of Mexico: evidence of sporadic ecological connectivity?

Gorgonian species richness and abundance were documented in a survey of 14 reefs across the Gulf of México. The results show two main patterns: (1) a westward decreasing gradient in gorgonian diversity and abundance at both supraspecific and specific levels, from the Yucatán straits to the southwestern Gulf of México reefs; and (2) a northward decreasing gradient in gorgonian diversity and abundance, only apparent at the reef group scale, in the southwestern Gulf of México. Even though the Campeche reef system consists of widely dispersed reefs, their gorgonian faunas are relatively rich, indicating that no isolation exists in this extensive area. In contrast, gorgonians in the continentally influenced reefs of the southwestern Gulf of México are rare, even though scleractinians are diverse and abundant. The author suggests that the observed patterns in gorgonian species richness and abundance cannot be explained solely by factors such as geographical setting, reef morphology, habitat suitability, storm effects, or anthropogenic impacts. Instead the observed patterns may primarily reflect limitations in ecological connectivity at the mesoscale, variably affected by ocean and shore current dynamics, as well as by continental and local effects.     

Reassessing the spatial relationship between sponges and macroalgae in sublittoral rocky bottoms: a descriptive approach

Because sublittoral sponges of temperate areas are usually more abundant at sites with low algal abundance, there is the widespread notion that macroalgae out-compete and displace sponges to habitats less suitable for algal proliferation. In this study, based on 292 sampling quadrats, we collected a total of 87 demosponge species and examined the level at which sponge distribution pairs with a variety of alga-dominated and animal-dominated habitats occurring in three zones located across a marked in-bay/out-bay environmental gradient. We found significant differences in sponge biomass, richness and diversity between the 18 habitats considered in the three zones, with abundance, richness and diversity being significantly higher in caves, vertical surfaces and overhangs out of the bay than in the remaining habitats. The cluster analysis and the unconstrained ordination consistently reflected the in-bay/out-bay environmental gradient. These analyses also revealed that the taxonomic distribution of sponge abundance is independent of the algal occurrence in the habitat, being more related to between-zone differences than to between-habitat differences. This trend was corroborated when the role of depth, algal abundance and substratum inclination in explaining total sponge abundance and diversity was examined by canonical correspondence analysis, regression analysis and mean comparisons. These analyses pointed to substratum inclination, rather than to algal abundance, as the factor explaining most variation in distribution of sponge abundance. These results, when discussed in the context provided by the outcome of other studies concerned with the spatial distribution of the sessile benthos in rocky temperate communities, strongly suggest a need to re-examine the idea that spatial distribution of sublittoral sponges largely results from competition with macroalgae.     

Multivariate statistical analysis of distribution of deep-water gorgonian corals in relation to seabed topography on the norwegian margin

Investigating the relationship between deep-water coral distribution and seabed topography is important for understanding the terrain habitat selection of these species and for the development of predictive habitat models. In this study, the distribution of the deep-water gorgonians, Paragorgia arborea and Primnoa resedaeformis, in relation to terrain variables at multiple scales of 30 m, 90 m and 170 m were investigated at Røst Reef, Traena Reef and Sotbakken Reef on the Norwegian margin, with Ecological Niche Factor Analysis applied. To date, there have been few published studies investigating this aspect of gorgonian distribution. A similar correlation between the distribution of P. arborea and P. resedaeformis and each particular terrain variable was found at each study site, but the strength of the correlation between each variable and distribution differed by reef. The terrain variables of bathymetric position index (BPI) and curvature at analysis scales of 90 m or 170 m were most strongly linked to the distribution of both species at the three geographically distinct study sites. Both gorgonian species tended to inhabit local topographic highs across all three sites, particularly at Sotbakken Reef and Traena Reef, with both species observed almost exclusively on such topographic highs. The tendency for observed P. arborea to inhabit ridge crests at Røst Reef was much greater than was indicated for P. resedaeformis. This investigation identifies the terrain variables which most closely correlate with distribution of these two gorgonian species, and analyzes their terrain habitat selection; further development of predictive habitat models may be considered essential for effective management of these species.     

Patterns of benthic mega-invertebrate habitat associations in the Pacific Northwest continental shelf waters: a reassessment

As human impacts and demands for ocean space increase (fisheries, aquaculture, marine reserves, renewable energy), identification of marine habitats hosting sensitive biological assemblages has become a priority. Epifaunal invertebrates, especially the structure-forming species, are an increasing conservation concern as many traditional (bottom-contact fishing) and novel (marine renewable energy) ocean uses have the potential to displace or otherwise impact these slow-growing organisms. The differences in mega-invertebrate species assemblages between high-relief rocks and low-relief sediments are well documented and likely hold for most marine environments. In anticipation of potential development of marine renewable energy faculties off Oregon and Washington (USA), a survey of the benthic invertebrate assemblages and habitats was conducted on three rocky reefs on the continental shelf of the Pacific Northwest, using video footage collected by remotely operated vehicle, to more finely characterize these assemblage–habitat associations. Benthic assemblages appeared to first group by depth (50–80 vs. 100–120 m), then by relief (consolidated rocks vs. unconsolidated rocks and soft sediments). Consolidated rocks were characterized at each site by a combination of various sponges, gorgonians, sea anemones and echinoderms; unconsolidated rocks were characterized at Grays Bank by sea anemones and burrowing brittle stars, and at Bandon-Arago by sponges and echinoderms; soft sediments were characterized at Grays Bank and Siltcoos Reef by sea whips and burrowing brittle stars, as well as pink shrimps and sea stars at Siltcoos Reef, and at Bandon-Arago by sponges, gorgonians and echinoderms. The results of this study will help classify and map the seafloor in a way that represents benthic habitats reflective of biological species assemblage distributions, rather than solely geological features, and support conservation and management planning.     

Reproductive cycle and trophic ecology in deep versus shallow populations of the Mediterranean gorgonian Eunicella singularis (Cap de Creus,northwestern Mediterranean Sea)

Coral Reefs - The annual gonad development of a shallow (20&nbsp;m depth) population of the Mediterranean gorgonian Eunicella singularis was found to be closely synchronized with that of a deep...     

Reef habitats and associated sessile-benthic and fish assemblages across a euphotic–mesophotic depth gradient in Isla Desecheo, Puerto Rico

Quantitative surveys of sessile benthos and fish populations associated with reef habitats across a 15–50 m depth gradient were performed by direct diver observations using rebreathers at Isla Desecheo, Puerto Rico. Statistically significant differences between depths were found for total live coral, total coral species, total benthic algae, total sponges and abiotic cover. Live coral cover was higher at the mid-shelf (20 m) and shelf-edge (25 m) stations, whereas benthic algae and sponges were the dominant sessile-benthic assemblage at mesophotic stations below 25 m. Marked shifts in the community structure of corals and benthic algae were observed across the depth gradient. A total of 119 diurnal, non-cryptic fish species were observed across the depth gradient, including 80 species distributed among 7,841 individuals counted within belt-transects. Fish species richness was positively correlated with live coral cover. However, the relationship between total fish abundance and live coral was weak. Abundance of several numerically dominant fish species varied independently from live coral cover and appeared to be more influenced by depth and/or habitat type. Statistically significant differences in the rank order of abundance of fish species at euphotic vs mesophotic stations were detected. A small assemblage of reef fishes that included the cherubfish, Centropyge argi, sunshine chromis, Chromis insolata, greenblotch parrotfish, Sparisoma atomarium, yellowcheek wrasse, Halichoeres cyanocephalus, sargassum triggerfish, Xanthichthys ringens, and the longsnout butterflyfish, Chaetodon aculeatus was most abundant or only present from stations deeper than 30 m, and thus appear to be indicator species of mesophotic habitats.     

Characteristics of the mesophotic megabenthic assemblages of the vercelli seamount (north tyrrhenian sea)

The biodiversity of the megabenthic assemblages of the mesophotic zone of a Tyrrhenian seamount (Vercelli Seamount) is described using Remotely Operated Vehicle (ROV) video imaging from 100 m depth to the top of the mount around 61 m depth. This pinnacle hosts a rich coralligenous community characterized by three different assemblages: (i) the top shows a dense covering of the kelp Laminaria rodriguezii; (ii) the southern side biocoenosis is mainly dominated by the octocorals Paramuricea clavata and Eunicella cavolinii; while (iii) the northern side of the seamount assemblage is colonized by active filter-feeding organisms such as sponges (sometimes covering 100% of the surface) with numerous colonies of the ascidian Diazona violacea, and the polychaete Sabella pavonina. This study highlights, also for a Mediterranean seamount, the potential role of an isolated rocky peak penetrating the euphotic zone, to work as an aggregating structure, hosting abundant benthic communities dominated by suspension feeders, whose distribution may vary in accordance to the geomorphology of the area and the different local hydrodynamic conditions.     

Modeling the habitat suitability for deep-water gorgonian corals based on terrain variables

The coral species Paragorgia arborea and Primnoa resedaeformis are abundant and widely distributed gorgonians in North Atlantic waters. Both species add significant habitat complexity to the benthic environment, and support a host of invertebrate species. Mapping their distribution is an essential step in conservation and resource management, but challenging as a result of their remoteness. In this study, three predictive models — Ecological Niche Factor Analysis, Genetic Algorithm for Rule-set Production and Maximum Entropy modeling (MaxEnt) were applied to predict the distribution of species' suitable habitat across a region of Røst Reef (Norwegian margin) based on multiscale terrain variables.All three models were successful in predicting the habitat suitability for both gorgonian species across the study area, and the MaxEnt predictions were shown to outperform other predictions. All three models predicted the most suitable habitats for both species to mainly occur along the ridges and on the upper section of the large slide, suggesting both species preferentially colonize topographic highs. Jackknife tests for MaxEnt predictions highlighted the seabed aspect in relation to P. arborea distribution, and the seabed relative position (curvature) in relation to the distribution of both species. Given the vulnerability of deep-water corals to anthropogenic impacts, further comparative study over a wider study area would be particularly beneficial for the management of the species.     

Newly dominant benthic invertebrates reshape competitive networks on contemporary Caribbean reefs

Competition is a fundamental process structuring ecological communities. On coral reefs, space is a highly contested resource and the outcomes of spatial competition can dictate community composition. In the Caribbean, reefs are increasingly dominated by non-scleractinian species like sponges, gorgonians, and zoanthids, yet there is a paucity of data on interactions between these increasingly common organisms and historically dominant corals. Here, we investigated interactions among these groups of sessile benthic invertebrates to better understand the role of spatial competition in shaping benthic communities on Caribbean reefs. We coupled surveys of competitive interactions on the reef with a common garden competition experiment to determine the frequency and outcome of interference competition among eight focal species. We found that competitive interactions were pervasive on Florida reefs, with 60% of sessile benthic invertebrates interacting with at least one other invertebrate. Increasingly common non-scleractinian species were some of the most abundant taxa and consistently outcompeted the contemporarily common scleractinian species Porites porites and Siderastrea siderea. The encrusting gorgonian, Erythropodium caribaeorum, was the most aggressive species, reducing the live area of its competitors on average 42% ± 7.04 (SE) over the course of 5 months. Surprisingly, the most aggressive species declined in size when competing, while some less aggressive species were able to increase or maintain area, suggesting a trade-off between aggressiveness and growth. Our findings suggest that competition among sessile invertebrates is likely to remain an important process in structuring coral reefs, but that the optimal strategies for maintaining space on the benthos may change. Importantly, many non-scleractinian species that now dominate reefs appear to be superior competitors, potentially increasing the stress on corals on contemporary reefs.