Gilberto M. Amado-Filho

Rhodolith Beds Are Major CaCO3 Bio-Factories in the Tropical South West Atlantic

Rhodoliths are nodules of non-geniculate coralline algae that occur in shallow waters (<150 m depth) subjected to episodic disturbance. Rhodolith beds stand with kelp beds, seagrass meadows, and coralline algal reefs as one of the worlds four largest macrophyte-dominated benthic communities. Geographic distribution of rhodolith beds is discontinuous, with large concentrations off Japan, Australia and the Gulf of California, as well as in the Mediterranean, North Atlantic, eastern Caribbean and Brazil. Although there are major gaps in terms of seabed habitat mapping, the largest rhodolith beds are purported to occur off Brazil, where these communities are recorded across a wide latitudinal range (2°N - 27°S). To quantify their extent, we carried out an inter-reefal seabed habitat survey on the Abrolhos Shelf (16°50′ - 19°45′S) off eastern Brazil, and confirmed the most expansive and contiguous rhodolith bed in the world, covering about 20,900 km2. Distribution, extent, composition and structure of this bed were assessed with side scan sonar, remotely operated vehicles, and SCUBA. The mean rate of CaCO3 production was estimated from in situ growth assays at 1.07 kg m−2 yr−1, with a total production rate of 0.025 Gt yr−1, comparable to those of the worlds largest biogenic CaCO3 deposits. These gigantic rhodolith beds, of areal extent equivalent to the Great Barrier Reef, Australia, are a critical, yet poorly understood component of the tropical South Atlantic Ocean. Based on the relatively high vulnerability of coralline algae to ocean acidification, these beds are likely to experience a profound restructuring in the coming decades.

An extensive reef system at the Amazon River mouth

A novel Amazonian reef biome was discovered, encompassing large rhodolith and sponge beds under low light, low oxygen, and high POC. Large rivers create major gaps in reef distribution along tropical shelves. The Amazon River represents 20% of the global riverine discharge to the ocean, generating up to a 1.3 × 106–km2 plume, and extensive muddy bottoms in the equatorial margin of South America. As a result, a wide area of the tropical North Atlantic is heavily affected in terms of salinity, pH, light penetration, and sedimentation. Such unfavorable conditions were thought to imprint a major gap in Western Atlantic reefs. We present an extensive carbonate system off the Amazon mouth, underneath the river plume. Significant carbonate sedimentation occurred during lowstand sea level, and still occurs in the outer shelf, resulting in complex hard-bottom topography. A permanent near-bottom wedge of ocean water, together with the seasonal nature of the plume’s eastward retroflection, conditions the existence of this extensive (~9500 km2) hard-bottom mosaic. The Amazon reefs transition from accretive to erosional structures and encompass extensive rhodolith beds. Carbonate structures function as a connectivity corridor for wide depth–ranging reef-associated species, being heavily colonized by large sponges and other structure-forming filter feeders that dwell under low light and high levels of particulates. The oxycline between the plume and subplume is associated with chemoautotrophic and anaerobic microbial metabolisms. The system described here provides several insights about the responses of tropical reefs to suboptimal and marginal reef-building conditions, which are accelerating worldwide due to global changes.

Extensive Rhodolith Beds Cover the Summits of Southwestern Atlantic Ocean Seamounts

Abstract PEREIRA-FILHO, G.H.; AMADO-FILHO, G.M.; MOURA, R.L.; BASTOS, A.C.; GUIMARÃES, S.M.P.B.; SALGADO, L.T.; FRANCINI-FILHO, R.B.; BAHIA, R.G.; ABRANTES, D.P.; GUTH, A.Z., and BRASILEIRO, P.S., 2012. Extensive rhodolith beds cover the summits of southwestern Atlantic Ocean seamounts. Calcium carbonate production by marine organisms is an essential process in the global budget of , and coralline reefs are the most important benthic carbonate producers. Crustose coralline algae (CCA) are well recognized as the most important carbonate builders in the tropical Brazilian continental shelf, forming structural reefs and extensive rhodolith beds. However, the distribution of CCA beds, as well as their role in mineralization in mesophotic communities and isolated carbonate banks, is still poorly known. To characterize the bottom features of several seamount summits in the Southwestern Atlantic (SWA), side-scan sonar records, remotely operated vehicle imagery, and benthic samples with mixed-gas scuba diving were acquired during two recent research cruises (March 2009 and February 2011). The tops of several seamounts within this region are relatively shallow (∼60 m), flat, and dominated by rhodolith beds (Vitória, Almirante Saldanha, Davis, and Jaseur seamounts, as well as the Trindade Island shelf). On the basis of abundance, dimensions, vitality, and growth rates of CCA nodules, a mean CaCO3 production was estimated, ranging from 0.4 to 1.8 kg m−2 y−1, with a total production reaching 1.5 × 10−3 Gt y−1. Our results indicate that these SWA seamount summits provide extensive areas of shallow reef area and represent 0.3% of the worlds carbonate banks. The importance of this habitat has been highly neglected, and immediate management needs must be fulfilled in the short term to ensure long-term persistence of the ecosystem services provided by these offshore carbonate realms.

Fish Biodiversity of the Vitória-Trindade Seamount Chain, Southwestern Atlantic: An Updated Database

Despite a strong increase in research on seamounts and oceanic islands ecology and biogeography, many basic aspects of their biodiversity are still unknown. In the southwestern Atlantic, the Vitória-Trindade Seamount Chain (VTC) extends ca. 1,200 km offshore the Brazilian continental shelf, from the Vitória seamount to the oceanic islands of Trindade and Martin Vaz. For a long time, most of the biological information available regarded its islands. Our study presents and analyzes an extensive database on the VTC fish biodiversity, built on data compiled from literature and recent scientific expeditions that assessed both shallow to mesophotic environments. A total of 273 species were recorded, 211 of which occur on seamounts and 173 at the islands. New records for seamounts or islands include 191 reef fish species and 64 depth range extensions. The structure of fish assemblages was similar between islands and seamounts, not differing in species geographic distribution, trophic composition, or spawning strategies. Main differences were related to endemism, higher at the islands, and to the number of endangered species, higher at the seamounts. Since unregulated fishing activities are common in the region, and mining activities are expected to drastically increase in the near future (carbonates on seamount summits and metals on slopes), this unique biodiversity needs urgent attention and management.

Reef fish and benthic assemblages of the Trindade and Martin Vaz Island group, southwestern Atlantic

The Trindade and Martin Vaz island group (TMVIG) is located at about 1,120 km off the Brazilian coast. Despite its importance, highlighted by the presence of several endemic fish species, the TMVIG lacks detailed information on the structure of fish and benthic assemblages. Presented here is the first quantitative assessment of reef fish and benthic assemblages of the TMVIG in a depth gradient ranging from 5 to 45 m. Additional qualitative information on reef assemblages between 45 and 100 m was obtained using advanced gas diving techniques (TRIMIX) and a remotely operated vehicle (ROV). Similarly to other Brazilian oceanic islands, the TMVIG possesses depauperated fish and benthic assemblages, possibly due to its isolation and small size in comparison to the mainland. Depth was the most important factor affecting the structure of fish assemblages, with the density of most fish species declining with depth. Deep reefs (> 45 m) were characterized by the presence of extensive rhodolith beds and rocky reefs sparsely covered with crustose coralline algae, black coral (Cirripathes sp.) and a few massive or plate-like reef corals. Part-time or obligatory planktivorous fishes (e.g. Cephalopholis furcifer and Clepticus brasiliensis) also dominated deep reefs. Similar characteristics were recorded in mesophotic reef ecosystems across the Western Atlantic. Evidence of overfishing (obtained here and in other recent studies), the presence of four endemic and restricted range fish species, as well as the increase in number of new (and still undescribed) endemic taxa, indicates that the adoption of precautionary conservation measures are urgently needed in order to maintain the fragile and unique ecosystems of the TMVIG.

Physiologic and metagenomic attributes of the rhodoliths forming the largest CaCO3 bed in the South Atlantic Ocean.

Rhodoliths are free-living coralline algae (Rhodophyta, Corallinales) that are ecologically important for the functioning of marine environments. They form extensive beds distributed worldwide, providing a habitat and nursery for benthic organisms and space for fisheries, and are an important source of calcium carbonate. The Abrolhos Bank, off eastern Brazil, harbors the world’s largest continuous rhodolith bed (of ∼21 000 km2) and has one of the largest marine CaCO3 deposits (producing 25 megatons of CaCO3 per year). Nevertheless, there is a lack of information about the microbial diversity, photosynthetic potential and ecological interactions within the rhodolith holobiont. Herein, we performed an ecophysiologic and metagenomic analysis of the Abrolhos rhodoliths to understand their microbial composition and functional components. Rhodoliths contained a specific microbiome that displayed a significant enrichment in aerobic ammonia-oxidizing betaproteobacteria and dissimilative sulfate-reducing deltaproteobacteria. We also observed a significant contribution of bacterial guilds (that is, photolithoautotrophs, anaerobic heterotrophs, sulfide oxidizers, anoxygenic phototrophs and methanogens) in the rhodolith metagenome, suggested to have important roles in biomineralization. The increased hits in aromatic compounds, fatty acid and secondary metabolism subsystems hint at an important chemically mediated interaction in which a functional job partition among eukaryal, archaeal and bacterial groups allows the rhodolith holobiont to thrive in the global ocean. High rates of photosynthesis were measured for Abrolhos rhodoliths (52.16 μmol carbon m−2 s−1), allowing the entire Abrolhos rhodolith bed to produce 5.65 × 105 tons C per day. This estimate illustrates the great importance of the Abrolhos rhodolith beds for dissolved carbon production in the South Atlantic Ocean.

Rhodolith-forming species of Lithophyllum (Corallinales; Rhodophyta) from Espirito Santo State, Brazil, including the description of L. depressum sp. nov.

Villas-Boas A.B., Riosmena-Rodriguez R., Amado-Filho G.M., Maneveldt G.W. and de O. Figueiredo M.A. 2009. Rhodolith-forming species of Lithophyllum (Corallinales; Rhodophyta) from Espírito Santo State, Brazil, including the description of L. depressum sp. nov. Phycologia 48: 237–248. DOI: 10.2216/08-35.1. The taxonomy of the rhodolith-forming species of Lithophyllum (Corallinales, Rhodophyta) found from Espirito Santo State, Brazil, were evaluated based on extensive subtidal sampling. A comparative analysis of the Brazilian material with recently described species lead us to conclude that three known species (Lithophyllum corallinae, L. johansenii, L. stictaeforme) and one species new to science (Lithophyllum depressum Villas-Boas, Figueiredo & Riosmena-Rodriguez) are present in the area. All four species can be distinguished primarily by their tetrasporangial conceptacle roof anatomy. This study has shown that Brazilian rhodolith beds are comparatively more diverse in number of Lithophyllum species than other areas of the world.

Baseline Assessment of Mesophotic Reefs of the Vitória-Trindade Seamount Chain Based on Water Quality, Microbial Diversity, Benthic Cover and Fish Biomass Data

Seamounts are considered important sources of biodiversity and minerals. However, their biodiversity and health status are not well understood; therefore, potential conservation problems are unknown. The mesophotic reefs of the Vitória-Trindade Seamount Chain (VTC) were investigated via benthic community and fish surveys, metagenomic and water chemistry analyses, and water microbial abundance estimations. The VTC is a mosaic of reef systems and includes fleshy algae dominated rhodolith beds, crustose coralline algae (CCA) reefs, and turf algae dominated rocky reefs of varying health levels. Macro-carnivores and larger fish presented higher biomass at the CCA reefs (4.4 kg per frame) than in the rhodolith beds and rocky reefs (0.0 to 0.1 kg per frame). A larger number of metagenomic sequences identified as primary producers (e.g., Chlorophyta and Streptophyta) were found at the CCA reefs. However, the rocky reefs contained more diseased corals (>90%) than the CCA reefs (~40%) and rhodolith beds (~10%). Metagenomic analyses indicated a heterotrophic and fast-growing microbiome in rocky reef corals that may possibly lead to unhealthy conditions possibly enhanced by environmental features (e.g. light stress and high loads of labile dissolved organic carbon). VTC mounts represent important hotspots of biodiversity that deserve further conservation actions.

Expansion of an invasive coral species over Abrolhos Bank, Southwestern Atlantic

Invasive coral species of the genus Tubastraea have been increasingly recorded in Southwestern Atlantic waters since the 1980s. Their invasion and infestation are mainly related to port and oil exploration activities. For the first time the presence of Tubastraea tagusensis colonies is reported in Espírito Santo State, colonizing a port shore area, and incrusting oil/gas platform structures situated in the southern Abrolhos Bank, which is part of the most important coral reef system of the South Atlantic Ocean. Tubastraea colonies exhibit fast growth and high recruitment rates, and colonized 40% of the analyzed structures in just four years. The projection of port and oil/gas industry growth for the Espírito Santo State (more than 300%) highlights an alert to the dispersal of this alien species to natural areas.

Traffic of secondary metabolites to cell surface in the red alga Laurencia dendroidea depends on a two-step transport by the cytoskeleton.

In Laurencia dendroidea, halogenated secondary metabolites are primarily located in the vacuole named the corps en cerise (CC). For chemical defence at the surface level, these metabolites are intracellularly mobilised through vesicle transport from the CC to the cell periphery for posterior exocytosis of these chemicals. The cell structures involved in this specific vesicle traffic as well as the cellular structures related to the positioning and anchoring of the CC within the cell are not well known. Here, we aimed to investigate the role of cytoskeletal elements in both processes. Cellular and molecular assays were conducted to i) determine the ultrastructural apparatus involved in the vesicle traffic, ii) localise cytoskeletal filaments, iii) evaluate the role of different cytoskeletal filaments in the vesicle transport, iv) identify the cytoskeletal filaments responsible for the positioning and anchoring of the CC, and v) identify the transcripts related to cytoskeletal activity and vesicle transport. Our results show that microfilaments are found within the connections linking the CC to the cell periphery, playing an essential role in the vesicle traffic at these connections, which means a first step of the secondary metabolites transport to the cell surface. After that, the microtubules work in the positioning of the vesicles along the cell periphery towards specific regions where exocytosis takes place, which corresponds to the second step of the secondary metabolites transport to the cell surface. In addition, microtubules are involved in anchoring and positioning the CC to the cell periphery. Transcriptomic analysis revealed the expression of genes coding for actin filaments, microtubules, motor proteins and cytoskeletal accessory proteins. Genes related to vesicle traffic, exocytosis and membrane recycling were also identified. Our findings show, for the first time, that actin microfilaments and microtubules play an underlying cellular role in the chemical defence of red algae.